Systematic revision of Australian and Indo-Pacific Lucinidae (Mollusca: Bivalvia): Pillucina,
Wallucina and descriptions of two new genera and four new species.
.Emily A. Glover & John D. Taylor 263
Revision of the Australian seahorses of the genus Hippocampus (Syngnathiformes:
Syngnathidae) with descriptions of nine new species. Rudie H. Kuiter 293
Snailfishes (Pisces: Liparidae) of Australia, including descriptions of thirty new species.
.David L. Stein, Nat alt a V. Chernova & Anatoly P. Andriashev 341
Larvae and juveniles of the deepsea “whalefishes” Barbourisia and Rondeletia
(Stephanoberyciformes: Barbourisiidae, Rondeletiidae), with comments on family relationships
.John R. Paxton, G. David Johnson & Thomas Trnski 407
Contents Volume 53. 427
Contents Supplements 26 and 27. 429
VOLUME 53 NUMBER 3 12 December 2001
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Records of the Australian Museum (2001) Vol. 53: 263-292. ISSN 0067-1975
Systematic Revision of Australian and Indo-Pacific
Lucinidae (Mollusca: Bivalvia): Pillucina, Wallucina and
Descriptions of Two New Genera and Four New Species
Emily A. Glover and John D. Taylor
Department of Zoology, Natural History Museum, London SW7 5BD, United Kingdom
emily. glover @ dial.pipex. com
j. tay 1 or @ nhm. ac. uk
Abstract. This taxonomic revision concerns the Australian and Indo-Pacific species of small lucinid
bivalves possessing a deeply inset internal ligament. Nine species of Pillucina are recognised of which
four occur around Australia including the two new species, P. pacifica and P. australis. Two other new
species are described; P. denticula from South Africa and P. mauritiana from Mauritius. Pillucina
vietnamica is common along the Queensland coast and P. symbolica the only species previously recorded
from Australia is considered conspecific with the wide ranging species P. pisidium. A new genus,
Chavania, includes two species; C. striata is widely distributed in the Indo-West Pacific including
eastern and western Australia, while C. erythraea is restricted to the Arabian Peninsula. Two species of
Wallucina live around Australia, W. assimilis is endemic occurring at southerly locations from New
South Wales to North West Cape, while the tropical species, W. fijiensis, is found in island habitats of
Queensland but is widely distributed in the Indo-West Pacific. Another new genus, Funafutia, is proposed
for the species, F. levukana, recorded from Australia for the first time. Details of anatomy are provided
for Pillucina vietnamica, W. assimilis and C. striata. Symbiotic bacteria are confirmed for the first time
in the lateral zone of gill filaments of Pillucina vietnamica and Wallucina assimilis.
Glover, Emily A., & John D. Taylor, 2001. Systematic revision of Australian and Indo-Pacific Lucinidae
(Mollusca: Bivalvia): Pillucina, Wallucina and descriptions of two new genera and four new species. Records of
the Australian Museum 53(3): 263-292.
The discovery of the symbiotic association between lucinid
bivalves and sulphide oxidising chemoautotrophic bacteria
has attracted much recent biological interest (Reid, 1990;
Distel, 1998). The symbiosis has now been confirmed in at
least 30 species of Lucinidae representing 18 different
genera from several distinct clades (Taylor & Glover, 2000)
and from this may it be extrapolated as present in most, if
not all, living taxa. This biological interest highlights the
neglect of lucinid systematics at all levels, with most taxa
being rather poorly defined (e.g., Chavan, 1969) and known
only from shells. The last major taxonomic review (Bretsky,
1976) was largely restricted to north American taxa and no
phylogenetic analyses, either morphological or molecular,
provide an evolutionary framework for the group.
Additionally, recent studies of tropical lucinids show that the
family is much more diverse than has been previously realised
(Glover & Taylor, 1997; Taylor & Glover, 1997a,b) and many
small species remain undocumented.
During a search for lucinids on the Queensland coast we
discovered populations of an abundant Pillucina species,
not previously reported from Australia, living on the outer
fringes of mangroves at Port Douglas. On attempting to
identify this species ( Pillucina vietnamica ) it soon became
clear that the taxonomy of Pillucina and related genera was
264 Records of the Australian Museum (2001) Vol. 53
in a state of confusion and, moreover, other species, either
undescribed or unrecorded from Australia, were present in
museum collections. None of these species were considered
in the recent synopses of the bivalve fauna of Australia
(Lamprell & Whitehead, 1992; Lamprell & Healy, 1998).
To identify and assess the status of the Australian species,
we embarked on a review of the Australian and Indo-West
Pacific species of Pillucina. Because of similarities of shell
form we also took into account species of Wallucina and
other genera having an internal ligament with the form
described by Allen (1960) for Loripes lucinalis. Genera with
internal ligaments were briefly reviewed by Oliver (1986)
when he introduced a new generic name for West African
species. Apart from Keletistes from West Africa (Oliver,
1986) and Loripes from northeast Atlantic, Mediterranean
and western Indian Ocean, the other genera are, at the
present day, confined to the Indo-West Pacific province.
These genera, Pillucina, Wallucina, Sydlorina are not
well defined (e.g., Iredale, 1930) and much confusion
concerns the assignment of species (Chavan, 1937, 1938,
1969). Most species are rather small and usually the figures
accompanying the original descriptions lack detail, as is
the case for Pillucina spaldingi Pilsbry, 1921, the type
species of Pillucina. Additionally, several species, such as
Lucina funafutica Finlay, 1927, cannot be satisfactorily
accommodated within any described genus. A preliminary
survey of museum collections also showed that some species
have considerably broader distributions within the Indo-
West Pacific province than previously realised.
Most species considered in this study are known only
from shells and little information is available concerning
their biology. However, chemosynthetic bacteria have been
previously described from the gills of Pillucina pisidium
from the Sea of Japan (Rodionov & Yushin, 1991). Also,
Barnes & Hickman (1999) record but do not illustrate
ctenidial bacteria and provide a detailed habitat description
for Wallucina assimilis from Western Australia.
The objectives of this study are to describe and review
the species of Pillucina, Wallucina and other genera
possessing internal ligaments living around Australia and
to review the other species of the Indo-West Pacific
province. Virtually all relevant primary type material
available has been examined and lucinid collections studied
in the museums of Sydney, Melbourne, Adelaide, Perth,
London, Cardiff and Paris. Additionally, we describe the
anatomy and life habit of some live-collected animals, and
confirm the presence of chemosymbiotic bacteria in the
ctenidia of two species.
Although we have live-collected material of a few
species, most samples in museum collections usually
comprise dead shells from beach collections or otherwise
picked from sediment samples. It is clear that the frequency
of these small white bivalves in collections is a function of
the effort devoted by particular museums (notably the
Australian Museum) to intensive sediment sorting. Thus,
the distribution maps included in this paper probably do
not reflect the complete ranges of these species but to some
extent are artifacts of the intensity of sampling effort.
Methods
Material for anatomical studies was fixed either in 4%
seawater formalin, or in a 2.5% solution of glutaraldehyde
in phosphate buffer. For serial thin sections, animals were
decalcified in EDTA, wax embedded, sectioned at 8 pm
and stained with Masson’s trichrome. For scanning electron
microscopy (SEM), animals were dissected, then critical
point dried, mounted on stubs and sputter coated with gold.
Most shells were imaged without coating using an
environmental SEM.
Most available type specimens have been examined as
indicated in the synonymies by the word “seen”. The frequently
used terms: shell height in mm (H), shell length in mm (L) and
tumidity of a single valve (T), have been abbreviated, as
have museums and institutions cited in the text:
AMS Australian Museum, Sydney
ANSP Academy of Natural Sciences, Philadelphia
BMNH The Natural History Museum, London
BPBM Bernice P. Bishop Museum, Honolulu
DK Henk Dekker, private collection
MCG Museo Civico, Genoa
MCZ Museum of Comparative Zoology, Harvard
MNB Museum fur Naturkunde, Berlin
MNHN Museum national d’Histoire naturelle, Paris
NMSA Natal Museum, South Africa
NMV National Museum of Victoria, Melbourne
NMW National Museum of Wales, Cardiff
NSMT National Science Museum, Tokyo
SAM South Australian Museum, Adelaide
UMUT University Museum, University of Tokyo
UMZC University Museum of Zoology, Cambridge, UK
WAM Western Australian Museum, Perth
ZISP Zoological Institute, St Petersburg, Russia
Systematic descriptions
In this review, we consider lucinid genera possessing an
internal ligament of the form described for Loripes lucinalis
by Allen (1960). This type of ligament (Fig. 1) has a broad,
wedge-like internal portion which broadens to the posterior
and is formed within a deep embayment of the mantle
epithelium. The groove in the hinge (resilifer) to
accommodate the ligament is deeply inset and is aligned
obliquely and laterally into the left and right valves.
Although no comprehensive phylogenetic analysis of the
Lucinidae has yet been attempted it is possible that this
internal ligament may prove be an apomorphy of a clade of
genera allied to Loripes. In a phenetic analysis, Bretsky
(1970, 1976) recognised a distinct “ Loripes group” based
on this form of ligament.
The genera possessing internal ligaments were briefly
reviewed by Oliver (1986) when he introduced the new
generic name, Keletistes, for West African species. In this
paper, we redefine some of the genera, based on reinvestig¬
ation of the type species and introduce two new genera for
species which could not be accommodated into any existing
taxa. The Indo-West Pacific genera Pillucina, Wallucina,
and two new genera Chavania and Funafutia are described
and illustrated in the systematic section below and their
characters summarised in Table 1. The name Sydlorina
Iredale is synonymised with Pillucina (p. 266). The main
characters we have used to discriminate the genera derive
from the external sculpture, hinge teeth and shape. Although
some characters may appear rather minor, after examination
of many specimens from around the Indo-Pacific we feel
that the groupings of species probably represent mono-
phyletic clades. The three other genera with internal
ligaments Loripes, Keletistes and the Miocene Microloripes
are briefly reviewed below and their characters also listed
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 265
Figure 1. Hinge of left valve of Pillucina pisidium showing the deeply inset internal ligament. Scale bar =1.0 mm.
in Table 1. Chavan (1938) regarded Microloripes as a “stem”
group from which Loripes, Pillucina and Wallucina were
derived, however, a phylogenetic analysis of all these genera
is needed to clarify their relationships.
Other genera possessing internal ligaments
Microloripes Cossmann, 1910 (type species: Lucina dentata
Defrance, 1823 a fossil from the L. Miocene of France
(Burdigalian)). Although Chavan (1938) considered
Microloripes close to and a likely sister taxon to Wallucina
and Pillucina, he later classified it (Chavan, 1969) as a
subgenus of Parvilucina despite the possession of an internal
ligament. The type species is similar to Chavania (described
herein p. 282) in general morphology but possesses a
sculpture of rather broad, flat, commarginal lamellae and a
strong anterior lateral tooth. Microloripes has been applied
to a Recent species, Lucina contrarius Dunker, 1846, from
Table 1. Summary of shell characters of lucinid genera considered in this review and other genera with internal ligaments.
shell sculpture
commarginal
radial ribs lamellae
marginal
crenulae
ligament
anterior
adductor
cardinal
hinge teeth
anterior
lateral
posterior
lateral
sulci
Chavania
faint riblets to
anterior and
posterior
fine, low, present,
anastomosing fine,
more prominent margin
to posterior thickened
internal,
very short
short,
detached for
1/3—1/5
of length
1 RV
large,
2 LV
RV absent
or weak,
LV absent
RV present,
low;
LV present,
low
shallow,
posterior
Funafutia
fine striae
between
lamellae
prominent,
thick
present,
fine
internal,
short
short, barely
detached
1 RV,
2 LV
RV present
prominent,
LV present
prominent
RV present,
LV present
absent
Pillucina
fine to coarse
ribs often
stronger to
posterior
and anterior
numerous,
fine to
prominent,
fluted in some
species
present,
fine to
coarse
internal,
short
short, detached
only slightly
or for less
than 1/2 length
1 RV,
2 LV
RV present
or absent,
LV present
or absent
RV present,
LV present
absent
Wallucina
absent or
fine striae
numerous, low
very fine
internal,
deeply inset
detached,
for 1/2 length
1 RV,
2 LV
RV v small RV present
to indistinct LV present
LV absent
very
shallow
posterior
other genera with internal ligaments
Keletistes
weak riblets
towards
anterior and
posterior
numerous,
fine
anastomosing
present,
fine
variable
internal,
short
short, detached
for 1/2
length
1 RV
v low,
2 LV
v low
RV low to
obsolete
LV low to
obsolete
low,
indistinct
absent
Loripes
weak radial
striations
fine growth
lines
absent
internal, long
deeply inset
long, detached
for 2/3
of length
1 RV,
2 LV
RV small
LV absent
to v small
RV present
LV present
present
posterior
Microloripes
faint radial
striations
broad, slightly
raised,
anastomosing
fine
internal,
very short
very short,
barely
detached
1 RV,
2 LV
RV present
LV present
RV low present
LV low posterior
and anterior
266 Records of the Australian Museum (2001) Vol. 53
West Africa (Chavan, 1937; Dell, 1964), however, von Cosel
(pers. comm.) considers this species as part of a broader
concept of Keletistes.
Keletistes Oliver, 1986 (type species: Loripes (Keletistes)
rhizoecus Oliver, 1986 from West Africa). This genus was
proposed for an unusual species, K. rhizoecus , which is
subquadrate in shape, with commarginal lamellae, slight
radial costae, a denticulate inner margin, and hinge teeth
including anterior laterals that become obsolete with age.
Another West African species, K. aberrans (Dautzenberg,
1910), was an included species and von Cosel (pers. comm.)
also considers the other West African species Lucina
contrarius and L. legouxi (Nickles, 1952) as part of the same
group. The relationships of the West African species to the
Indo-Pacific Chavania, Wallucina and Pillucina need to be
examined in more detail.
Loripes Poli, 1791 (type species Tellina lactea Poli, 1791
(non Linnaeus, 1758) = Amphidesma lucinalis Lamarck,
1818 from the eastern Atlantic and Mediterranean). The type
species has a thin subcircular shell, slightly inflated, smooth,
with a sculpture of fine growth lines only, a hinge with strong
anterior lateral teeth, an obliquely inset internal ligament, a
long anterior adductor muscle scar detached from the pallial
line for about 2/3 of its length and a smooth inner shell
margin. Loripes lucinalis is common in the Eastern Atlantic
and Mediterranean and the morphologically rather different,
Loripes clausus (Philippi, 1850), occurs in the Red Sea,
along the east African coast and Madagascar.
Family Lucinidae Fleming, 1828
Genus Pillucina Pilsbry, 1921
Pillucina Pilsbry, 1921: 382. Type species: original designation
Pillucina spaldingi Pilsbry, 1921 (= P. hawaiiensis Smith,
1885).
Sydlorina Iredale, 1930: 390. Type species: original designation
Sydlorina symbolica Iredale, 1930
Diagnosis. Shells small (H to 14), inflated, sculpture of fine
to broad radial ribs that often bifurcate, crossed by fine,
low, commarginal lamellae. Sculpture usually more
prominent on anterior and posterior parts of shell. Hinge
with two cardinal teeth in left valve and a single cardinal
tooth in right valve. Posterior lateral tooth present in both
valves, anterior lateral tooth present or absent. Ligament
internal, located on elongate resilifer widening to the
posterior. Anterior adductor muscle scar short and detached
from pallial line for about 1/4—1/2 of length. Inner shell
margin finely to coarsely crenulate.
Remarks. The type species, which possesses relatively fine
radial ribs, would seem to be rather different in morphology
from some of the other species we regard as congeneric, in
particular the coarsely ribbed Pillucina vietnamica. However,
a complete range in the strength of radial ribbing is seen
amongst Pillucina species and we have found no objective
criteria for separating them. Pillucina differs from Wallucina,
Chavania and Funajutia in the presence of radial ribs which
cross the commarginal lamellae and which are particularly
prominent to the anterior and posterior parts of the shell.
Distribution. Indo-West Pacific, Red Sea to Hawaii.
Pillucina hawaiiensis (Smith, 1885)
Fig. 2h,i
Lucina (Codakia) hawaiiensis Smith, 1885: 183-184, pi. 13, figs.
8, 8a. Three syntypes: BMNH 1887.2.9.2780; unrecognisable
from serious decay from Byne’s disease. Type locality:
Honolulu Harbour, Hawaii. Seen
Pillucina hawaiiensis (Smith).-Dali, Bartsch & Rehder, 1938: 134,
pi. 36, figs. 9-12.
Loripes (Pillucina) spaldingi Pilsbry, 1921: 381, fig. 17. Lectotype:
ANSP 127929; H 8.6, L 7.6. Type locality: Kaneohe Bay, Oahu,
Hawaii.
Pillucina spaldingi (Pilsbry).-Dali, Bartsch & Rehder, 1938: 133—
134, pi. 36, figs. 7 & 8.
Description. Small, H to 10, robust, inflated shell, higher
than long (H/L 1.1), low commarginal growth increments,
with fine radial ribs more prominent in the anterior and
posterior parts of shell. Lunule, short, broad, slightly
impressed. Hinge plate sinuously curved with cardinals
located on a downward projection. Right valve with strong,
single cardinal tooth under umbo, short anterior lateral tooth
lying close to the cardinal tooth and elongate posterior
lateral. Left valve with two cardinal teeth, anterior is larger
and elongate; anterior and posterior lateral teeth present.
Ligament internal, located on elongate resilifer widening
to the posterior. Anterior adductor scar short, detached
ventrally from pallial line for about 1/5 of length, posterior
scar ovate. Inner shell surface with faint radial grooves. Pallial
line continuous. Shell outside of pallial line slightly thickened.
Shell margin finely and evenly crenulate. Colour white.
Distribution. Endemic to Hawaiian Is (12-20 m), see Kay
(1979).
Remarks. Although P. hawaiiensis and P spaldingi have
been usually considered as separate species (Dali, Bartsch
& Rehder, 1938; Kay, 1979), they are similar in most
characters except shape and we regard them as variants of
a single species. Specimens usually named P. hawaiiensis
are slightly longer than high compared with the higher P.
spaldingi. Such shape variation is common within Indo-
Pacific Pillucina species.
Material examined. Hawaiian Islands: off Waikiki, Oahu
(BPBM); entrance to Honolulu Harbour, Oahu (BPBM);
Midway, (BPBM).
Pillucina pacifica n.sp.
Figs. 2a-g, 3
Type material. Holotype: AMS C355685; H 5.5, L 5.5.
Paratypes: AMS C380464, 3 paired valves. Paratypes:
BMNH 2000204,3 single valves. Type locality: Michaelmas
Cay, Great Barrier Reef, Queensland, Australia, 16°36'S
145°59'E, collected by Iredale and Whitley, May 1926.
Description. Shell small (H to 8), robust, inflated, slightly
higher than long (H/L 1.1). Juveniles are proportionately
more anteriorly extended (Fig. 2f) and outline of adults is
variable from much higher than long to more circular in
shape. Sculpture of fine radial ribs (>50) that are slightly
more prominent in anterior of the shell and are slightly
broader posteriorly. Radial ribs are crossed by thread-like,
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 267
Figure 2. Pillucina pacifica n.sp. and P. hawaiiensis (Smith), a-c, Pillucina pacifica holotype AMS C355685,
Michaelmas Cay, Great Barrier Reef, Queensland; a, exterior of left valve; b, interior of left valve; c, interior of
right valve, d-f, paratypes AMS C380464; d, exterior of left valve; e, interior of right valve; f, exterior of right
valve of juvenile specimen, g, detail of hinge of left valve of holotype. h,i, Pillucina hawaiiensis (Smith), off
Waikiki, Oahu, Hawaii (BPBM 22068); h, exterior of left valve; i, interior of right valve. Scale bars =1.0 mm.
commarginal lamellae giving a reticulate ornament.
Ribbing is often worn in the middle part of the shell. No
sulci present. Lunule is short, broad and slightly
impressed. Hinge plate sinuous, with cardinal teeth
located on downward projection. Right valve with single
prominent cardinal tooth with a thin, elongate, posterior
lateral tooth and an indistinct anterior lateral peg. Left
valve with two cardinal teeth of which the anterior is
larger. Posterior lateral tooth a thin groove, anterior
lateral indistinct. Ligament internal, short. Anterior
adductor scar short and barely detached from the pallial
line, posterior scar ovate. Pallial line continuous and shell
outside pallial line thickened. Shell margin finely and
evenly crenulate. Colour white.
268 Records of the Australian Museum (2001) Vol. 53
Habitat. Intertidal to shallow subtidal (20-30 m).
Distribution. Northern Australia on the Great Barrier Reef
and offshore islands to Micronesia (Fig. 3).
Remarks. Pillucina pacifica is similar to P. hawaiiensis
but the latter has a longer lunule, a more robust hinge area,
and in the right valve a less prominent anterior lateral tooth
which is located at a greater distance from the cardinal tooth.
The radial ribbing of P. pacifica is more prominent
compared to P. hawaiiensis and the marginal crenulations
are coarser.
Material examined. Australia: Queensland, Murray I., Torres
Strait (AMS); Thursday I., Torres Strait (AMS); Albany Passage,
Cape York (AMS); Flinders I., Princess Charlotte Bay (AMS);
Lizard I. (AMS); Eagle I. (AMS); Low Isles, 22 m (AMS);
Michaelmas Cay 16°36'S 145°59'E (AMS); Green I., Cairns
(AMS); Ellison Reef, near Kurrimine (AMS); Juno Bay, Fantome
I., Palm Group (AMS); North East Bay, Great Palm I., 6 m (AMS);
Nara Inlet, Hook I., Whitsunday Group (AMS); Heron I.,
Capricorn Group (AMS); Lady Elliott I., Bunker Group (AMS);
Lady Musgrave I., Bunker Group 23°54'S 152°25'E lagoon (AMS);
NE of Cape Moreton Light 26°55'S 153°33'E (AMS). New
Guinea: Lolorual., Port Moresby, 13-18 m (AMS); SW of Losuia,
Kiriwina I., Trobriand Is, mudflats (AMS). Kiribati: Tarawa
Lagoon (AMS). New Caledonia: Baie des Isoles, Ouemo,
Magenta mudflats (AMS); Noumea (MNHN); Touho, 11m, 20°5'S
165°8'E (MNHN).
Pillucina pisidium (Dunker, 1860)
Figs. 4a-f, 8
Lucina pisidium Dunker, 1860: 227; figured by Dunker, 1861:
28, pi. 3, fig. 9. Two syntypes: badly eroded, MNB. Type
locality: Dejima, Nagasaki City, Japan. Seen.
Lucina parvula Gould, 1861: 36. Lectotype: MCZ169284. Type
locality: Port Lloyd, Bonin Is (see Johnson, 1964: 122, pi. 28,
fig. 5) (non Lucina parvula Muenster, 1835).
Codakiapisidium (Dunker).-Hedley, 1914: 699, figs. 25-28.
Sydlorina symbolica Iredale, 1930: 390. Syntypes: AMS C032175;
Hedley’s figured specimen plus many other syntypes. Type
locality: Sydney, NSW, Australia (introduced as new name for
Hedley’s material and figure). Seen.
Loripes (Pillucina) pisidium (Dunker).-Chavan, 1937: 226.
Pillucina (Pillucina)pisidium (Dunker).-Kuroda, Habe & Oyama,
1971: 393, pi. 118, fig. 14; Habe, 1977: 126, pi. 24, figs. 5 & 6.
Description. Shell small, H to 7, moderately inflated, longer
than high (H/L 0.98). Sculpture of fine, close set,
commarginal lamellae crossed by fine radial ribs which are
more distinct at posterior and anterior parts of the shell.
Shell surface slightly nodulose at junctions of radial and
commarginal sculpture. Posteriorly, there is usually a very
shallow sulcus with commarginal sculpture only. Lunule
shallowly impressed. Hinge plate sinuous with cardinal teeth
located on a downward projecting buttress. Right valve with
prominent single cardinal tooth, no anterior lateral, and a
low, short, posterior lateral. Left valve with two cardinal
teeth, no anterior lateral and a low short, posterior lateral.
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 269
Figure 4. Pillucina pisidium (Dunker); a,b, exterior of left valve and interior of right valve of P. pisidium, Hizen,
Hirado, Kyushu, Japan (BMNH 1009.6.9.60); c,d, exterior of left valve and interior of right valve, Sydney, NSW
(BMNH 1963534); e,f, exterior and interior of right valve, a less mature individual than c, Port Jackson, Sydney,
NSW (AMS C013982). Scale bars = 1.0 mm.
Ligament internal, sitting on broad, obliquely extended
triangular resilifer. Anterior adductor muscle scar short, narrow,
detached for about one third of length. Posterior adductor
muscle scar ovate. Pallial line patchily discontinuous with
ventral margin slightly thickened and finely crenulate. Inner
shell surface dull, chalky, pustulate within pallial line. Pallial
blood vessel scar visible. Colour white.
Remarks. Hedley (1914) identified the Pillucina species
from Sydney as Codakia pisidium although he had not seen
the type and the original illustration is poor (Dunker, 1861).
However, Iredale (1930) later considered that the Australian
shells were only slightly similar to L. pisidium from Japan
and introduced a new genus and species, Sydlorina
symbolica. Despite Iredale’s opinion, the Japanese and
270 Records of the Australian Museum (2001) Vol. 53
Australian shells are very similar and we consider them as
conspecific. We have examined the type material of L. pisidium
Dunker in Berlin, other material from Japan and the Indo-
Pacific and conclude that P. pisidium is widely distributed
species which includes the eastern Australian shells.
Pillucina pisidium can be distinguished from P. australis
(Fig. 6) from southern and southwestern Australia, by the
absence of an anterior lateral tooth in the right valve, by
the more numerous and finer radial ribs and its larger size.
It is also similar to P. neglecta, also from Japan, which is
smaller, more inflated and with a strong anterior lateral tooth
in the right valve.
Two types of bacteria have been reported in bacteriocytes
from the gills of P. pisidium from the Sea of Japan
(Rodionov & Yushin, 1991). One type resembles the
sulphide-oxidising, chemosymbiotic bacteria found in other
lucinids, while the other type with cytoplasmic inclusions
is dissimilar to other bivalve symbionts.
Habitat. Intertidal and shallow water (to 90 m) in mud and
fine sand.
Distribution. Indo-West Pacific, Japan to East Africa and
Madagascar (Fig. 8).
Material examined. Kenya: Ras Anzuani, near Shimoni
(BMNH); Kilifi (NMSA). Tanzania: Kunduchi Beach, 6 km N
of Dar es Salaam (AMS); Ras Fumba, Zanzibar (DC). Mada¬
gascar: Tulear (several lots MNHN). Mozambique: Conducia
Bay; Bazzaruto Is (ZMSA). Reunion: Souris Chaude (ZMSA).
Mauritius: Le Goulet, Tombeau Bay (ZMSA). Seychelles:
Aldabra Atoll, Main Channel (BMNH); Cascade, Mahe (BMNH).
Maldives: Gan lagoon, Addu Atoll (BMNH). Ceylon: Trincomali
(BMNH). Andaman Islands: Aves I. (BMNH). Thailand: Ko
Chang, Trat Province, Gulf of Thailand (DC). Australia:
Queensland: Macoma Inlet, 20°9.6'S 148°55.3'E (AMS); New
South Wales: 12 km E of Cakora Point, S of Yamba (AMS);
Dudley, Newcastle (AMS); Toowoon Bay, The Entrance (AMS);
Port Stephens, 44 m (AMS); Careel Bay, Pittwater (AMS); Broken
Bay, Pittwater (AMS); Broken Bay, Patonga (AMS); Port Jackson
(BMNH); Balmoral, Middle Harbour, Sydney, 5 m (AMS); off
Morts Dock, Balmain, Port Jackson (AMS); Middle Harbour,
Chinaman’s Beach (AMS); Narrabeen Lagoon, Sydney (AMS);
Quarantine Bay, North Head, Sydney Harbour (AMS); Collaroy
Beach Sydney (AMS); Port Hacking, Bundeena, Simpsons Beach
(AMS); Port Hacking, Gunnamatta Bay (AMS); Cronulla (AMS);
off Montague Is, Narooma (AMS); Boydtown Beach, Twofold
Bay (AMS). Philippines: 11°43'N 122°34'E, 93 m (MNHN);
Magellan Bay, Mactan I., Cebu (BMNH). New Caledonia: Bay
of Ouanap, 0-2 m, 20°34'S 164°16'E (MNHN). Japan: Hirado
Hizen, Kyushu (BMNH); Kamakura Beach, Sagami Bay, Honshu
(AMS); Okinawa, Ryukyu Is (AMS). China: Hoi Ha Wan, Tolo
Channel, Hong Kong (NMW).
Pillucina neglecta Habe, 1960
Figs. 5a-d, 8
Pillucina neglecta Habe, 1960: 282, figs. 7-9. Holotype: H 2.4, L
2.2, and paratypes NSMT—Mo38710. Type locality: Tanabe
Bay, Wakayama Prefecture, Japan.
Description. Shell very small (H to 2.5) and rather inflated.
Sculpture of 30-40 radial ribs which often bifurcate, crossed
by fine commarginal lamellae, both of which are more
pronounced towards the posterior and anterior. Median part
of shell often worn and sculpture indistinct. Lunule smooth
and distinctly impressed. Left valve with two solid cardinal
teeth and anterior and posterior lateral teeth. Right valve
with a single, solid cardinal tooth, a large peg-like anterior
lateral tooth and long, posterior lateral tooth. Anterior
adductor muscle scar short and only slightly separated from
pallial line. Interior margin finely crenulate.
Remarks. This species is similar to P. pisidium but can be
distinguished by the smaller size, greater inflation, more
deeply impressed lunule and the presence of a strong lateral
tooth in the right valve. It is also similar to P. australis but
is longer, with a shorter, deeper lunule and fewer ribs.
Habitat. Intertidal mud (Habe, 1960).
Distribution. Japan, see Higo, Callomon & Goto (1999).
Material examined. Japan: Okinawa, Ryukyu Is (AMS).
Pillucina australis n.sp.
Figs. 6a-f, 8
type material. Holotype: NMV F83659; H 2.9, L 3.1.
Paratype: H 3.3, L 3.2, NMV F87506 (other specimens in
lot F87507). Type locality: Port Lincoln, Eyre Peninsula,
South Australia, 34°44'S 135°52'E, J. Veitch collection.
Description. Shell very small, H to 4.8, moderately inflated,
circular to sub circular in outline (H/L 1.02). Sculpture of
fine, closely-spaced commarginal lamellae crossed by 35-
45 low radial ribs. Radial ribs sometimes bifurcating
particularly at the anterior of the shell, more prominent and
slightly broader towards the anterior and posterior. Radial
sculpture absent from the umbones. Lunule shallowly
impressed and lanceolate in outline. Ligament internal,
extremely short. Hinge plate narrow, right valve with single
large cardinal tooth and single anterior and posterior lateral
teeth. Left valve with two cardinal teeth, anterior slightly
larger; posterior lateral is a narrow groove and anterior
lateral a shallow socket. Anterior adductor scar short and
rounded, only barely detached from pallial line. Shell
margin finely crenulate, often coarser towards the posterior.
Colour white.
Habitat. Intertidal to shallow water in sand.
Distribution. South Australia to Western Australia as far
north as Port Hedland and Scott Reef (Fig. 8).
Remarks. Although specimens of this species from
localities in South Australia and Western Australia are
present in museum collections (AMS, NMV, WAM) it has
not been mentioned in earlier literature (see Cotton &
Godfrey, 1938), or even recorded as Pillucina pisidium (=
symbolica) which it resembles. However, it differs from P
pisidium in its smaller size, fewer radial ribs, about 35-45
compared to more than 50 ribs on P. pisidium, and an
anterior lateral tooth in the right valve which is lacking in
P. pisidium. The two species also have a disjunct distribution
with seemingly no Pillucina species recorded from Victoria
and Tasmania, either in collections or literature.
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 271
Figure 5. Pillucina neglecta Habe, Okinawa, Japan (AMS C355657); a, exterior of left valve; b, exterior of left
valve; c, interior of right valve; d, interior of right valve. Scale bar =1.0 mm.
Material examined. Australia: South Australia: Outer
Harbour Beach, Adelaide (AMS); Largs Bay, St Vincents Gulf
(AMS); Point Sinclair (AMS); Tumby Bay, Spencer Gulf (AMS);
Arno Bay, Eyre Peninsula, Spencer Gulf (AMS); Hardwicke Bay,
Spencer Gulf (AMS); Henley 34°56'S 138°31'E (AMS); Western
Australia: E of Observatory Point, Recherche Archipelago
(AMS); Esperance (AMS); Oyster Harbour, Emu Point, Albany
(AMS); South Point, Two Peoples Bay, Albany (AMS); Yallingup
33°39'S 115° l'E, intertidal; Dunsborough 33°36'S 115°6'E(AMS);
Peppermint Grove beach near Capel, 6 m (AMS); Mandurah,
32°32'S 115°43'E (AMS); Point Peron, S of Perth (AMS); Ricey
Beach, Rottnest I. (WAM); E of Buller I., Grey, 6 m (WAM);
Cervantes, Hansen Bay (WAM); W of Green Head 29°50'S
114°7'E, 50 m (AMS); Dongara 29°9'S 114°43'E (AMS); Jurien,
Murchison River (WAM); Hamelin Pool, Shark Bay (WAM); 10
km SE of Faure I., Shark Bay (AMS); Freycinet Reach, Shark
Bay, intertidal sand (WAM); Bernier I., S of Redcliff Point, Shark
Bay, 10 m (WAM); Carnarvon, 6 m (AMS); North West Cape
21°44'S 114°19'E (AMS); North West Cape, S of Vlamingh
(WAM); Port Hedland (AMS).
Pillucina denticula n.sp.
Fig. 7a-g
Type material. Holotype: NMSAB310/T1758; H 3.5, L
3.5. Paratypes: NMSA V8402/T1759; H 2.8, L 2.9; H 2.9,
L 2.8; H 3.1, L 3.1. BMNH 2000377; H 3.7, L 3.5; H 3.5, L
3.5. Type locality: Durban Bay, South Africa.
Description. Shell small (H to 3.7), robust, inflated (H/L
1.0). Circular in outline. Sculpture of fine, closely spaced,
commarginal lamellae crossed by low, rounded radial ribs
that are prominent and broader towards anterior and
posterior. Ribs inconspicuous in central part of shell. Lunule
long, shallowly impressed and lanceolate in outline.
Ligament internal, short. Right valve with single cardinal
tooth and prominent anterior and posterior lateral teeth. Left
valve with two cardinal teeth, and anterior and posterior
lateral teeth. In addition to the major teeth the hinge plate
between the internal ligament and posterior lateral tooth
bears subsidiary denticles and sockets. Left valve has 6-8
sawtooth-like denticles and a posterior narrow ridge
overlying the lateral tooth (Fig. If), and right valve has
corresponding series of irregular sockets (Fig. 7g). Anterior
adductor scar short and barely detached from the pallial
line. Shell margin crenulate, with crenulations more widely
spaced to anterior and posterior of shell. Colour white.
272 Records of the Australian Museum (2001) Vol. 53
Figure 6. a-f, Pillucina australis n.sp.; a, exterior right valve, holotype NMV F83659, Port Lincoln, South Australia;
b, paratype NMVF 87506, exterior of left valve; c,d, interior of left and right valves of holotype; e,f, interior of
right valve and exterior left valve of specimen from Jervoise Groyne, 1.5 km south of Woodmans Point, Cockburn
Sound, Western Australia (AMS C355475). Scale bars =1.0 mm.
Etymology. The name denticula refers to the denticles on
the hinge plate.
Habitat. Subtidal sand 18-50 m.
Distribution. Eastern South Africa (Fig. 8).
Remarks. Pillucina denticula is similar in shell form to P.
australis from southern and western Australia but easily
distinguished by the presence of the unusual denticles on
the hinge plate.
Material examined. South Africa: Kwazulu Natal, St
Lucia Lighthouse, 50 m (NMSA); Durban Bay Head, 18-
22 m (NMSA).
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 273
Figure 7. a-g. Pillucina denticula n.sp.; a,b,c, exterior of left valve and interior of right and left valves, holotype
NMSA, Durban Bay, South Africa; scale bar =1.0 mm. d,e, exterior of right valve and interior of left valve,
paratype NMSA; scale bar =1.0 mm. f,g, detail of hinges of left and right valves of holotype showing the unusual
denticles on the hinge plate; scale bar = 250 pm.
Pillucina vietnamica Zorina, 1978
Figs. 9a-g, 10-13, 14a-d,f, 15a,c,d, 16
Pillucina vietnamica Zorina, 1978: 195, figs. 3 & 6 (fig. 6 seems
to be the interior of fig. 3 rather than the cited fig. 4 which is
the inside of fig. 5). Syntypes: ZISP, 13 whole shells and 1
valve, L 5.5-8.9. Type locality: intertidal, south coast of Hainan,
China. Seen
Lucina fischeriana Issel, 1869: 83-84, pi. 1, fig. 8 (non L.
fischeriana Orbigny, 1845, a Jurassic fossil). Five syntypes:
MCG, possible figured syntype, H 10.7, L 12.1. Type locality:
Suez, Egypt. Seen.
Lucina concinna H. Adams, 1871: 791, pi. 48, fig. 14 (non L.
concinna Deshayes, 1857, Eocene fossil). Holotype: ZMC; H
9.1, L 9.2. Type locality: Gulf of Suez, Red Sea. Seen.
Pillucina fischeriana (Issel).-Oliver, 1992: 98, pi. 20, fig. 4; Oliver,
1995: 236, fig. 1026.
Description. Shell small, H to 14, moderately inflated, shell
longer than high (H/L 0.9-0.95). Shell slightly translucent
and waxy in appearance. Extended anteriorly. Sculpture of
many, fine, low commarginal lamellae and fine radial ribs
which are broader and more prominent towards the anterior
and posterior. Intersection of commarginal and radial ribs
is finely cancellate in central part of shell and conspicuously
fluted where commarginal lamellae cross the broader radial
ribs towards anterior and posterior. Shell margin can appear
“scalloped” towards anterior and posterior. Lunule elongate,
274 Records of the Australian Museum (2001) Vol. 53
Figure 8. Map showing distribution of Pillucina pisidium (Dunker), P. neglecta Habe, P. australis n.sp. and P.
denticula n.sp.
lanceolate and impressed, slightly asymmetrical, with right
valve overlapping left. Hinge plate slightly sinuous, right
valve with single, narrow, cardinal tooth, anterior and
posterior lateral teeth short and peg-like. Left valve with
two narrow cardinal teeth, small, anterior lateral tooth and
posterior lateral tooth comprises a low, short ridge. Ligament
internal, short, situated on a broadly triangular resilifer.
Anterior adductor muscle scar medium-long, but narrow,
detached for slightly more than half of its length from pallial
line. Posterior scar ovate. Inner shell surface is fluted from
impressions of radial ribs especially in younger specimens.
Shell margin slightly thickened and crenulate, with
crenulations coarser towards anterior and posterior. Colour
yellowish to white.
Anatomy. The anatomy of P. vietnamica from Port Douglas,
Queensland was studied by dissection (Fig. 11), serial thin
sections and by SEM of critical point dried specimens. The
gross anatomy is generally similar to other species of
Lucinidae (Allen, 1958; Taylor & Glover, 2000).
Mantle. Three main folds of the mantle edge are present
(Fig. 12). There is a small outer fold, divided by the
periostracal groove from a middle fold which is composed
of two unequal lobes, the outer surface of the smaller outer
lobe forms the edge of the periostracal groove, whilst the
innermost lobe is large and muscular. Some distance back
from the mantle margin there is a small inner fold forming
a narrow, low ridge. Within the outer fold there is a large
blood space and within the inner part of the mantle edge
just below the line of pallial attachment there are groups of
large, blue-staining, subepithelial gland cells. Groups of
gland cells in this position within the inner mantle are seen
in other lucinids (Allen, 1958, fig. 2; Taylor & Glover, 2000,
fig. 7).
Mantle gills (putative respiratory structures situated near
the anterior adductor muscle) or a mantle septum as
described for lucinids such as Codakia, Phacoides, Lucina
and Anodontia (Taylor & Glover, 2000) are absent.
However, the inner surface of the mantle in the area
surrounding the anterior and ventral end of the anterior
adductor muscle and also the surface of the muscle itself
are covered by abundant ciliary tufts (Fig. 15c).
Ctenidia. As in most lucinids, the ctenidia consist of inner
demibranchs only, these are large and thick. The posterior
end of each ctenidial demibranch is joined laterally and
ventrally to the mantle edge by a thin tissue connection
which effectively partitions the inhalant and exhalant
apertures (Fig. 11). The gill filaments are each around 20-
25 pm in width. In section, there is a short outer ciliated
zone with the usual bands of frontal, laterofrontal and lateral
cilia (Fig. 14a,b). This is followed inwards by a narrow
intermediate zone of 2-3 cells, and then the thick lateral
zone comprising bacteriocytes and intercalary cells which
compose the greater part of the filament. The symbiotic
bacteria in P. vietnamica are contained in bacteriocytes and
have the form of elongate rods 6-8 pm long and 1.5-2 pm
in width (Fig. 14c,d). The bacteriocytes also contain
spherical granules around 5-7 pm in diameter which are
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 275
Figure 9. Pillucina vietnamica Zorina and P. angela (Melvill). a-d, P. vietnamica syntypes south coast of Hainan,
China, ZISP; a, exterior of right valve; b, interior of left valve; c, interior of right valve; d, exterior of left valve, e-
g, P. vietnamica, Jazirat as Sa’diyat, Abu Dhabi BMNH; e, interior of left valve; f, exterior of left valve; g, detail of
hinge of left valve. h,i, Pillucina angela (Melvill) figured syntype. Gwadur, Pakistan, BMNH 1899.12.18.20; h,
interior right valve; i, exterior of left valve. Scale bars =1.0 mm.
known in other lucinids to be sulphur-rich (Fig. 14f). The
labial palps of Pillucina are very small, as in most Lucinidae,
and consist of small, ciliated folds at the edge of the elongate
ciliated lips (Fig. 15d).
Foot. The foot is long and subcylindrical and lacks the
heel present in some lucinids. In section, it is composed of
layers of longitudinal, radial and circular muscles surrounding
a central blood space and is capable of considerable protraction.
The distal end of the foot is ciliated with abundant
subepithelial gland cells similar to those described for other
lucinids (Allen, 1958; Taylor & Glover, 1997a).
Mantle fusion and the posterior apertures. Characters
of the posterior apertures vary considerably between lucinid
genera (Allen, 1958, fig. 9) and may have value in
phylogenetic analysis. One of these characters is the extent
of mantle fusion ventral to the inhalant aperture and this
may vary from very short as in Codakia, to long as in
Anodontia. In Pillucina vietnamica, the fused section of
mantle is relatively long with only the inhalant aperture
edged with 6 papillae on either side (Fig. 15a), similar to
276 Records of the Australian Museum (2001) Vol. 53
Figure 10. Pillucina vietnamica Zorina from Queensland, Australia, a-d, from Port Douglas, Queensland BMNH
accession 2394; a, exterior of of right valve; b, exterior of left valve; c, interior of right valve; d, interior of left
valve. e,f, specimen from Redland, Moreton Bay, Queensland (BMNH accession 2394); e, interior of right valve;
f, exterior of left valve. Scale bars =1.0 mm.
the condition in Lucinoma borealis (Allen, 1958, fig. 9c).
Sections of the fused mantle near the inhalant aperture (Fig.
13b) show the extent of mantle fusion involving both the
inner and most of the middle mantle folds. The large blood
spaces indicate that this part of the mantle may be
considerably expanded and contracted. The exhalant
aperture comprises a thin-walled retractable tube which in
sections was seen inverted into the suprabranchial space
between the two inner demibranchs (Fig. 13b).
Distribution. Tropical Indo-West Pacific from Red Sea to
China and southern Queensland (Fig. 16).
Habitat. Intertidal to shallow water, sand and silt on the
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs
Figure 11. Pillucina vietnamica Zorina from Port Douglas, Queensland, general anatomy with right gill demibranch
nearly completely removed. Abbreviations: aa, anterior adductor muscle; au, auricle; bw, body wall; ct, ctenidia;
ext, exhalant tube; f, foot; ina, inhalant aperture; k, kidney; lig, ligament; lp, labial palp; me, mantle connection to
gill; me, mantle edge; pa, posterior adductor muscle; r, rectum.
Figure 12. Pillucina vietnamica Zorina from Port Douglas, Queensland, section through the mantle edge showing
the structure of the mantle folds. Abbreviations: bs, blood space; glc, subepithelial gland cells; imf, inner mantle
fold; lm, longitudinal muscles; mmf, middle mantle fold; omf, outer mantle fold; p, periostracum; pat, pallial
attachment; pg, periostracal groove; pn, pallial nerve; rm, radial muscles.
278 Records of the Australian Museum (2001) Vol. 53
Figure 13. a,b, Pillucina vietnamica Zorina from Port Douglas, Queensland; a, semi-diagrammatic drawing of
transverse serial section through median part of the body; b, semi-diagrammatic drawing of transverse section
through the posterior of body. Abbreviations: au, auricle; bs, blood space; ct, ctenidia; dmi, dorsal mantle isthmus;
elig, external ligament; ext, exhalant tube; f, foot; fm, fused mantle; ilig, inner ligament; int, intestine; k, kidney; m,
mantle; me, mantle edge; ov, ovary; pat, pallial attachment; pedal ganglion; prm, pedal retractor muscle; r, rectum.
fringes of mangroves; a species of continental margins rather
than offshore islands.
Remarks. This species is usually known from localities
around the Arabian Peninsula as Pillucina fischeriana (Issel,
1869) (see Oliver, 1992; 1995), however, the name is
unfortunately preoccupied, as is the name L. concinna H.
Adams, 1871. The next available name appears to be
Pillucina vietnamica Zorina, 1978 with the type locality of
Hainan I., China. We have examined the syntype series of
this species and they are similar in shell morphology to
samples from the Arabian Peninsula, Southeast Asia and
northern and eastern Australia and we regard them as
conspecific. Oliver (1992) regarded Pillucina concinna as
a distinct species from the Red Sea, however, we have
examined the type material and consider it a slightly more
inflated variant of P. vietnamica.
Individuals of P. vietnamica from the northern Red Sea
are larger than those from other localities as are shells from
Moreton Bay, Queensland at the southern edge of its range.
Pillucina vietnamica has never previously been recorded
from Australia although unidentified shells in museum
collections and our own sampling of live animals from
mangroves show that this species is widely distributed on
the Queensland coast.
Material examined. Egypt: Suez (BMNH). Yemen: Aden (BMNH).
Kuwait: (BMNH). Bahrain: Tubli Bay (NMW). Emirate of Abu
Dhabi: Dhafra Beach, Jebel Dhanna (BMNH). Emirate of Ras A1
Khaymah: Rams (BMNH). Saudi Arabia: Tarut, Arabian Gulf
(BMNH). Oman: Masirah I. (BMNH); Umm Rusays, Masirah;
Quriyat, Gulf of Oman (NMW); Qurm, Muscat. Somalia: Loyada
Beach (MNHN). Pakistan: Karachi (BMNH). India: Madras
(BMNH); Krusadai, Kutical Bay (BMNH); Kunugal Bay (BMNH).
Sri Lanka: Trincomalee (BMNH & MNHN). Malaysia: Kuantan,
intertidal sandflats 3°48'N 103°20'E (AMS); Country Club, Pulau
Langkawi, NW Malaysia (AMS). Cambodia: 5 km E of Port
Sihanoukville (BMNH). Australia: Western Australia: Parry
Harbour, Kimberley 13°58'S 126°05'E (WAM); Mission Bay, Napier
Broome Bay, Kimberley; intertidal (WAM). Northern Territory:
East Point, Darwin (BMNH); Snake Bay, near Timarambu Creek,
Melville I. (BMNH); Cape Consor, Melville I. (AMS); Crocodile
Research Station, Maningrida, Amhemland (AMS); Port Essington
(AMS); Crab Cay, Boucot Bay (AMS); Gove (NMV); Groote Eylandt,
Gulf of Carpentaria (AMS). Queensland: Forsyth I., Gulf of
Carpentaria; Friday I., Torres Strait 10°36'S 142°10'E (AMS);
Somerset, Cape York (AMS); Seisia, Bamaga, Cape York (NMV);
Caims Reef lagoon, 9-18 m, 14°42'S 145°30E, N of Cooktown
(AMS); Craigbe, Port Douglas (BMNH); Buchans Point, N of Caims
16°44'S 145°40'E (AMS); Halfmoon Bay, Yorkeys Knob (AMS);
Michaelmas Cay, Cairns (AMS); Port Denison, Bowen (AMS);
Mission Beach, N of Tully (AMS); Lucinda (AMS); Kings Beach,
Bowen (AMS); Seaforth, Mackay (AMS); Sarina beach (AMS);
Yeppoon (AMS); Quoin Is, Port Curtis, 2-5 m (AMS); Pialba, Hervey
Bay 25°17'S 152°50'E (AMS); Dundowran Beach, Hervey Bay
(AMS); Point Vernon, Hervey Bay (AMS); Tin Can Bay, NE of
Gympie (AMS); Noosa Heads (AMS); Redland Bay (BMNH);
Nudgee Beach, Moreton Bay (BMNH); Coochiemudlo I., SW of
Cleveland, Moreton Bay (AMS); Sandgate, Moreton Bay (NMV);
Woody Point, Moreton Bay (NMV); Peel I., Moreton Bay (AMS);
Southport, 27°58'S 153°25'E (AMS). China: Hong Kong (BMNH);
Daya Bay, Guangdong Prov (BMNH); Hainan (MNHN).
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 279
Figure 14. a-d. Pillucina vietnamica Zorina from Port Douglas, Queensland, all SEMs of critical point dried
preparations; a, distal tip of gill filament showing frontal, laterofrontal and lateral cilia, scale bar =10 pm; b,
section through three gill filaments showing ciliated zone, intermediary zone and bacteriocyte zone, scale bar = 20
pm; c, section through bacteriocyte showing rod shaped bacteria, scale bar = 5 pm; d, section through gill filament
showing bacteria in vacuoles within bacteriocytes, scale bar =10 pm. e, Wallucina assimilis Jervis Bay, NSW,
section through gill filaments showing bacteria in bacteriocyte zone, scale bar =10 pm. f, Pillucina vietnamica
spherical granules in bacteriocytes, scale bar =10 pm. Abbreviation: cz, ciliated zone; iz, intermediate zone; lz,
lateral zone.
Pillucina angela (Melvill, 1899)
Fig. 9h,i
Lucina (Codakia) angela Melvill, 1899: 98, pi. 2, fig. 8. Two
syntypes: BMNH 1899.12.18.20-21; figured syntype: H 7.9,
L8.1,T 1.9(lv); 1 syntype: NMW 1955.158.684. Type locality:
Gwadur, Pakistan, 8 fathoms. Seen.
Divaricella cypselis Melvill, 1918: 156, pi. 5, fig. 33. Holotype:
BMNH 1921.1.28.42; H 5.1, L 5.2, T 3.5 (2v). Type locality:
Karachi, Pakistan. Seen.
Pillucina angela (Melvill)-Oliver, 1995: 236, fig. 1025.
Description. Shells small (H to 8.0), subcircular (H/L 0.91),
inflated. Sculpture of strong radial ribs which are broader
and more widely spaced to the anterior and posterior. Ribs
steeply divaricate in anterior third of shell. Ribs crossed by
fine, closely spaced, commarginal lamellae which curve
over ribs producing a scalloped appearance. Anterodorsal
area extended above hinge line. Right valve with single
large cardinal tooth and short anterior and posterior lateral
teeth. Left valve with two cardinal teeth and anterior and
posterior lateral teeth. Ligament, internal, short. Anterior
adductor scar narrow, elongate, detached from pallial line
for about half of length. Inner shell margin coarsely
crenulate.
Distribution. Oman, northern Arabian Sea.
280 Records of the Australian Museum (2001) Vol. 53
Figure 15. a, Pillucina vietnamica Zorina, Port Douglas, Queensland, posterior apertures showing mantle fusion,
scale bar =1.0 mm. b, Wallucina assimilis, Jervis Bay, NSW posterior apertures, scale bar =1.0 mm. c,d, Pillucina
vietnamica ; c, inner surface of anterior mantle showing ciliary tufts and ciliated surface of anterior adductor muscle,
scale bar = 200 pm; d, ventral view of mouth, lips and labial palps, scale bar = 200 pm. Abbreviations: exa, exhalant
aperture; fm, fused mantle; ina, inhalant aperture; lp, labial palps; m, mouth.
Figure 16. Map showing distribution of Pillucina vietnamica Zorina.
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 281
Remarks. This species is similar to Pillucina vietnamica
and may represent an extreme morphological variant of that
species. However, from the evidence of the few shells we
have seen it differs from P. vietnamica in the stronger
divaricate radial ribbing, the extended anterodorsal area,
the more robust cardinal teeth and the longer and more
detached anterior adductor muscle scar. The holotype of
Divaricella cypselis Melvill, 1918 from Karachi is similar.
Material examined. Oman: Sur Masirah, sand flats (NMW
1993, figured in Oliver, 1995) plus 5 other lots from Masirah
(NMW). Pakistan: Gwadur (BMNH, NMW); Karachi
(BMNH).
Pillucina mauritiana n.sp.
Fig. 17
Loripes erythraeus (Issel, 1869).-Viader, 1951: 132, pi. 3, fig. 12.
Type material. Holotype: BMNH 20000079 paired valves
H 11.6, L 11.2, T. 4.0 (single valve). Type locality: dredged
Grande Passe, Cannoniers Point, Mauritius, 1947, Viader
collection. Paratypes: BMNH 20000080/1-7. Six paired
valves and single left valve, locality as holotype.
Etymology. Named for the island of Mauritius.
Description. Shell small (H to 11.6), thin, subcircular in
outline (H/L 1.0), some individuals slightly higher than long,
not inflated (T/H 0.3), although larger individuals tend to
be more inflated than juveniles. Sculpture of extremely fine,
low, commarginal lamellae with variable, fine, radial striae
that are slightly broader and more prominent near the dorsal
margin on both sides of the shell. Lunule narrow, lanceolate
to heart-shaped, slightly impressed. Hinge plate shallow.
Right valve with a single prominent cardinal tooth, an
elongate, posterior lateral tooth and a peg-like anterior
lateral tooth. Left valve with two cardinal teeth, a low
posterior lateral tooth and a socket for the anterior lateral
tooth of the right valve. Ligament internal, fairly short,
deeply impressed into each valve. Anterior adductor muscle
scar short, broad and detached from the pallial line for about
1/2 of its length; the posterior scar is reniform. Pallial line
discontinuous and impression of pallial blood vessel faint.
Shell outside of the pallial line glossy with fine radial
striations. Shell margin finely denticulate.
Remarks. Viader placed this species from Mauritius in what
we now call Chavania erythraea (see below) but it differs
in shape, hinge and external sculpture. Although the type
material is slightly worn the shells are distinctive and differ
from other Pillucina species. The species is most similar to
Pillucina hawaiiensis and P. pacifica, but it is larger, less
Figure 17. Pillucina mauritiana n.sp., holotype BMNH 20000079, Grande Passe, Cannoniers’ Point, Mauritius: a,
exterior of left valve; b, interior of left valve, c, exterior of right valve; d, interior of right valve; scale bar =1.0 mm.
282 Records of the Australian Museum (2001) Vol. 53
inflated, with a prominent anterior lateral tooth in the right
valve, a longer anterior adductor muscle scar and finer radial
ornament.
Distribution. Known only from Mauritius.
Material examined. Mauritius: Mauritius (MNHN).
Chavania n.gen.
Type species. Lucina erythraea Issel, 1869.
Diagnosis. Shells small, robust, H to 12. Outline generally
circular to slightly higher than long but rather variable.
Sculpture of fine, closely-spaced, low commarginal lamellae
which anastomose occasionally (Fig. 18), fine riblets
somet im es visible on anterior and posterior of shell and
also fine radial striations visible within the outer shell layer.
Shallow posterior sulcus present on both valves (Fig. 18).
Cardinal teeth large, with a single projecting tooth in the
right valve and two cardinal teeth in left valve of which the
anterior is larger. Anterior lateral teeth absent or vestigial
ridge. Ligament internal and short, deeply inset into valve.
Anterior adductor scar short and rounded and detached from
pallial line for about 1/5 to 1/3 of length. Shell margin broad,
thickened with the inner margin finely crenulate.
Remarks. Differs from Pillucina in the lack of prominent
radial ribs (Table 1), from the Miocene Microloripes in form
of the commarginal lamellae and the absence of an anterior
lateral tooth, and from Wallucina in having a more robust,
inflated shell, and coarser and anastomosing, commarginal
sculpture.
Etymology. Named in recognition of Andre Chavan for his
work on lucinid systematics.
Distribution. Indo-West Pacific (Fig. 21)
Chavania erythraea (Issel, 1869)
Figs. 18i-k, 21
Lucina erythraea Issel, 1869: 84, pi. 1, fig. 9. Three syntypes:
MCG, largest H 4.1, L 4.5. Type locality Suez. Some Byne’s
disease but identifiable. Seen.
Lucina crosseana Issel, 1869: 255, pi. 3, fig. 3. Types: ?Turin
Museum, not located.
ILucina elegans H. Adams, 1871: 791, not figured. Types: not
located 7BMNH or ?MZC.
Phacoides malcolmensis Melvill, 1918: 156-157, pi. 5, fig. 32.
One syntype: BMNH 1921.1.28.41, Malcolm Inlet, Arabian
Gulf, 35 fathoms; 1 syntype: NMW Z. 1955.158.686. Seen.
Wallucina erythraea (Issel).-Oliver, 1992:98, pi. 20, fig. 3; Oliver,
1995: 236, fig. 1027.
Description. Shell small, H to 12, sub-circular, anterior and
posterior shell margins straight, posterior slightly concave.
Sculpture of fine, low, closely-spaced, commarginal
lamellae which occasionally intersect and divide; fine radial
riblets sometimes visible to anterior and posterior. Posterior
dorsal area demarcated by broad shallow sulcus and less
prominent commarginal lamellae. Lunule heart shaped and
impressed. Hinge plate narrow, right valve with single
cardinal tooth, elongate posterior lateral tooth and no
anterior lateral tooth. Left valve with two cardinal teeth, an
elongate posterior lateral tooth and no anterior lateral.
Ligament internal, short, deeply inset into the valve. Anterior
adductor scar short, rounded and detached from pallial line
for about 1/3 of its length. Inner shell margin finely
denticulate. Colour white.
Habitat. Shallow water to 200 m.
Distribution. Red Sea, Arabian Gulf, Arabian Sea (Fig. 21).
Remarks. This species is similar to Chavania striata
(Tokunaga) but can be distinguished by the less inflated
shell and the quadrate anterior and posterior shell margins.
Issel’s original description was based on three specimens
from Suez and Savigny’s figure (1817, pi. 8, fig. 8). Bouchet
& Danrigal (1982, fig. 20) illustrate a shell from Savigny’s
collection, but this is a Ctena species whereas Issel’s
specimens (MCG) match the original description. We have
not seen the type material of L. crosseana (Issel) but follow
Lamy (1916) and Oliver (1992) in synonymising this species
with C. erythraea. Evidence from the description suggests
that Lucina elegans H. Adams may be a junior synonym
but there was no original figure and the type material has
not been located.
Material examined. Egypt: Suez (BMNH); Gulf of Suez
(NMW); Tubya al-Bayda, Safaga Bay. Kuwait: (BMNH).
Oman: Urf, Masirah (NMW); Sur Masirah, Masirah
(NMW). Yemen: al-Murk island, Red Sea (DC); Aden
(MNHN); Ra’s Qusay’ir, Hadramawt (DC). Djibouti:
(MNHN).
Chavania striata (Tokunaga, 1906)
Figs. 18a-h, 19, 21
Lucina contraria Dunker, 1882: 215, pi. 13, figs. 12-14. Holotype:
MNB Type locality: Japan (non Lucina contraria Dunker, 1846
a West African species). Seen.
Lasaea striata Tokunaga, 1906: 53-54, pi. 3, fig. 14. Twenty-one
syntypes: UMUT CM13599 and CM13753-13772 size range
H 7.1-9.0, L 7.0-9.4. Type locality: Oji, Tokyo City
(Pleistocene fossil). Seen.
Phacoides minutus Thiele, 1930: 592, pi. 4, fig. 72. Three syntypes:
MNB 67727, H 2.5. Type locality: Warnbro Sound, Western
Australia. Seen, with Byne’s disease.
Wallucina lamyi Chavan, 1938: 227, fig. 19. A replacement
name for L. contraria Dunker, 1882. Habe, 1977: 126, pi.
24, figs. 1 & 2.
Pillucina (Wallucina) striata (Tokunaga, 1906).-Kuroda, Habe
& Oyama, 1971: 392, pi. 118, fig. 15.
Wallucina aff jacksoniensis (Smith, 1885).-Hickman & Barnes,
1999: 240, fig. la-e.
Description. Shell small, H to 12, sub-circular, but shape
variable (mean H/L 1.02), posterior margin often somewhat
truncate. Sculpture of fine, low, closely-spaced, com¬
marginal lamellae which occasionally intersect and divide;
commarginal lamellae slightly higher on posterior dorsal
margin. Fine radial riblets somet im es visible on anterior
ventral margin. Posterior dorsal area demarcated by a narrow
sulcus. Lunule heart shaped and impressed. Hinge plate
narrow, right valve with single cardinal tooth, elongate
posterior lateral tooth and no anterior lateral. Left valve
with two cardinal teeth, an elongate posterior lateral tooth
and no defined anterior lateral although there is sometimes
a vestigial ridge in that position. Ligament internal, short,
deeply inset into the valve. Anterior adductor scar short,
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 283
Figure 18. Chavania striata (Tokunaga) (a-h) and C. erythraea (Issel) (i-k). a,b, Lasaea striata Tokunaga syntype
UMUT CM13754, exterior and interior of left valve. c,d, Lasaea striata syntype UMUT CM13763, interior and
exterior of right valve, e-f, Chavania striata Japan, BMNH 1873.8.6.196; e, interior of right valve; f, exterior of
left valve. g,h, C. striata Atoll de Huon, New Caledonia, station 439, 39 m, MNHN; g, interior of right valve; h,
exterior of left valve, i, Chavania erythraea, Suez, Red Sea, exterior of right valve BMNH. j,k, C. erythraea Sur Masirah,
Masirah, Oman, NMW 1993.061.00425a; j, exterior of right valve; k, interior left valve. Scale bars = 1.0 mm.
rounded and detached from pallial line for about 1/5 of its
length. Inner shell margin finely crenulate. Colour white.
Anatomy. The only material available for study was a single
poorly-fixed individual from the Easter Group, Houtman
Abrolhos Is, Western Australia (BMNH). With a cylindrical
foot, large gill and tiny labial palps, the gross anatomy of
the animal (Fig. 20) is similar in most respects to Pillucina
vietnamica. A tissue connection occurs between the
demibranchs and posterior mantle margin. The fused mantle
anterior to the inhalant aperture is shorter than in P.
vietnamica or Wallucina assimilis and papillae are lacking
284 Records of the Australian Museum (2001) Vol. 53
Figure 19. Chavania striata (Tokunaga) from Australia, a-c, Watsons Bay, Lizard I., Queensland, 10.5 m, AMS
C355806; a, exterior of left valve; b, interior of right valve; c, interior of right valve, d-g, Fremantle, Western
Australia, 18-22 m, NMV TD15644; d, exterior of left valve; e, exterior of left valve; f, interior of right valve; g,
interior of right valve. Scale bars =1.0 mm.
around the aperture. An elongate glandular strip is located
along the mantle margin ventral to the anterior adductor
muscle.
Distribution. Central Indo-West Pacific—India to Japan,
Marshall Is, intertidal to 250 m (Fig. 20).
Remarks. The earliest available name is Lasaea striata
Tokunaga, a Pleistocene fossil from Japan. We have
examined 21 syntypes of this species that are similar to
living shells from the Indo-West Pacific. This species has
been known as Wallucina lamyi Chavan (a replacement
name for the preoccupied L. contraria Dunker) but Chavan
(1938) was apparently unaware of the earlier name.
Phacoides minutus Thiele seems to be the same species
although the original figure is poor and the syntypes are
corroded by Byne’s disease.
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 285
Figure 20. Chavania striata (Tokunaga) from the Houtman Abrolhos Is, Western Australia, showing general anatomy
and detail of exhalant and inhalant apertures. Abbreviations: aa, anterior adductor muscle; ct, ctenidia; exa, exhalant
aperture; f, foot; fm, fused mantle; gs, glandular strip; ina, inhalant aperture; k, kidney; lp, labial palp; me, mantle
connection; me, mantle edge; pa, posterior adductor muscle.
• C. striata
A C. erythraea
Figure 21. Map showing distribution of Chavania striata (Tokunaga) and C. erythraea (Issel).
286 Records of the Australian Museum (2001) Vol. 53
Where larger samples from single localities (for example
a few sites from northern Queensland) are available,
Chavania striata shows considerable shape variation and
this is likely typical of the species throughout its range.
However, most of our records listed below comprise only
single or few dead shells.
This is the species referred to by Hickman & Barnes
(1999, fig. la-e) as Wallucina aff. jacksoniensis (Smith)
from the Late Quaternary Herschell Limestone of Rottnest
I., Western Australia. They suggest that the species is now
extinct in the area but shells are frequent in offshore
sediment samples and live animals have been dredged
around the Abrolhos Is.
Material examined. India: Tuticorin (BMNH). Sri Lanka:
Mannar (BMNH). Thailand: Ko Samui, Gulf of Thailand
(DC). Australia: Western Australia: Cockbum Sound, 20
m (AMS); Rottnest I., 18 m (BMNH); Fremantle, 25 m (NMV);
Houtman Abrolhos Is, (many lots) (WAM); North West shelf
230 m, 14°50'S 121°49.4’E (AMS); North West Shelf, north
of PortHedland, 150 m, 18°25’S 119°0’E (AMS); North West
Shelf, NNW of Dampier, 110m, 19°29'S 116°E (AMS); North
West Shelf, N of Cape Leveque, 194 m, 14°12’S 123°2'E
(AMS). Queensland: Murray I., Torres Strait (AMS); Albany
Passage, Cape York Peninsula (AMS); Palfrey I., Lizard I.
(AMS); Bird Islet, Lizard I. (AMS); Rocky Point, Lizard I.
(AMS); SE of Lizard I., 35 m (AMS); Carter Reef (AMS);
Low Isles (AMS). New Caledonia: lie St Marie, Noumea
(AMS); Atoll de Surprise 18°25'S 163°11’E (MNHN); Atoll
de Huon 18°07’S 162°55’E (MNHN); Plateau de Chesterfield,
56 and 51 m, 19°25’S 158°22’E (MNHN); Secteur de Belep
19°43’S 163°57’E (MNHN); Isle des Pins, 31 m (MNHN).
Marshall Islands: Kwajalein Atoll (MNHN). China: Mirs Bay,
Hong Kong (AMS). Japan: Sea of Japan, 34°13’N 136°13'E
(BMNH); Bungo Strait, between Shikoku & Kyushu, dredged
(BMNH).
Literature record. Cocos Keeling Atoll: lagoon as
IWallucina sp. Maes, 1967: 59, pi. 22D.
Funafutia n.gen.
Type species. Lucina levukana Smith, 1885 (= L. oblonga
Hedley, 1899).
Diagnosis. Shells small, anteriorly extended, ornament of
prominent commarginal lamellae with finely striated
interspaces. Umbones prominent, lunule lanceolate,
impressed. Ligament internal, short, triangular. Two cardinal
teeth in left valve, one in right valve; anterior and posterior
lateral teeth prominent in both valves. Shell margin finely
crenulate.
Comparison with other genera. This genus is distinguished
from others with an internal ligament by the anteriorly
extended shell, the prominent, rounded commarginal
lamellae and the strong anterior and posterior lateral teeth
in both valves.
Etymology. Named for Funafuti Atoll, the type locality of
L. oblonga.
Funafutia levukana (Smith, 1885)
Figs. 22a-d, 23
Lucina levukana Smith, 1885: 181, pi. 13, fig. 6. Holotype: BMNH
1887.2.9.2778 (type with extreme Byne’s disease). Type
locality: Levuka, Fiji.
Lucina (Codakia) oblonga Hedley, 1899: 497, fig. 51. Two
syntypes: AMS C6157 2 right valves, H 3.1, L 4.5. Type
locality: Lagoon beach, Funafuti Atoll, Tuvalu, (non Lucina
oblonga Philippi, 1836).
Lucina funafutica Finlay, 1927: 529. Replacement name for L.
oblonga Hedley.
Description. Small, H to 6, longer than high (H/L 0.89)
and extended anteriorly. Sculpture of prominent, rounded
commarginal lamellae that are irregularly spaced. Within
the relatively wide interspaces, fine radial striations (Fig.
22a,b) may be more prominent in eroded specimens. Lunule
long, lanceolate and impressed; umbonal area prominent.
Right valve with single large cardinal tooth and
prominent posterior and anterior lateral teeth. Left valve
with two cardinal teeth and prominent sockets for the
lateral teeth of the right valve. Ligament internal, with
short, narrow, triangular resilifer. Anterior adductor scar
short and rounded and barely detached from the pallial line,
posterior scar ovate. Pallial line continuous. Shell margin
outside the pallial line finely and evenly crenulate. Colour
white or pink.
Habitat. Shallow water to 40 m.
Distribution. Indian Ocean and West Pacific, ?Japan (in
Higo et al., 1999 as Anodontia oblonga ). A species of
offshore island habitats, not found on continental margins
(Fig. 23).
Remarks. The two syntypes of Lucina levukana Smith,
1885 have entirely decayed from Byne’s disease and the
figures do not illustrate the distinctive hinge teeth or
sculpture. However, in the detailed description Smith
highlights the extreme anterior extension of the shell, the
prominent commarginal sculpture, the deep internal
ligament and the strong cardinal and lateral teeth. These
are all features shared by Lucina oblonga Hedley.
Material examined. Seychelles: Mahe, off Anse aux Pins,
20 m (BMNH); Aldabra Atoll, Main Channel (BMNH).
Madagascar: Tulear (MNHN). Reunion: 310 m, 21°22'S
55°47'E (MNHN). Mauritius: Riambel Lagoon; off Trou aux
Biches; Pointe Radeau (NMSA). Australia: Queensland:
Murray I., Torres Strait, 9-15 m (AMS); Friday I., Torres
Strait (AMS); Euston Reef, 21 m, 16°40’S 146°13’E (AMS);
Lindeman I., 37 m (AMS); North West I., Capricorn Group
(AMS); Masthead I., Capricorn Group 31-37 m (AMS);
Lady Elliott I. (AMS); Lady Musgrave I., Bunker Group
(AMS). New Caledonia: Loyalty Is, Lifou (AMS); Touho,
Grand Recif (MNHN); Passe de Koumac, 20°39’S 164° 14’E;
Chesterfield Bank (MNHN). Kiribati: Abaing lagoon
(AMS). Tuvalu: Funafuti Lagoon Beach, Funafuti Atoll
(AMS). Wallis Islands: 13°19’S 176°17'W (MNHN).
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 287
Figure 22. Funafutia levukana (Smith), a-d, Masthead I., Capricorn Group, Queensland, 23°32'S 151°45'E AMS
C019420, 31-37 m, Hedley collection; a, exterior of right valve; b, exterior of left valve; c, interior of left valve; d,
interior of right valve. Scale bars = 0.5 mm.
Genus Wallucina Iredale, 1930
Wallucina Iredale, 1930: 390.
Type species. Wallucina jacksoniensis Smith, 1885 (= W.
assimilis Angas, 1867).
Diagnosis. Shells small, subcircular in outline. Moderately
inflated, relatively thin shelled. Sculpture of fine closely
spaced commarginal lamellae, sometimes slightly raised to
posterior, fine radial striations sometimes present. Hinge
plate narrow. Right valve with single cardinal tooth and
posterior lateral tooth, anterior lateral absent or very small.
Left valve with two cardinal teeth and a posterior lateral
tooth. Ligament internal, set in elongate, oblique resilifer.
Anterior adductor muscle scar elongate, detached from
pallial line for about 1/2 to 1/3 of length. Pallial line
discontinuous. Inner shell margin finely crenulate.
Distribution. Central Indo-West Pacific and southern and
western Australia.
Comparison with other genera. Wallucina differs from
Pillucina in absence of radial ribs and from Chavania by
the much thinner shell, narrow hinge and absence of
anastomosing commarginal lamellae.
Wallucina assimilis (Angas, 1867)
Figs. 24c-g, 25, 26
Loripes assimilis Angas, 1867: 910, pi. 44, fig. 8. Two syntypes:
BMNH 1870.10.26.34; H 11.6, L 11.8, T 3.8; H 11.0, L 11.1,T
4.8. Type locality: as cited in original description, New South
Wales and Hobson’s Bay, Port Phillip, Victoria. Seen.
Lucina jacksoniensis Smith, 1885: 185, pi. 13, figs. 11 & lib.
Two syntypes: BMNH 1887.2.9.2784-5, (2 complete shells);
larger shell H 9.3, L 9.2, T 3.2. Type locality: Port Jackson,
288 Records of the Australian Museum (2001) Vol. 53
NSW, 6-15 fathoms. Seen.
Wallucina jacksoniensis (Smith).-Iredale, 1930: 390.
Wallucina icterica (Reeve, 1850).-Cotton & Godfrey, 1938: 203,
fig. 214. Lucina icterica Reeve, 1850 is a Semele (BMNH
1998170—see Ludbrook, 1955: 51).
Description. Shells small (H to 15), subcircular,
anteriorly extended, not inflated and rather thin-shelled.
Sculpture consists of extremely fine, low concentric
growth increments with variable, faint, fine radial folds
between the commarginal lamellae. Microscopically,
shells often have fine radial striations which have been
described as external sculpture but are in fact the internal
shell microstructure of the outer radial composite prismatic
layer. Lunule narrow, lanceolate to heart-shaped, not
impressed (in type) and slightly asymmetrical. Anterior
dorsal area demarcated by narrow, often indistinct sulcus.
Hinge plate shallow, right valve with single prominent
cardinal tooth and two smaller posterior laterals, anterior
lateral tooth tiny to indistinct. Left valve with two
cardinals, of which anterior cardinal is trigonal and
pronounced, posterior lateral small and anterior lateral
indistinct. Ligament internal, short and deeply impressed
into each valve, resilifer oblique. Anterior adductor
muscle scar short, rounded and detached for about 1/2
of its length. Pallial line usually discontinuous with
pallial blood vessel impression often faint. Shell margin
finely crenulate. Typically interior of shell chalky with
narrow radial ridges inside pallial line, whilst outside
pallial line shell margin is often glossy with fine radial
striations. Shell white.
Anatomy. The anatomy was studied from dissections and
SEM of critical point dried specimens from Jervis Bay, New
South Wales (BMNH). In general anatomical features (Fig.
25), W. assimilis is similar in most respects to Pillucina
vietnamica including the posterior mantle connection to the
ctenidia. The fused mantle ventral to the inhalant aperture
is relatively longer and the inhalant aperture lacks the
papillae of P. vietnamica (Fig. 15b). Sections of the lateral
zone of the gill filaments confirm the presence of abundant
rod-shaped bacteria (about 3-5 pm long and 1-1.5 pm wide)
contained in bacteriocytes (Fig. 14e).
Remarks. Samples of W. assimilis from Western Australia,
South Australia and NSW all show considerable variation
in shell outline, lunule depth and thic kn ess of the hinge and
size of cardinal teeth. The shells are often rather irregularly
shaped with a crumpled appearance. Because of the high
variation in shell morphology we can see no reason to
support Cotton & Godfrey’s (1938) separation of the
southern and western Australian shells as a separate species
W. icterica.
Distribution. From northern NSW to southern Western
Australia as far north as Ningaloo (Fig. 26). Barnes &
Hickman (1999) highlight the association of this species
with seagrass beds around Rottnest I., Western Australia
where they recorded densities of up to 1048 m' 2 amongst
Posidonia and Amphibolis. We have also collected living
W. assimilis from shallow seagrass beds at various localities
along the New South Wales coast.
Material examined. Australia: New South Wales: Port
Stephens (AMS); Pittwater (BMNH); Middle Harbour, Sydney
(NMV); Narrabeen lagoon (BMNH); Gunnamatta Bay, Port
Hacking (AMS); Jervis Bay (BMNH); Gerringong (AMS); Kiama
(AMS); Sussex Inlet (AMS); Lake Conjola (AMS); Twofold Bay
(AMS); Bendalong, Ulladulla (AMS); Narooma (NMV);
Merimbula (NMV); PambulaLake (AMS). Victoria: Mallacoota
(NMV); Point Nepean (NMV); Cable I., Bass Strait (NMV);
Flinders (NMV); Corio (NMV); Lakes Entrance, 20 fathoms
(NMV); Frankston (NMV); Port Albert (NMV); off Wilsons
Promontory (NMV); Point Cook, Port Philip Bay (NMV); Point
Leo, Mornington Peninsula (NMV). TASMANIA: Adelaide Bay,
Flinders I. (AMS); Logan’s Lagoon, Flinders I. (AMS); Tinderbox,
Derwent Estuary (AMS); Randalls Bay, Houn Estuary (NMV);
Eaglehawk Bay (AMS); Long Bay (AMS); Cremorne, Frederick
Henry Bay (AMS); Pittwater, Cambridge (AMS). SOUTH
Australia: Normanville, St Vincents Gulf (AMS); Outer
Harbour, Adelaide (SAM); St Vincents Gulf (SAM); Edithburg, 9
fathoms (SAM); Point Germein (SAM); Boston I. 10 fathoms
(NMV); St Francis I. (SAM). Western Australia: Kalgan
Bridge (AMS); Two Peoples Bay, Albany (AMS); E of Hood Point,
79 m, 34°21'S 121°16'E (AMS); King George Sound, Albany
(SAM); Windy Harbour (AMS); Nornalup inlet (AMS);
Cowaramup (AMS); Eagle Bay, W of Dunsborough (AMS);
Yallingup (SAM); Bunbury (SAM); Hopetown beach (SAM);
Swan River estuary (AMS); Cockburn Sound (AMS); Rottnest I.
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 289
Figure 24. Wallucina fijiensis (Smith) and W. assimilis (Angas). a,b, Wallucina fijiensis Eclipse I., Claremont
Group, Queensland, 13°29'S 143°43'E, AMS 007256; a, exterior of left valve; b, interior of right valve. c,d, W.
assimilis Narrawallee lagoon, Ulladulla (BMNH); c, exterior of right valve; d, interior of right valve, e, hinge of
right valve of W. assimilis Pittwater, New South Wales (BMNH). f,g, W. assimilis Robe, South Australia (SAM
D15648); f, exterior of right valve; g, interior of right valve. Scale bars =1.0 mm.
(SAM); Cervantes (AMS); Port Denison near Dongara (AMS); Wallucina fijiensis (Smith, 1885)
Geraldton (SAM); Horrocks (AMS); West Wallabi I., Houtman
Abrolhos (AMS); Kalbarri (AMS); Dirk Hartog I. (AMS); Point Figs. 24a,b, 26
Peron (AMS); Ningaloo Reef (AMS).
Lucina (Codakia) fijiensis Smith, 1885: 184, pi. 13, fig. 9.
Holotype: single valve with severe Byne’s disease BMNH
1887.2.9. Type locality: off Levuka, Fiji, 12 fathoms. Seen.
290 Records of the Australian Museum (2001) Vol. 53
Figure 25. Wallucina assimilis (Angas) from Jervis Bay, NSW, showing general anatomy with the gill intact.
Abbreviations: aa, anterior adductor muscle; bw, body wall; ct, ctenidia; exa, exhalant aperture; f, foot; g, gonad;
ina, inhalant aperture; lig, ligament; lp, labial palp; me, mantle connection to gill; pa, posterior adductor muscle; r,
rectum.
Figure 26. Map showing distribution of Wallucina fijiensis (Smith) and W. assimilis (Angas).
Glover & Taylor: Australian and Indo-Pacific lucinid molluscs 291
Lucina (?Loripes) gordoni Smith, 1885: 186, pi. 13, fig. 12.
Holotype: BMNH 1887.2.9.2786 single valve (with Byne’s
disease). Type locality: off Levuka, Fiji, 12 fathoms. Seen.
Loripes haddoni Melvill & Standen, 1899: 200, fig. 12. Holotype:
BMNH 1899.2.23.12; H 12.6, L 13.1, T 13.5. Type locality:
Warrior I., Torres Strait, Australia. Seen.
Wallucina gordoni (Smith).-Maes, 1967: 159, pi. 22, fig. e.
Description. Shell small, thin, H to 15, circular to
subcircular in outline; posterior shell margin slightly
truncate. Sculpture of extremely fine, low concentric growth
increments with variable, faint, fine, radial striae between
commarginal lamellae. Commarginal lamellae are slightly
broader and more elevated near dorsal margins. Lunule
narrow, lanceolate to heart-shaped and slightly excavated.
Anterior dorsal area often with irregular folds. Hinge plate
narrow; right valve with prominent cardinal tooth, two small
posterior lateral teeth and tiny anterior lateral tooth. Left
valve with two cardinals, anterior cardinal is trigonal and
larger, posterior laterals are low and anterior lateral absent.
Ligament internal, deeply impressed into each valve.
Anterior adductor muscle scar short, broad and detached
from the pallial line for about 1/2 of length. Pallial line
discontinuous and pallial blood vessel impression faint.
Inner surface of shell inside pallial line with indistinct
narrow, radial ridges, shell outside the pallial line glossy
with fine radial striations. Shell margin finely denticulate.
Colour white.
Distribution. Northern Queensland, Great Barrier Reef
islands, western Pacific, tropical southeast Asia, shoals and
islands on the north Western Australian shelf (Fig. 26).
Remarks. This species is similar to W. assimilis. The shells
are larger, with a more quadrate posterior margin and the
commarginal lamellae often elevated on the posterodorsal
margin. The lunule is deeper and shorter than in W. assimilis
and the ligament is longer. The two species have non¬
overlapping ranges; W. assimilis has temperate southerly
distribution ranging from NSW to Ningaloo in Western
Australia whilst W. fijiensis is a tropical species (Fig. 26).
The specimens from Cocos Keeling are larger, H to 17.5,
and have a yellow interior but in other respects are closely
similar.
Material examined. Cocos Keeling Islands: Pulo Bras, 2 m
(BMNH); Prison I., Cocos Keeling (WAM); reef slope off North
West I. (WAM). Australia: Western Australia: Bedwell I.,
Rowley Shoals (WAM); Eastern I., Ashmore Reef (WAM).
Queensland: Yorke I. (AMS); Bramble Cay (AMS); Friday I.,
Torres Strait (AMS); Cairncroft I., Torres Strait (BMNH);
Thursday I. (AMS); Murray I. (AMS); Hope I. (AMS); Lizard I.
(AMS); Two Isles, south of Cape Flattery (AMS); Eagle I., N of
Cooktown, 5.5 m (AMS); Low Isles (AMS); Wonga Beach
(BMNH); Opal Reef, N of Cairns (AMS); Green Is (AMS); Ellison
Reef off Kurrimine (AMS); Palm I. (AMS); Eclipse I. (AMS);
Hayman I. (AMS); Michaelmas Cay (AMS); Lady Elliott I.
(AMS); Facing I., Port Curtis (AMS). Thailand: Ban Thap Lamu,
Thai Muang, Phangna province (BMNH). Philippines: Magellan
Bay, west side of Punta Engano, Mactan I., Cebu (BMNH). New
Caledonia: Touho, Grand Recif Mengalia, 20°45'S 165°16'E
(MNHN). New Hebrides: (MNHN). Kiribati: Tarawa lagoon,
Kiribati 1°25'N 173°00'E (AMS); Kingsmill I. (BMNH); Teirio
Abaiang Atoll (AMS). Guam: East Agana Bay (AMS). Japan:
Edatekujima I., Anami Is, Japan. (BMNH).
Species excluded from Pillucina and Wallucina
Pillucina yamakawai (Yokoyama, 1920)
Lucina yamakawai Yokoyama, 1920: 135, pi. 10, fig. 9. Type
locality: Plio-Pleistocene fossil, Shimo-Miyata, Yokosuka City,
Honshu.
Pillucina (Sydlorina) yamakawai (Yokoyama) -Kuroda, Habe &
Oyama, 1971: 393, pi. 118, fig. 16; Habe, 1977: 126, pi. 24,
figs. 3 & 4.
This species probably belongs to the genus Parvilucina and
can be easily distinguished from Pillucina by the absence
of an internal ligament.
Wallucina izuensis Okutani & Matsukuma, 1982
Wallucina izuensis Okutani & Matsukuma, 1982: 174-175, pi.
10, figs. 7 & 8, text fig. 2.
This species from Japan has an external ligament and cannot
be included in our concept of the genus Wallucina.
Acknowledgments. We are grateful to the Australian
Biodiversity Resources Study (ABRS) which partially funded this
study. We thank Ian Loch (AMS), Sue Boyd (NMV), Wolfgang
Ziedler (SAM), Fred Wells and Shirley Slack Smith (WAM),
Philippe Bouchet, Virginie Heros, Rudo von Cosel (MNHN),
Graham Oliver (NMW), Matthias Glaubrecht (MNB), Boris
Sirenko (ZISP), Takenori Sasaki (UMUT), Guiliano Doria (MCG),
Reggie Kawamoto (BPBM), Dick Kilburn (NMSA), Richard
Preece (ZMC) and Henk Dekker for allowing us access to
collections in their care and for generous loan of types and other
material. Dave Cooper made the thin sections, Harry Taylor took
the macrophotographs, Rudo von Cosel arranged the photography
of the syntypes of Pillucina vietnamica and Alex Ball and Chris
Jones of the EM Unit (NHM) gave much assistance.
References
Adams, H., 1871. Descriptions of twenty-six new species of shells
collected by Robert Me Andrew, Esq., in the Red Sea. Proceedings
of the Zoological Society of London 1870: 788-793.
Allen, J.A., 1958. On the basic form and adaptations to habitat in
the Lucinacea (Eulamellibranchia). Philosophical Transactions
of the Royal Society of London B 241: 421^-84.
Allen, J.A., 1960. The ligament of the Lucinacea. Quarterly
Journal of Microscopical Science 101: 25-37.
Angas, G.F., 1867. Descriptions of thirty-two new species of
marine shells from the coast of New South Wales. Proceedings
of the Zoological Society of London 1867: 110-117.
Barnes, P.A.G., & C.S. Hickman, 1999. Lucinid bivalves and
marine angiosperms: a search for causal relationships. In The
Seagrass Flora and Fauna of Rottnest Island, Western
Australia , ed. D.I. Walker & F.E. Wells, pp. 215-238. Perth:
Western Australian Museum.
Bouchet, P, & F. Danrigal, 1982. Napoleon’s Egyptian campaign
(1798-1801) and the Savigny collection of shells. Nautilus 96:
9-24.
Bretsky, S.S., 1970. Phenetic and phylogenetic classifications of
the Lucinidae (Mollusca, Bivalvia). Bulletin of the Geological
Institute of the University ofUpsala, new series 2: 5-23.
Bretsky, S.S., 1976. Evolution and classification of the Lucinidae
(Mollusca; Bivalvia). Palaeontographica Americana 8(50):
219-337.
Chavan, A., 1937-1938. Essai critique de classification des lucines.
Journal de Conchyliologie 81: 237-281; 82: 215-241.
Chavan, A., 1969. Superfamily Lucinacea Fleming, 1828. In Treatise
on Invertebrate Paleontology, Part N, Mollusca 6, Bivalvia, vol.
292 Records of the Australian Museum (2001) Vol. 53
2, ed. R.C. Moore, pp. N491-N518. Boulder, Colorado: Geological
Society of America and University of Kansas.
Cotton, B.C., & F.K. Godfrey, 1938. The Molluscs of South
Australia. Parti. The Pelecypoda. Adelaide: South Australian
Branch of the British Science Guild.
Dali, W.H., R Bartsch & H.A. Rehder, 1938. A manual of the
Recent and fossil marine pelecypod mollusks of the Hawaiian
Islands. Bulletin of the Bernice P. Bishop Museum 153: 1-233.
Dell, R.K., 1964. Antarctic and subantarctic Mollusca: Amphineura,
Scaphopoda and Bivalvia. Discovery Reports 33: 93-250.
Distel, D.L., 1998. Evolution of chemoautotrophic endosymbioses
in bivalves. Bioscience 48: 277-286.
Dunker, W., 1860. Neue japanische Mollusken. Malakozoologische
Blatter 8: 150-154.
Dunker, W., 1861 . Mollusca Japonica Descripta et Tabulis Tribus
Iconum. Stuttgart, 1-36.
Dunker, W., 1882. Index Molluscorum Maris Japonici. Novitates
Conchologicae, Supplement 7: 1-301.
Finlay, H.J., 1927. New specific names for Austral Mollusca.
Transactions of the New Zealand Institute 57: 488-533.
Glover, E.A., & J.D. Taylor, 1997. New species and records of
Rastafaria and Megaxinus from the western Indian Ocean and
Red Sea, with a reappraisal of Megaxinus. Journal of
Conchology 36: 1-18.
Gould, A., 1861. Descriptions of shells collected in the North Pacific
Exploring Expedition under Captains Ringgold and Rodgers.
Proceedings of the Boston Society of Natural History 8: 33^4-0.
Habe, T., 1960. Eleven new bivalves from Tanabe Bay, Wakayama
Pref., Japan. Publications of the Seto Marine Biological
Laboratory 8: 281-288.
Habe, T., 1977. Systematics of Mollusca in Japan, Bivalvia and
Scaphopoda. Tokyo: Hokuryukan.
Hedley, C., 1899. The Mollusca of Funafuti. Part II. Pelecypoda and
Brachiopoda. Memoirs of the Australian Museum 3: 491-510.
Hedley, C., 1914. Studies on Australian Mollusca. Part XII.
Proceedings of the Linnean Society of New South Wales 39:
695-755.
Hickman, C.S., & P.A.G. Barnes, 1999. Fossil lucinid bivalves of
Rottnest Island: anomalous Late Quaternary geographic
distributions. In The Seagrass Flora and Fauna of Rottnest
Island, Western Australia, ed. D.I. Walker & F.E. Wells, pp.
239-245. Perth: Western Australian Museum.
Higo, S., P. Callomon & Y. Goto, 1999. Catalogue and
bibliography of the marine shell-bearing Mollusca of Japan.
Osaka: Elle Scientific Publications.
Iredale, T., 1930. More notes on the marine Mollusca of New
South Wales. Records of the Australian Museum 17: 384-407.
Issel, A., 1869. Malacologia del Mare Rosso, richerche zoologiche
e paleontologiche. Pisa, Italy.
Johnson, R., 1964. The Recent Mollusca of Augustus Addison Gould.
Bulletin of the United States National Museum 239: 1-182.
Kay, E.A., 1979. Hawaiian marine shells. Bernice P. Bishop
Museum Special Publication 64 (4): 1-652.
Kuroda, T., T. Habe & K. Oyama, 1971. The Seashells ofSagami
Bay. Tokyo, Maruzen.
Lamprell, K., & J. Healy, 1998. Bivalves of Australia Volume 2.
Leiden: Backhuys Publishers.
Lamprell, K., & T. Whitehead, 1992. Bivalves of Australia,
Bathurst, Australia: Crawford House Press.
Lamy, E., 1916. Les Lucines et les Diplodontes de la Mer Rouge
(d’ apres les materiaux recueillis par M. le Dr Jousseaume). Bulletin
du Museum d’Histoire naturelle 22: 145-155, 183-190.
Ludbrook, N.H., 1955. The molluscan fauna of the Pliocene strata
underlying the Adelaide plains. Transactions of the Royal
Society of South Australia 78: 18-87.
Maes, V.O., 1967. The littoral marine mollusks of Cocos Keeling
Islands (Indian Ocean). Proceedings of the Academy of Natural
Sciences of Philadelphia 119: 93-217.
Melvill, J.C., 1899. Notes on Mollusca from the Arabian Sea,
Persian Gulf, and Gulf of Oman, mostly dredged by Mr. F.W.
Townsend, with descriptions of twenty-seven species. Annals
and Magazine of Natural History, series 7, 4: 81-101.
Melvill, J.C., 1918. Descriptions of thirty-four species of marine
Mollusca from the Persian Gulf, Gulf of Oman and Arabian
Sea, collected by Mr. F.W. Townsend. Annals and Magazine
of Natural History series 9, 1: 137-158.
Melvill, J.C., & R. Standen, 1899. Report on the marine Mollusca
obtained during the first expedition of Professor A.C. Haddon
to the Torres Strait in 1888-89. Journal of the Linnean Society,
Zoology 27: 150-206.
Okutani, T., & A. Matsukuma, 1982. Some interesting molluscs
dredged from the shelf around the southern coast of the Izu
peninsula, Honshu, with descriptions of two new species.
Memoirs of the National Science Museum 15: 163-180.
Oliver, P.G., 1986. A new lucinid bivalve from the Niger Delta
and an appraisal of the Loripes group (Bivalvia, Lucinacea).
Basteria 50: 47-64.
Oliver, P.G., 1992. Bivalved seashells of the Red Sea. Wiesbaden &
Cardiff, Verlag Christa Hemmen & National Museum of Wales.
Oliver, P.G., 1995. Bivalvia. In Seashells of Eastern Arabia ed.
S.P Dance. Dubai, Motivate Publishing.
Pilsbry, H.A., 1921. Marine Mollusks of Hawaii—XIV, XV.
Proceedings of the Academy of Natural Sciences of Philadelphia
72:360-382.
Reeve, L.A., 1850. Monograph of the genus Lucina. Conchologica
Iconica vol 6. London: Reeve, Benham & Reeve.
Reid, R.G.B., 1990. Evolutionary implications of sulphide-
oxidising symbioses in bivalves. In The Bivalvia—Proceedings
of a Memorial Symposium in Honour of Sir Charles Maurice
Yonge, Edinburgh, 1986, ed. B. Morton, pp. 127-140. Hong
Kong: Hong Kong University Press.
Rodionov, I.A., & V.V. Yushin, 1991. Procaryotic symbionts in
gill cells of the bivalve mollusc Pillucina pisidium. Biologia
Morya N 1: 39-46 (in Russian).
Smith, E.A., 1885. Report on the Lamellibranchiata collected by
H.M.S. Challenger during the years 1873-76. Report of the
Scientific Results of the Voyage of H.M.S. Challenger 1873-
76. 13: 1-341.
Taylor, J.D., & E.A. Glover, 1997a. A chemosymbiotic lucinid
bivalve (Bivalvia: Lucinoidea) with periostracal pipes:
functional morphology and description of a new genus and
species. In The Marine Flora and Fauna of the Houtman
Abrolhos, Western Australia, ed. F.E. Wells, pp. 335-361. Perth:
Western Australian Museum.
Taylor, J.D., & E.A. Glover, 1997b. The lucinid bivalve genus
Cardiolucina (Mollusca, Bivalvia, Lucinidae): systematics,
anatomy and relationships. Bulletin of the Natural History
Museum, London (Zoology) 63: 93-122.
Taylor, J.D., & E.A. Glover, 2000. Functional anatomy,
chemosymbiosis and evolution of the Lucinidae. In The
Evolutionary Biology of the Bivalvia, ed. E.M. Harper, J.D.
Taylor & J.A. Crame, Geological Society Special Publication
111: 207-225.
Thiele, J., 1930. Gastropoda und Bivalvia. In Die Fauna
Sudwestaustraliens, ed. W. Michaelson & R. Hartmeyer, pp.
561-596. Jena: Gustav Fischer.
Tokunaga, S., 1906. Fossils from the environs of Tokyo. Journal of
the College of Science, Imperial University, Tokyo 21: 1-96.
Viader, R., 1951. New or unrecorded shells from Mauritius and
its dependencies. Mauritius Institute Bulletin 3: 127-153.
Yokoyama, M., 1920. Fossils from the Miura Peninsula and its
immediate north. Journal of the College of Science, Imperial
University, Tokyo 39: 1—198.
Zorina, I.P, 1978. New species of bivalve molluscs (Bivalvia) of
the Gulf of Tonkin (South China Sea). Trudy Zoological Institut,
Leningrad 61: 193-203 (in Russian).
Manuscript received 9 June 2000, revised 12 October 2000 and accepted
13 October 2000.
Associate Editor: G.D.F. Wilson.
© Copyright Australian Museum, 2001
Records of the Australian Museum (2001) Vol. 53: 293-340. ISSN 0067-1975
Revision of the Australian Seahorses of the Genus
Hippocampus (Syngnathiformes: Syngnathidae)
with Descriptions of Nine New Species
Rudie H. Kuiter
Ichthyology, Museum Victoria, Melbourne Victoria 3001, Australia
rudie.kuiter@zoonetics.com • syngnathiformes@zoonetics.com
Abstract. Australian species of seahorses (genus Hippocampus ) are reviewed and descriptions of nine
new species from Australian and adjacent waters are presented: H. jugumus n.sp. (Lord Howe Island),
H. biocellatus n.sp. (Shark Bay, Western Australia), H. alatus n.sp. (northern Australia), H. semispinosus
n.sp. (Timor Sea, southern Indonesia), H. montebelloensis n.sp. (Monte Bello Islands, Western Australia),
H. procerus n.sp. (tropical eastern Australia), H. multispinus n.sp. (northern Australia), H. hendriki n.sp.
(northeastern Queensland), and H. grandiceps n.sp. (Gulf of Carpentaria). A total of twenty-four species
have been collected in Australian waters, and additional species may be found as these are known to
occur in neighbouring waters. Diagnosis and a key for the Australian species are provided.
Kuiter, Rudie H., 2001. Revision of the Australian seahorses of the genus Hippocampus (Syngnathiformes:
Syngnathidae) with descriptions of nine new species. Records of the Australian Museum 53(3): 293-340.
In compiling information for a book on the syngnathid fishes
of the world (Kuiter, 2000), it became clear that despite
recent studies and the publication of a book on the taxonomy
of seahorses (Lourie et al., 1999), confusion remains.
Eschmeyer (1998) lists 112 nominal species worldwide,
many representing the same taxon and others wrongly
synonymised since their initial original publication. An
identification guide to the world’s syngnathid fishes (Kuiter,
2000) suggests over 60 valid seahorses species worldwide.
Applying the correct scientific name to seahorse species
has always been problematic. In Australia, Whitley & Allan
(1958) recognised eight species and suggested that there
are about 100 valid species worldwide. Paxton et al. (1989)
recognised 9 Australian species; and, Gomon, 1997, added
a new species. An identification guide to the world’s species
(Lourie et al., 1999) recognised only 32 species worldwide,
but did not deal with species-complexes. The difficulty in
working with seahorses stems primarily from their peculiar
morphology, with the absence of many of the characters
that fish taxonomists traditionally rely on to distinguish
species. In examining material from Australian collections
for this revision, specimens of the same taxon were
frequently found to be identified as a variety of species,
often in relation to their relative presence or absence of
spines. This work revealed that some species with little spine
development in adults have spiny juveniles. The names H.
kuda long applied to many smooth species and H. histrix
frequently used for spiny species in tropical regions has
given rise to the perception of wide-spread distributions,
but it is clear that most species are highly localised and that
there are a number of species-complexes whose members
are variously distributed in different faunal regions.
Australia supports a number of different bioregions that
range from temperate to tropical, and have Pacific or Indian
Ocean origins. The pelagic regions are generally divided
into four zones: north, south, east and west, whilst demersal
294 Records of the Australian Museum (2001) Vol. 53
areas are often localised to small coastal sections that are
determined by their different geological and biological
make-ups, the affect of currents, tides and temperatures,
and interconnection or isolation. Some 17 different demersal
bioregions were identified in a recent CSIRO study, based
on the distribution of fish species around Australia (Interim
Marine Bioregionalisation for Australia, 1999). Much of
Australia’s diversity may be attributed to the climatic
changes that occurred during glacial periods, affecting sea-
levels and temperatures and causing populations to move
or adapt to altering conditions in different places.
Populations on the east and west coasts were able to move
latitudinally when temperatures changed thus presenting
little or no pressure for change. However, on the south coast,
populations became isolated on the eastern or western sides
or were split allowing for speciation over time. Seahorses
living in estuaries were most likely to have remained in
their original areas, as their offspring are not often dispersed
by ocean currents, and were likely to have adapted to
changing temperatures and the other conditions. The effect
of these environmental changes are probably reflected in
the differences in the length of snout, maximum adult size
and colour patterns that currently exist among populations.
Reproduction in seahorses, with males brooding the eggs
and producing highly advanced young, reinforces
localisation. Of the few species with pelagic young, some
are estuarine and rather restricted in distribution, and others
coastal and wide-ranging within a region. Australian species
are variously distributed along the continental coast, with
more than one species usually found in any given area, but
usually in different habitats. Despite this, no seahorses have
been recorded from the southern-most part of Western
Australia.
The unique morphology of seahorses is known even to
the non-specialist. Likewise, their reproductive strategy,
shared with other members of the syngnathid family, is
familiar to most. Rather than males taking sperm to the eggs
of females, as occurs in the vast majority of animals, female
syngnathids take their eggs to the source of sperm, placing
the eggs on or in the male’s body for incubation. In seahorses
and a few pipefishes, this involves a fully enclosed pouch
in which the eggs are brooded, the male truly becoming
pregnant. After three to four weeks a male seahorse gives
birth to between 50 to 400 fully formed seahorses,
depending on species (personal observation of aquarium-
reared H. breviceps, H. whitei and H. bleekeri ). Some
temperate species have only a few large broods in summer
months, but tropical species may reproduce more often, and
breeding appears to coincide with the wet season or phases
of moon which produce large tides and food for offspring.
Most species go through changes with growth, usually
reflected in length of their snout, spines or tubercles, and
colour. Males develop a brood-pouch well before maturity,
often when only a few months old, and usually start breeding
before becoming fully grown. Pelagic young of H.
abdominalis, H. bleekeri, H. histrix, H. breviceps, H.
tuberculatus and H. bargibanti have relatively long snouts.
The long snout eventually becomes very short in the adults
of H. abdominalis, H. breviceps, H. tuberculatus and H.
bargibanti, but in H. bleekeri and H. histrix the snout
remains long and may even lengthen further with age. Many
species have young that settle at birth and they have a
relatively short snout which usually becomes longer with
growth. The length of the snout relates to the form of prey
targeting: different forces of suction are required for free
swimming or crawling prey. A small seahorse feeding on
plankton needs a long snout for success, whilst one clinging
to the substrate can target crawling prey, and a short snout
is more practical in negotiating complex, three-dimensional
habitat, such as Sargassum or sponges. The general
appearance of the seahorses in the environment is one of
effective camouflage. A species that lives on vegetated or
sponge reefs can adapt to the environment by changing
colour or growing appendages. These adaptations are slow
processes that occur when individuals move to different
surroundings. This may be controlled through the food-
chain, as most feed on small Crustacea that in turn feed on
the weeds or sponges which dominate most seahorse
habitats. Some species can change colours rapidly, but
usually these are temporary and relate to courtship. Although
colour is highly variable within most species, there are
diagnostic features in patterns and the range of colours, that
can be useful in distinguishing between even closely related
species. Species with long spines live on more open
substrates and are often brightly coloured to match the soft-
corals or sponges to which they cling. They are not found
in dense weed or algae habitats, as the spines would
probably foul and be a hindrance to movement. A good
example is the rather smooth Western Australian H.
elongatus, which apart from its reduced spines is virtually
identical to the more tropical spiny H. angustus. I speculate
that H. angustus and H. elongatus shared a spiny ancestral
form and the spines reduced in the southern population as
reefs became more algae dominant during the cooling
periods.
Although seahorses have been of little interest to western
society, except as curiosities or aquarium pets, dried
seahorses have been used for medicinal purposes in the
Orient as part of Traditional Chinese Medicine (TCM) for
centuries. Whilst trade in dried seahorses uses millions of
specimens it involves only a few species worldwide. Threats
to the survival of any of the species collected in the wild
are, in my opinion, generally overstated or sensationalised.
Whilst many have a limited geographical range, commercial
collecting is very localised with many areas remaining
untouched. This includes Indonesia and Philippines, where
most collecting takes place (Lourie et al., 1999). Collecting
specimens from the wild in Australia is not economical,
except for supporting a small aquarium trade that is closely
monitored. Since aquaculture is much more economical it
is unlikely that catch rates will increase. Threats are more
likely to come from the rapid increase of human populations
which cause environmental pollution and habitat destruction
in estuaries and coastal waters. Seahorses can be used as
indicator species of ecosystem health and, because they are
habitat-specific, can tell us much about how habitats differ
or relate to each other. As shown here, species are more
localised than previously thought, and preserving habitats
is the most important factor in protecting seahorses for the
future.
As information on the distribution of some Australian
species is very limited, because they dwell at greater depths,
distributions provided may only reflect a small portion of
their true range. The author welcomes correspondence on
Hippocampus species (syngnathiformes@zoonetics.com)
which may contribute to our understanding of their biology.
Kuiter: Australian Hippocampus revision 295
Materials and methods
Descriptions of species are primarily based on specimens
housed in collections of Australian institutions. Those taken
from previous works are so acknowledged.
Institutions are abbreviated as follows:
AMS Australian Museum, Sydney
CSIRO CSIRO Marine Laboratories, Hobart
MNHN Museum national d’Histoire naturelle, Paris.
NMV Museum Victoria, Melbourne
NTM Museum and Art Galleries of the Northern
Territory, Darwin
QM Queensland Museum, Brisbane
SAMA South Australian Museum, Adelaide
WAM Western Australian Museum, Perth
ZMB Universitat Humboldt, Museum fur Naturkunde,
Berlin
Observations and photographs of species taken while
diving, in aquariums, and based on freshly collected
specimens were important in determining some of the
species. Despite this, most photographs presented were
taken from preserved material. A simplified terminology
and the avoidance of abbreviations has been employed here
for two reasons: 1, seahorses have unusual characters that
many workers may not recognise, and 2, the unusual vertical
posture and shape of seahorses may cause confusion when
applying the terms “anteriorly”, “posteriorly”, “dorsally”
and “ventrally” (see Fig. 1). Terminology used in the text
are depicted in Figs. 1 and 2. Measurements are kept to the
natural posture of the fish as shown in Fig. 3. Ridge and
ring definitions follow Dawson, 1985.
Figures of type-material, male, female and juvenile stages
are provided for each species where material was available
for photography.
A pictorial key is provided for species of Hippocampus
known to occur in Australia (Fig. 4). All species presented
on a page of the key are listed together with their meristic
values at the bottom of the page in the caption.
Morphology (Figs. 1, 2). The body of seahorses and other
syngnathids is covered by series of bony segments each
having a raised centre or edges, they together form rings
across the body and ridges that run the length of the body.
Where rings and ridges cross, the junctions normally rise
and form tubercles or spines. Tubercles may become large
in some species and have rugose tops. Spines may have
rugose edges, and may be blunt or sharp. The absence,
presence or size of spines or tubercles on the head and body,
are often diagnostic features, but these may vary between
different stages of a species or between the sexes. In general,
juveniles are more spiny than adults and females are more
spiny then males.
Trunk and tail ridges. Seahorses have 7 trunk ridges, 1
mid ventrally and 3 laterally on each side, and 4 tail ridges,
2 laterally on each side. Principal ridges (Fig. 3) are very
similar among species. All trunk ridges originate on the first
body ring but each one ends on different rings coinciding
with the origin of the tail: superior ridges usually continue
on to the first one or two tail rings, but the lateral ridge
usually ends on the penultimate trunk ring.
Trunk rings. The first trunk ring starts where the neck-
ridge divides laterally into the superior trunk ridges (see
Fig. 1) and the last trunk ring is the ring on which the seven-
■jl ::;ih i i :j
vc rrl rally
dersally
|KlhbHHiHy
I j-hiI ili’-i+idkTa
7
1 k'r'a
i.ilv
L
| aorty dniiiFj ii
F^crrrl Y
Figure 1. General morphology.
| writ sprk?]
l^arebdiirwl tfc|
I-JU..H un i-
I: an
| .rliwuinj*;'
tlHIII
■ii i r*: n-:L|i-
Figure 2. Spine, ring and ridge terminology.
296 Records of the Australian Museum (2001) Vol. 53
Figure 3. Measurements.
sided body changes to the four-sided tail. The anus is
situated ventrally on the last trunk ring.
Subdorsal rings. The dorsal fin is usually based over a
number of trunk rings and 1 or 2 tail rings. The rings are
termed as subdorsal and presented as 3+1, when the base
reaches over three trunk rings and one tail ring. In variable
species the origin may range in front or behind a certain
trunk ring and the rings are presented as a range (e.g., 1-2,
2-3).
Tail rings. The first tail ring follows the last trunk ring.
The superior tail ridge is usually preceded by a spine on the
last trunk ring below the superior trunk ridge running along
the base of the dorsal fin onto the first tail ring. The last
few tail rings are often difficult to count, especially when
covered with thick skin, and are best judged by the spacing
of the preceding rings, assuming the same rate of
proportional reduction posteriorly. The posterior part of the
tail is usually rolled tightly near the tip, where rings are the
smallest. The resulting skin folds on the inside are easily
mistaken for rings and may cause small errors in counts. In
most cases the tail-ring count is not useful in differentiating
closely related species.
Tubercles. Tubercles are raised nodes at the intersections
of rings and ridges. They are usually shaped like a small
pimple or with broad, rugose tips. Taller tubercles with
rounded tops resemble low, blunt spines. These are often
present as spines in juveniles, but usually become blunt and
relatively smaller with growth.
Spines. Spines are essentially extended and better developed
tubercles. They range in size from small to moderate in
length, in shape from thorn-like to long and slender, and
from blunt to pointed at the tip. Tips or edges are sometimes
rugose. Males are usually less spiny than females and
juveniles are more spiny than adults. In most cases males
have reduced spines on the superior trunk rings, but these
can be long in females. In long-spined species, deformities
are common with doubled or even tripled spines angled
randomly. Occasionally spines that may be of diagnostic
value are broken, regrown or missing due to damage from
predators or the methods of capture.
Head spines. The lower shoulder-ring spine is termed
“cheek-spine” by some authors, but true cheek spines are
located below the eye. Because of this confusion the term
is omitted here and “spines below eye” is employed in
referring to spines in the cheek region.
Subdorsal spines. The posterior and anterior parts of the
superior trunk and tail ridges, respectively, overlap each
other below, and for the length of the dorsal-fin base; spines
on the intersection of superior ridges and ring joints (nodes)
in this region are termed “subdorsal spines”. The superior
trunk ridge ends below the dorsal fin and often bears several
enlarged spines on the subdorsal rings, usually one per ring.
The superior tail ridge arises on the trunk, below the superior
trunk ridge, and may or may not bear spines on each ring
below the dorsal-fin base, depending on the species. Usually,
the intersection of the tail ridge with the second subdorsal
ring has an enlarged spine, and intersections of the tail ridge
with the first or third subdorsal rings occasionally bear
smaller spines. When species have a long dorsal-fin base,
there may be one or two additional intersections posteriorly.
The subdorsal-spine counts are presented as follows:
number of superior trunk ridge spines separated by a
diagonal (/) from the superior tail ridge spine formula. The
number of superior trunk-ridge subdorsal-spines is
represented by a value, and a variable number by a range
of values (e.g., 3-4). The superior tail-ridge subdorsal-spine
formula comprises the values for presence, absence and
position of superior tail-ridge spines per subdorsal
intersection of the superior tail ridge. The presence of a
spine is represented by 1, the absence of a spine by 0, a
small or poorly developed spine by 0.5, and variable
presence is indicated by a range (e.g., 0-1 or 0-0.5). Values
are separated by commas (e.g., 0,1,0). The presence of
successive spines is represented by separate numbers (e.g.,
1,1) and successive absence by naughts (e.g., 0,0). Atypical
count would be shown as 0,1,0, or 0,1,1 (no spine on 1st
node in both examples, only one spine on the 2nd node in
former, and spines on 2nd and 3rd nodes in latter), but in
variable species may appear as 0,1,0-1 (3rd node without
or with spine) or 0-0.5,1,0-1,1 (1st node without or with
small or poorly developed spine; 3rd node without or with
spine; the last value representing 4th node with spine).
Kuiter: Australian Hippocampus revision 297
Fin rays. Fin-ray counts provided represent the number of
rays having a single base. Rays split at the base are rare in
seahorses and were counted as one. In some specimens,
fins were observed that may have been damaged at an early
stage in life and may have re-grown in an aberrant way so
that they included additional rays or missing parts. An
attempts was made to adjust counts when abnormalities were
recognised. They usually manifested themselves as an
irregular spacing between rays, with several rays crowded
at the base, or by the presence of an unusually wide space
between the rays. Differences in pectoral-fin counts between
the left and right sides are common and variations by one
or even two rays were regarded as normal.
Measurements (Fig. 3). Height. A straight-line measure¬
ment from upper surface (ignoring spines) of first trunk
ring, to tip of tail as shown in Fig. 3. Coiled tails were either
straightened or replicated by bending a thin metal wire into
the shape of the tail, and straightening it for measurement.
Trunk length. Measured as for “height” from upper surface
of first trunk ring, vertically downward to last trunk ring,
employing tail-ridge sub-dorsal spine (usually present and
enlarged) at the last trunk ring as a termination point. Trunk
depth. Measured perpendicular to trunk axis, its maximum
dorsoventral depth. It is not shown in Fig. 3, as this
measurement is at different sections of the trunk, depending
on species, sex or age. Head length. Distance from tip of
snout to gill opening. Postorbital length. Distance from rear
of orbit to gill opening. Snout length. Distance from tip of
snout to front of orbit. Snout depth. Minimal depth of snout.
Total length. Combined length of head and height, measured
by bending a metal wire in a similar fashion to that described
for “Height”. Though not recorded for this study, values
are occasionally reported when quoting published data
(abbreviated as TL). Standard length. Length of fish from
tip of snout to posterior edge of hypural (joint between
caudal skeleton and caudal fin). This is a standard
measurement for the length of fishes with caudal fins
(abbreviated as SL) but, except for Lourie et al. (1999), is
normally not used for seahorses.
Although standard ichthyological terminology has been
employed wherever practical, because of the extraordinary
morphology in these fishes, additional terms as “neck” or
“shoulder” are employed. Whitley (1958) used “shoulder¬
ring” for the ridge-like feature that runs in front of the
pectoral-fin base and the term is retained here, along with
several of his other descriptive terms. Because of the head
position, the direction “anteriorly” and “posteriorly” (Fig.
1) can be confusing for seahorses. The angle of the head to
the body can vary greatly among species. In some species
the heads of adults are strongly angled down, whilst the
heads of pelagic young are almost in line with the body.
When used in reference to the head, anterior is directed
towards the tip of the snout, while posterior is toward the
gill-opening. For the rest of the body, posterior is orientated
toward the tip of the tail, while anterior is directed toward the
back of the head. The term “nose” refers to the nasal area.
Material examined. Since most specimens were registered
under different names, only those specimens identified as
the actual species are listed in the descriptions. Specimens
examined from outside Australia, representing species not
included in this revision, are not listed and only remarked
on when relevant.
Genus Hippocampus Rafinesque
Hippocampus Rafinesque, 1810: 18. Masculine. Type species
Syngnathus heptagonus Rafinesque, 1810 ( =Syngnathus
hippocampus Linnaeus 1758).
Macleayina Fowler, 1907: 426. Feminine. Type species
Hippocampus bleekeri Fowler, 1907.
Farlapiscis Whitley, 1931: 313. Masculine. Type species
Hippocampus breviceps Peters, 1869.
Jamsus Ginsburg, 1937: 584. Masculine. As subgenus of
Hippocampus. Type species Hippocampus regulus Ginsburg, 1937.
Hippohystrix Whitley, 1940: 414. Feminine. Type species
Hippocampus spinosissimus Weber, 1913.
Diagnosis. Body encased in ring-like rigid plates; body
usually maintained in vertical posture with head bent
forwards and downwards in sharp angle, usually forming
an angle of less than 90° to body in adults; snout tubular,
with small mouth at tip, jaws lacking teeth; tail prehensile;
males with brood pouch below anterior part of tail; trunk
heptagonal in cross section, each side with superior ridge
dorsally, lateral ridge along upper side, and inferior ridge along
lower side, as well as single mid-ventral ridge (sometimes
termed “keel”); tail four-sided with two superior ridges dorsally,
and two inferior ridges ventrally; trunk usually with 11 rings,
but occasionally with 12 or 13 rings; tail usually with 30 or
more rings; dorsal and pectoral fins moderately large, usually
with about 10-20 rays; anal fin small and occasionally
absent in adults; ventral and caudal fins absent.
Remarks. Only a single genus is presently recognised for
this highly diverse group of fishes. The absence of a clear
separation between any of the various forms, based on
external features, makes it difficult to justify the recognition
of one or more of the current synonyms at the generic level.
Although there appear to be distinct groups, such as
“smooth” and “spiny” forms, additional characters need to
be investigated prior to any taxonomic splitting of the genus,
if it is indeed warranted. Two of the species treated here are
only known from single specimens in Australian waters.
Hippocampus jugumus is a very distinctive species from
Lord Howe I. but no additional material is known. A single
specimen of H. kampylotrachelos was found on Ashmore
reef, which is well away from the Australian mainland. This
species is known to occur in Indonesian waters from Java
to the Timor Sea. As H. semispinosus was reported only as
coming from the “Timor Sea”, it is unclear whether the
paratype came from Indonesian or Australian waters. The tiny
H. spinosissimus is known only from 2 specimens trawled at a
depth of 70 m depth in Sapeh Strait which runs east of Komodo,
Indonesia. Occurrence off this species in Australian waters
is unconfirmed. Since the name is commonly used for spiny
species in Australia, its description, translated from the
original German publication, is included. It may be unique
to the Komodo region as a few other species of fish (e.g.,
Apogon komodoensis Allen, 1998) appear to be endemics
there. The occurrence of H. histrix in Australian waters is
highly likely even though no specimens were found in the
collections of the Australian institutions. This species occurs
from Japan to Bali, Indonesia, along the Wallace’s Line,
and eastward along the Papuan New Guinea coast, ranging
to at least New Caledonia in the Coral Sea. As this is one of
the most widespread species of the genus, expatriates
transported by current during the pelagic stage can be
expected on the north coast or as inhabitants of the inner
parts of the Great Barrier Reef.
298 Records of the Australian Museum (2001) Vol. 53
//. bargibanti
-► !
to 25 mm
► 1 (p. 299)
II. jugumus
-►
to 44 mm
2 (p. 301)
Figure 4. Key to Australian Hippocampus species. Hippocampus minotaur (p. 304), D7, Pll, R8+41, southeastern
Australia; H. jugumus (p. 306), D20, PI6, R12+37, known only from a single Lord Howe I. specimen, height 44
mm; H. bargibanti (p. 305), D13-15, P10, R11-12+31-34, Qld and west Pacific.
Kuiter: Australian Hippocampus revision 299
1 (p. 298)
H upper shoulder-ring spine near gill opening |
z.
I
upper shoulder-ring spine near fin base
3 (p. 302)
often small or
as low ridge
moderate
spine or
tubercle
coronet low or
poorly developed
nape spine \
directed upward~
coronet very low
lower shoulder-ring
spine always single
nose profde low,
curving into snout
H. kampylotrachelos
nape spine
directed forward
coronet small to
moderate, usual¬
ly well defined
/ '
| dorsal rays 19 |
i
k
i
| dorsal rays 21—24
1
r
snout short, to 40%
of head, or equal to
postorbital length
to 22 cm
nose profile straight
rising c. 45° from snout
in
_ H.dahli
\ snout long, to 50% of head [
P
lower shoulder-ring
spine (or tubercle)
single or double
nose profile slightly up,
rising c. 45° from snout
to 12 cm
fy
snout spotted (fresh)
trunk longer than
twice its depth (adult)
//. planifrons
to 10 cm
nose profile straight
rising c. 45° from snout
snout mostly plain
trunk shorter than
twice its depth (adult)
4 (p. 300)
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus kampylotrachelos (p. 307), D19,
P16, Rll+39, Indonesia and northwestern Australia; H. dahli (p. 308), D21-22, P17-18, R11+37-40, Qld and NT;
H. planifrons (p. 310), D23-24, P18-19, Rll+37-38, WA; H. biocellatus (p. 311), D22-23, P16-18, Rll+36-38,
Shark Bay, WA.
300 Records of the Australian Museum (2001) Vol. 53
T
4 (p. 299)
coronet-apex with¬
out posterior spines
D20—22; P13—15
coronet
poster
-apex with
ior spines
D 16—
19; P16—19
1. Coronet rounded, head and body
semi-fleshy, no spines visible.
2. Low tubercles below dorsal fin
and anteriorly on superior tail ridge.
//. breviceps
to 10 cm
*3
1. Coronet square to rounded, head
and body bony, spines visible.
2. Large tubercles below dorsal fin
and anteriorly on superior tail ridge.
II. tuberculatus
to 8 cm
superior ridge
maybe slightly
spiny in small
juveniles
snout-length equals
postorbital length,
much less than 1/2
head length
tail not spiny
spines maybe
reduced to low
tubercles in
large adults
Coronet back-swept with
5 diverging small spines,
or 3 posterior spines
■D 17—18; P 16
Lower shoulder-ring-spine
single
D 18—19; P 18—19
Lower shoulder-ring spines as
double blunt, rounded tubercles
snout-length about 1/2 head length or longer
tail usually spiny, sometimes as tubercles in large adults
► 5(p. 301)
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus breviceps (p. 312), D21-22, P13-14,
R11+38-42, SA, Vic., Tas.; H. tuberculatus (p. 313), D20-21, P14-15, Rll+36-37, WA; H. taeniopterus (p. 314),
D17-18, P16, R11+34-35, NT, Qld and west Pacific; H. tristis (p. 316), D18-19, P18-19, Rll+35-37, southern
Qld, northern NSW and Lord Howe I.
Kuiter: Australian Hippocampus revision
//. semispinosus
to 14 cm
to 30 cm
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus alatus (p. 317), D15-
18, P16-18, Rll+34-36, northern Australia; H. queenslandicus (p. 319), D17-18, P16-19, Rll+35-
36, Qld; H. semispinosus (p. 320), D18, P16-17, R11+35-36, southern Indonesia; H. abdominalis (p.
322), D25-28, P15-16, R12-13+44-45, NSW and NZ; H. bleekeri (p. 323), D27-30, P14-16, R12-
13+44-48, SA, Vic. and Tas.
II. bleekeri
H. abdominalis
to 18 cm
filaments usually
present in adults
5 (p. 300)
ribbon-like filaments
snout-length ~l/2 head length,
and snout-depth 4x in its length
trunk ridges with enlarged broad-
based blunt spines on rings 1, 4, 7
and below dorsal fin, others as low
tubercles, ventral ridge smooth.
laterally directed,
wing-like spines
nape and lateral head spines with ribbon-like filaments H. alatus
snout-length ~l/2 head length,
and snout-depth 3.5x in its length
trunk ridges with broad-based small
spines on nearly all intersections,
some enlarged. Juveniles more
spiny, enlarged spines on rings 1, 4,
7 and below dorsal fin.
II. queenslandicus
to 12 cm
spines on superior trunk
and tail ridges directed
laterally, slightly dorsally
spines on superior trunk
and tail ridges directed
dorsally, slightly laterally
to 12 cm
snout longer than 1/2 head length,
and snout-depth 5x in its length
tmnk ridges with few broad-based
blunt spines on rings 1, 4, 7 and
below dorsal fin, others as low
tubercles, ventral ridge smooth.
occasional fila¬
ments, usually
absent in adults
2 (p. 298)
T
snout 2.0—2.5 in head (juv—adult respect.)
snout 2.7—3.2 in head (juv—adult respect.)
head 1.25—1.4 in tmnk
head 1.3—1.6 in tmnk
302 Records of the Australian Museum (2001) Vol. 53
3 (p. 299)
no distinct transverse stripes | — H zebra-like pattern of alternating white and dark stripes | -
H. zebra
to 8 cm
1. spines small & blunt
2. neck spines absent
3. two sub-equal spines
//. montebelloensis
-►
to 8 cm
1 spines on front of coronet-base [
1. spines moderate & sharp
2. neck spines present, but
low in large males
3. single spine, 2nd in front
usually small or absent.
II. whitei
to 10 cm
II. procerus
-►
to 11 cm
- |nose-profile straight without spine| - ftail rings 37| —
nose-profile
angular or
spiny
tail rings
-I no spines on front of coronet-base [■
6 (p. 303)
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus zebra (p. 325), D17-18, P15-16,
R11+37-39, Qld; H. montebelloensis (p. 326), D18-19, P15-16, Rll+37, WA; H. whitei (p. 327), D16-17, P15-
17, R11+33-34, NSW; H. procerus (p. 328), D17-19, P16-18, Rll+34-35, Qld.
6 (p. 302)
Kuiter: Australian Hippocampus revision
1. coronet with rounded flanges between tips
H. elongatus
2. trunk mainly with tubercles
to 20 cm
K
H. spinosissimus
/IV, * ,
to 7 cm
snout shorter than
1/2 head length
1. coronet with sharp spines
2. trunk spiny _
1. length ~ eye-diameter
2. long & slender _
'1
snout longer than
1/2 head length
1. shorter than eye-diameter
2. short & angular
IP
nape spine
directed forward,
placed away
from coronet
/
//. multispinus 1 f ■
-► '
to 14 cm
* l _ ■- -^1
i _
shoulder-ring
spine double
nape spine
directed upward
and placed close
to coronet
» to 15 cm
shoulder-ring spine single
vJr jf *
II. histrix
7 (p. 30 z
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus elongatus (p. 329), D18, P17-18,
R11+33-34, southern WA; H. spinosissimus (p. 330), D17, P15, Rll+34, Indonesia; H. multispinus (p. 331), D18,
P16-18, Rll+30-35, northern Australia; H. histrix (p. 333), D17-19, P17-18, Rll+33-34, west Pacific.
304 Records of the Australian Museum (2001) Vol. 53
W 7 (p. 303)
head length about equal to trunk length}
moderate lateral-
ridge-spines on
3rd & 5th ring
^^►- phcad length to about 80% of trunk length
//. grandiceps
-►
to 11 cm
_
| no lateral-ridge-spines on 3rd & 5th ring
' r
II. hendriki
to 10.5 cm
H. angustus
to 16 cm
Figure 4 (continued). Key to Australian Hippocampus species. Hippocampus hendriki (p. 333), D17-18, P16-17,
Rll+34, northern Qld; H. angustus (p. 335), D18-19, P15-20, Rll+31-32, WA; H. grandiceps (p. 336), D18,
P17-18, R11+32-33, Gulf of Carpentaria, Qld.
Hippocampus minotaur
Bullneck Seahorse
Fig. 5
Hippocampus minotaur Gomon, 1997, off Eden, New South Wales,
Australia.
Diagnosis (after Gomon, 1997). Dorsal-fin rays 7; pectoral-
fin rays 11; anal-fin rays 4; no exaggerated constriction
dividing head and body; head and body extremely fleshy
without recognisable bony segments, spines or other
ornamentation; ventral trunk ridges undeveloped. Fin rays:
dorsal 7; pectoral 11; anal 4. Rings: subdorsal obscured;
trunk 8; tail 41. Spines or tubercles: head and body fleshy
without recognisable bony segments, spines or other
ornamentation. Lateral line: obscured. Coronet: appearing
as a smooth raised hump. Colour in life: unknown. Colour
in alcohol: mostly cream, speckled with tiny brown dots
having cream centres or plain brownish overall.
Kuiter: Australian Hippocampus revision 305
Figure 5. Hippocampus minotaur. A & B, holotype, 6, NMV A192, Eden, NSW. C, paratype, $, AMS IA3509,
coast of NSW. D, paratype, juvenile, NMV A14161, Cape Patterson, Vic. A—pouch modified digitally; B & C—
after Gomon.
Measurements (after Gomon, 1997). The holotype
(paratype measurements in brackets) has a total length of
48.7 mm (19.2-52.6). Head length 16.4% (16.7-21.3) TL;
snout very short, 29.3% (28.0-31.1) HL; trunk short, 31.0%
(31.3-36.4) TL.
Distribution (Fig. 6). Southern New South Wales to the
Bass Strait region in a depth range from about 70 to 110 m.
Only known from the type material.
Remarks. Readily distinguished from other congeners by
the short-based dorsal fin with just 7 rays and features listed
in the diagnosis. Specimens reported by Gomon (1997) as
being taken in environmental monitoring surveys off
Wollongong appear to be lost. Although no details of the
specimens are known, they are more likely to represent an
undescribed species of pipehorse Idiotropiscis sp. The latter
Figure 6. Hippocampus minotaur. Collection sites of specimens
examined.
is a small seahorse-like species that ranges from the Sydney
region to Jervis Bay (Kuiter, ms in preparation).
Material examined. Holotype: NMV A192, 9, 48.7 mm TL,
off Eden, New South Wales, depth 35-40 fm, Danish seine trawl,
R. Slack-Smith, 30 Dec 1960. Paratypes: AMS IA3509, 9,42.2
mm TL, NSW coast, trawled, K. Moller, registered 27 Jun 1927.
AMS IA3560, $, 52.6 mm TL, 8 mi E Eden, NSW, depth 50-60
fm, H. Howell, registered 7 Oct 1927. NMV A14161, juvenile,
19.2 mm TL, Bass Strait, 38 km SW Cape Patterson, 38°56.4'S
145°16.6'E, 70 m, fine sandy bottom, RV Tangaroa, R. Wilson,
12 Nov 1981.
Hippocampus bargibanti
Pygmy Seahorse
Fig. 7
Hippocampus bargibanti Whitley, 1970. New Caledonia, Noumea.
Diagnosis (after Gomon, 1997). Dorsal-fin rays 13-15;
pectoral-fin rays 10; anal fin absent in adults; very weak
constriction separating head and body; head and body
extremely fleshy without recognisable bony segments, body
ornamentation in the form of prominent bulbous tubercles
in adults; ventral trunk ridges poorly developed. Fin rays 1 :
dorsal 13-15; pectoral 10; anal fin absent in adults. Rings 1 :
subdorsal obscured; trunk 11-12; tail 31-34. Spines or
tubercles: head and body extremely fleshy without
recognisable bony rings, spines or tubercles not visible and
become overgrown with soft tissue, forming large bulbous
or wart-like lumps in adults. Lateral line: obscured. Coronet:
hump-like. Colour in life: pale grey, bluish grey, yellow or
pink with yellow, orange to red lumps; body striated in
adults. Colour in alcohol: mostly cream.
1 Counts after Gomon (1997, fig. 4), and Lourie et al. (1999).
306 Records of the Australian Museum (2001) Vol. 53
Figure 7. Hippocampus bargibanti. A, lectotype, 2, AMS 115418-002, Noumea, New Caledonia (after Gomon,
1997). B, $, Sulawesi, Indonesia. C, 8 , Sulawesi, Indonesia. D, maturing 8, showing striations on body, Sulawesi,
Indonesia, photo by Tony Wu. E juvenile, probably young 2, showing wart-like development on the principal
points of the trunk and head, Sulawesi, Indonesia, photo by Roger C. Steene.
Figure 8. Hippocampus bargibanti. Distribution in Australia,
based on localities of specimens photographed underwater.
Measurements. The five specimens in the Australian
museum collection range in size from 19.5-24.2 mm TL.
The largest specimens seen in the wild were estimated as
20 mm in height and about 30 mm in total length.
Distribution (Fig. 8). Since its discovery in Noumea, diver-
photographers have found Hippocampus bargibanti in many
places between Australia and Japan. It is now known from
Bali, and both north and south Sulawesi in Indonesia,
Ogasawara Is of southern Japan, Papua New Guinea and
Solomon Is. It was first discovered in Australia by Alan
Power in the Capricorn region, soon after it was described.
Since then, it has been photographed in several locations
on the northern Great Barrier Reef. Hippocampus bargibanti
is usually found in depths between 30 and 60 m, but occurs
as shallow as 10 m in Indonesia and the Solomon Is.
Remarks. This is a remarkable, small species that has
adapted to live on gorgonian corals, on which it feeds and
spends its adult life. Young are pelagic and may disperse
over great distances. Post pelagic young settle on a variety
of differently coloured gorgonian species at various depths,
quickly adopting a similar colour and changing shape. In
shallow water where gorgonians are diverse, young H.
bargibanti vary in colour depending on the species to which
they attach, and may differ completely from adults that seem
to prefer the gorgonian Muricella, usually occurring in
depths of 20 m or more. Those settling in the shallower
depths can range from yellow to brown or red, and have
warts that match retracted gorgonian polyps in size. Fish
appear to adapt to their hosts by eating their hosts’ tissue.
These seahorses become perfectly camouflaged, taking on
the colour and developing the polyp-like lumps of their host,
in addition its skin reacts by growing, as if following an
encoded message in the host flesh that they ingest. Young
settling on the deep-water gorgonian Muricella have the
same colours as most adults, and those in the shallows are
likely to move to deeper water and change colours after
having first settled on different coloured gorgonians. The
occasional young that does not match its host in colour and
shape, may represent an individual that only recently moved
to a new host. Gomon (1997) provided a detailed description
of the species.
Type material. Lectotype: AMS 115418-002, 2 ,20.9 mm
TL, Noumea, New Caledonia, depth 30 m, collected with
gorgonian coral Muricella sp., Georges Bargibant, Jul 1969.
Paralectotype: AMS 115418-001, 2, 19.5 mm TL,
Noumea, New Caledonia, depth 30 m, collected with
gorgonian coral Muricella sp., Georges Bargibant, Jul 1969.
Hippocampus jugumus n.sp.
Collared Seahorse
Fig. 9
Hippocampus histrix (non Kaup) Whitley & Allan, 1958: fig. 6-3.
Hippocampus sp. B, Allen et al., 1976.
Type material. Holotype: AMS IA2424, height 44 mm,
$(?), Lord Howe Island, off New South Wales, 31°31’S
159°05'E, collected by A. McCulloch, 1925.
Diagnosis. Dorsal-fin rays 20; pectoral-fin rays 16; trunk
rings 12; tail rings 37; subdorsal spines 4/0,1,1,1,1; nose
spine moderately large, height about equal to pupil diameter;
double spine above eyes moderately long, slightly longer
than pupil diameter; lateral head spine double and large;
coronet slightly raised; gill openings extending upwards,
Kuiter: Australian Hippocampus revision 307
Figure 9. Hippocampus jugumus. Holotype, AMS IA2424. A, head; B, head and trunk; and C, entire specimen.
almost joined at neck ridge; shoulder rings confluent over
neck-ridge forming continuous collar.
Description. Head small, c. 50% of trunk length; snout
short, length about equal to postorbital length; dorsal fin
with 20 rays, its base over 3 trunk and 2 tail rings; pectoral-
fin rays 16; trunk rings 12; tail rings 37; long double spine
above eyes; slightly larger double lateral head spine,
pointing outwards; nape spine of moderate length,
perpendicular to forehead; spine behind eye of moderate
length; 2 separate spines below eye; 2 moderately large
spines on shoulder ring, upper spine at level of last pectoral-
fin ray, and lower spine at ventral extent of ring, directed
perpendicular from the curve; coronet slightly raised with
6-7 short spines diverging in various directions; neck ridge
with small spine immediately behind shoulder ring and one
at posterior end; superior trunk ridge with enlarged blunt
spines on 1st, 4th, 7th and 11—12th rings; lateral trunk ridge
with small spine on each trunk ring apart from the first;
inferior trunk ridge similarly with small spine on each trunk
ring apart from the first; ventral trunk spines small, directed
downwards, and mainly on last few rings; subdorsal spines
4/0.4; superior tail ridge spines well developed anteriorly,
except on first ring, gradually reducing in size, with enlarged
spines on 5th and 9th rings bearing dermal flaps; inferior
tail ridge continuous with inferior trunk ridge, spine on each
ring gradually reducing in size to 9th tail ring; lateral line
with pores visible to about 24th tail ring; gill-openings
extending upwards nearly to top of neck-ridge, the latter
narrowly separating the two. Lateral line : not detected.
Colour in life : unknown. Colour in alcohol : plain light
brown.
Distribution. Only known from a single specimen collected
at Lord Howe I.
Remarks. This species is named jugumus, derived from
the Latin jugum, meaning yoke, in reference to the yoke¬
like ridge formed by the shoulder rings joined over the nape.
Hippocampus jugumus is unusual in having a the high
number of trunk rings, a yoke-like shoulder ring, gill-
openings reaching upwards to the neck ridge, and double¬
spines on the head. The specimen may be immature, but no
other nominal species that might occur in the area could be
linked, and it does not resemble any other Indo-Pacific
species with regard to the full suite of diagnostic characters.
Nothing is known about the history of the holotype. The
fins are held stiffly against the body, suggesting it may have
been dried before being preserved. This is typical of beach-
washed specimens. Judging by the broad angle of the head
to the body it may be pelagic, a feature that is found in the
pelagic stages of H. abdominalis and H. bleekeri. It appears
to have no close relatives.
Hippocampus kampylotrachelos
Smooth Seahorse
Fig. 10
Hippocampus kampylotrachelos Bleeker, 1854b, Priaman,
Sumatra, Indonesia.
Hippocampus trimaculatus (non Leach) Lourie et al., 1999, in
part.
Diagnosis. Dorsal-fin rays 19; pectoral-fin rays 16; tail rings
39; dorsal profile of head evenly concave to tip of snout;
nose-profile without spine and gradually curving onto snout
with no obvious inflection; subdorsal spines on superior
trunk ridge not greatly enlarged; subdorsal spine on second
node of superior tail ridge enlarged, without rings on either
side. Fin rays: dorsal 19; pectoral 16; anal 4. Rings:
subdorsal 2+1; trunk 11; tail 39. Spines or tubercles:
subdorsal 3/0,1,0, lower spine enlarged and obvious (see
Fig. 10A); small spine above eye, directed back and
outwards; nose ridge slightly raised, spine absent; nape spine
small, directed upward; small spine behind eye; low double
spines below eye; upper shoulder-ring spine appearing as
slightly raised ridge beside gill-opening; tubercle of
moderate size on shoulder ring at level of last pectoral-fin
308 Records of the Australian Museum (2001) Vol. 53
Figure 10. Hippocampus kampylotrachelos. A, 9, NTM S12060-001, Ashmore Reef, Timor Sea. B, 6, Bali,
Indonesia, photo by Akira Ogawa.
ray; lower shoulder-ring spine of moderate size and
recurved; lateral head spine recurved; body tubercles of
small to moderate size along dorsal and lateral ridges, but
enlarged and pointed at regular intervals, especially at
dorsal-fin base and along superior tail ridges, gradually
becoming smaller on tail. Lateral line : not detected.
Coronet : low, directed backwards, with 5 blunt diverging
spines in a star at apex. Colour in life : dusky brownish grey
to almost black, covered with fine white and black spots
and scribbles, especially on head where these form a pattern
radiating from eyes, and lines along lower part of
operculum; dorsal fin with two longitudinal lines, one
marginally and one centrally. Colour in alcohol : nearly
black with traces of spots and lines on head in the pattern
described for living specimens.
Measurements. Specimen illustrated in Fig. 10 from Bali
was estimated at 22 cm in height. The NTM SI2060-001
specimen from Ashmore reef is about 96 mm in height but
is missing the end of the tail. Snout is about equal to half
length of head.
Distribution. Found in Sumatra (type locality), Bali and
Timor Sea. It occurs on muddy substrates and either in very
sheltered estuaries, or in deep coastal waters.
Remarks. Hippocampus kampylotrachelos is known from
only a few specimens, but is a distinctive species that is
unlikely to be confused with other congeners. The head has
an elongate look and is less angled to body compared with
most other seahorses. A specimen was found on land
amongst nesting birds at Ashmore Reef, Western Australia,
fits Bleeker’s description of H. kampylotrachelos perfectly,
and is the only record for Australia. It may have been a
casualty of trawling and was either picked-up by birds or
washed ashore. Counts and measurements include the
Ashmore Reef specimen, underwater photographs, and
Bleeker, 1983, pi. 449, fig. 2.
Material examined. NTM S12060-001, 9, height c. 96 mm,
Ashmore Reef, WA.
Hippocampus dahli
Low-crown Seahorse
Fig. 11
Hippocampus lenis de Vis, 1908. Nomen nudum.
Hippocampus dahli Ogilby, 1908. Noosa, Queensland.
Hippocampus planifrons (non Peters) Whitley, 1952.
Hippocampus planifrons (non Peters) Paxton et al., 1989.
Hippocampus planifrons (non Peters) Johnson, 1999.
Hippocampus trimaculatus (non Leach) Lourie et al., 1999: in
part.
Diagnosis. Dorsal-fin rays 21-22 (usually 21); pectoral-
fin rays 17-18 (usually 17); tail rings 37-40 (usually 39);
nose without spine, profile straight, raised from snout at
55-60° angle; coronet very low, front and lateral edges
rugose, posterior edge with 3 short backward directed
spines; single, recurved lower shoulder-ring spine; lateral
line distinct to 19th-23rd tail ring. Fin rays: dorsal 21-22
(usually 21); pectoral 17-18 (usually 17); anal 4. Rings:
subdorsal 2-3 + 1-2, (usually a total of 4); trunk 11; tail
37-40 (usually 39). Spines or tubercles: subdorsal 3/0,1,1
or 4/0,0,1,1; small recurving spine above eye; nape spine
small, directed upwards; shoulder-ring spines small,
uppermost low and directed upwards next to gill-opening,
central spine tubercular at level of last pectoral-fin ray,
lowermost as recurved spine; lateral head spine as low a
tubercle; body tubercles of small to moderate size along
dorsal and lateral ridges, few slightly enlarged, especially
along dorsal-fin base. Lateral line: a small pore just above
lateral ridge on trunk at each ring, continuing to about 19th-
23rd tail ring, intermittent over last few rings. Coronet: very
low, front and lateral edges rugose, posteriorly with 3 short
backward directed spines. Colour in life: females mostly
pale brown to almost black, usually with small black scribble
marks that sometimes form lined, zebra-like patterns. Males
usually dark brown nearly to black with black scribbles.
Colour in alcohol: similar to life colours, with pale colours
more brown and dark colours almost black.
Kuiter: Australian Hippocampus revision 309
Figure 11. Hippocampus dahli. A, holotype, 9, QM1788, Noosa, Qld. B, 9, QM114027, off Shorncliffe, Moreton
Bay, Qld. C, $, QM 114068, off Shorncliffe, Moreton Bay, Qld. D, $, AMS 119655-001, Gulf of Carpentaria, NT.
E, 9, NTM S10106-001, Darwin, NT. F, 9, NMV A21620, Townsville, Qld. G, S, NTM S13790-001, Beagle
Gulf, NT. H, (3, QM 131366, Fraser I., Qld.
Measurements. Largest specimen examined with a height
of 116 mm. Length of tail about 60% of height; trunk about
38% of height; head 78% of trunk. Trunk depth highly
variable, about 50-80% of head, depending on size and
sex. Juveniles and non-gravid females more slender than
males and gravid females. Length of snout is about half
length of head.
Distribution (Fig. 12). Occurs in coastal waters of
northeastern Australia from Moreton Bay to Darwin. Most
specimens in collections were trawled in shallow waters on
rubble substrates, with 21 m the greatest depth recorded.
Remarks. Ogilby described Hippocampus dahli on the basis
of a specimen from the Queensland Museum collection that
was labelled as H. lenis de Vis. However, no description of
H. lenis was ever published and the name is regarded as
merely a museum name. Since Whitley (1952) synonymised
the names H. dahli and Western Australian H. planifrons,
specimens with a low coronet were regarded as a single
species that was broadly distributed in tropical Australian
waters. This study has revealed that H. dahli is a valid
species restricted to northeastern Australia between about
Moreton Bay and Darwin. Most specimens in collections
Figure 12. Hippocampus dahli. Collection sites of specimens
examined.
were trawled in shallow coastal waters on rubble substrates.
In Moreton Bay, where this species was common,
populations appear to have declined significantly since the
1960s (Johnson, 1999, as H. planifrons ). Specimens from
310 Records of the Australian Museum (2001) Vol. 53
northern waters compared to those from southern
Queensland have fewer tail rings (usually 37-38 versus 39-
40) and a slightly different colouration. Northern males are
very dark with series of small white spots instead of dark
brown with black scribbles. These differences appear minor
and are regarded as clinal variations.
Material examined. QM1788, holotype, 9 , height 80 mm, beach,
Noosa, Qld, Mrs Birkbeck, 1912. QM 114027, 9, height 86 mm,
Shomcliffe, Qld, trawled in 5 fm, W Croft, Apr 1977. QM 114067,
6 , height 100 mm, Shomcliffe, Moreton Bay, Qld, trawled, W. Croft,
May 1977. QM 114068, 9 , height 93 mm, Shomcliffe, Moreton Bay,
Qld, trawled, W. Croft, May 1977. QM 126679, 9 , height 116 mm,
central Moreton Bay, Qld, trawled, H. Weng, 8 Aug 1974. QM 131366,
8, height 86 mm, Eurong beach, Fraser I., Qld, beach washed, R.
Hobson, 2 Feb 1999. NTM S10106-001, 9, height 70.5 mm, Van
Diemen Gulf, NT, NT Fisheries, 17 Jan 1978. NTM S10284-003, 9,
height 82.5 mm, off Mickett Ck, Shoal Bay, NT, 12°18'S 130°58'E,
depth 3 m, NT Fisheries, 20 Oct 1977. NTM SI 1664-001, 8, height
55.6 mm, Kahlin Beach, Darwin Harbour, NT, 12°27. l'S 130°49. l'E,
depth 3-10 m, R. Williams, 2 May 1985. NTM S13717-002, 9, height
81.5 mm, Haycock Reach, Darwin Harbour, NT, 12°36.8'S
130°56.4F, depth 4 m, R. Williams, 14 Jul 1993. NTM SI3790-001,
8 , height 82 mm, Bynoe Harbour, Beagle Gulf, NT, 12°41 'S 130°33E,
depth 9 m, R. Williams, 7 Oct 1993. AMS 115557-076, 9, height
100 mm, Gulf of Carpentaria, Qld, 17°24'S 140°09'E, CSIRO prawn
trawl, 27 Nov 1963. AMS 115864-001, 9, height 115 mm, SE North
Keppel I., Qld, 21 m, 23°05'S 154°45E, W. Ponder, 25 Jul 1969.
AMS 119655-001, 9, height 95 mm, Gulf of Carpentaria, Qld, 15°29'S
141°29'E, CSIRO, 8 Dec 1976. AMS 123676-002, juvenile, height
60 mm, and AMS 123677-001, juvenile, height 70 mm, both with
data: power station screens, Gladstone, Qld, 23°52'S 151°16'E, P.
Saenger, 2 Jan 1979. NMV A21620, 9, height 125 mm, off
Townsville, Qld, M.L. Home, Feb 1999.
Hippocampus planifrons
Flat-face Seahorse
Fig. 13
Hippocampus planifrons Peters, 1877. Shark Bay, Western Australia.
Hippocampus trimaculatus (non Leach) Lourie et al., 1999: in part.
Diagnosis. Dorsal-fin rays 23-24 (usually 23); pectoral-
fin rays 18-19; tail rings 37; subdorsal spines 4/0,0,1,1 or
5/0,0,0,1,1; single lower shoulder-ring spine, directed
outward; snout short, about equal to postorbital length with
dark lateral spots. Fin rays : dorsal 23-24 (usually 23);
pectoral 18-19; anal 4. Rings : subdorsal 3-4 + 1, dorsal
origin variable in position, from in front of, on, or behind
8th trunk ring; trunk 11; tail 37-38. Spines or tubercles :
subdorsal 4/0,0,1,1 or 5/0,0,0,1,1; small outwards curving
spine above eye; nape spine small, pointing up and forward;
shoulder ring spines small, uppermost spine low and
directed outward next to gill-opening, central spine at level
just below level of centre of pectoral-base, lowermost spine
directed outward; small lateral head spine; small spine
behind eye; body spines of small to moderate size along
dorsal and lateral ridges, few enlarged, most prominently
along dorsal-fin base. Lateral line : pores small but distinct,
situated just above lateral ridge on trunk at each ring,
reaching about 18th tail ring. Coronet : slightly raised, 2
small front spines, posteriorly 3 spines of moderate size
and middle spine enlarged and recurved. Colour in life :
females greenish brown with dark blotching over dorsal
region of trunk and tail. Snout with dark lateral spots. Colour
in alcohol : mainly brown with dark spots on snout.
Measurements. Largest specimen examined with height
of 70.2 mm. Length of trunk about 36% of height; length
Figure 14. Hippocampus planifrons. Collection sites of specimens
examined.
Figure 13. Hippocampus planifrons. A & B, 9, WAM P26674-006 (A, preserved; B, fresh), Shark Bay, WA. C,
juvenile, AMS IA4276, Broome, WA.
Kuiter: Australian Hippocampus revision 311
of head 68% of trunk. Trunk depth 60% of head length.
Length of snout about equal to postorbital length.
Distribution (Fig. 14). Only known from Shark Bay and
Broome. Records from the Northern Territory and
Queensland are misidentified Hippocampus dahli.
Remarks. A photograph of the holotype of Hippocampus
planifrons , provided by Sara Lourie, shows a specimen virtually
identical to Fig. 13A. The holotype is slightly smaller (head
length 16 mm, versus 17 mm), and the original description
indicates that it has of 23 dorsal-fin rays and 19 pectoral-fin
rays, which conforms with the specimens in Australian
museums. This species is similar to H. dahli , but the coronet is
not as low; the spines on the trunk and tail ridges are more
developed; is nape spine is directed well forward and away
from the coronet, rather than upwards; and the coloration is
distinctly different, especially the presence of spots laterally
on the snout, that are absent in H. dahli.
Material examined. ZMB 9387, holotype, photograph,
Naturalists Channel, NW Australia, 10 fm, Apr 1876. WAM
P26674-006, 9, height 70.2 nun, Dirk Hartog I., WA, 26°08'S
113°10'E, rockpool, 0.5 m, J.B. Hutchins, 18 Apr 1979. WAM
P9403, 9, height 54 mm, Shark Bay, WA, N.E. Milward, no date.
AMS IA4276, juvenile, height 40 mm, Broome, WA, 18°01'S
122°12'E, A.A. Livingstone, 3 Sep 1929.
Hippocampus biocellatus n.sp.
False-eyed Seahorse
Fig. 15
Hippocampus sp. 5 Kuiter, 2000, p. 50.
Type material. Holotype: WAM P9398, height 94.8 mm,
S, Kok Island to Quobba Point, Shark Bay, Western
Australia, W. & W. Poole Bros, Bluefin, 23-30 Aug 1958.
Paratypes: WAM P28611-005, height 98 mm, Shark Bay,
WA, 25°25'S 113°30'E, RV Peron, Mar 1966. WAM
P30081-002, height c. 60 mm, Shark Bay, WA, 26°09’S
113°13'E, box trawl, 1.0-1.5 m, J.B. Hutchins et al., 29
Mar 1990. WAM P14584, height 108 mm, Shark Bay, R.J.
McKay, 3 Apr 1962. WAM P9402, height 68 mm, Shark
Bay, R.J. McKay, 1962. WAM P30998-001, height c. 95
mm, Dorre I., off Quoin Bluff, Shark Bay, dredge, 12 m, L.
Marsh, 13 May 1995.
Diagnosis. Dorsal-fin rays 22-23; pectoral-fin rays 16-18;
anal-fin rays 4-5, the rays divided at base; tail rings 36-38;
subdorsal spines 4/0,0,1,1 or 5/0,0,1,1,1; nose ridge slightly
raised, without a spine; spine above eyes moderately large,
angled back and laterally outward; lateral head spine
moderately large and recurved; spines behind and below
eye small; coronet slightly raised, apex with 5 blunt
diverging spines; upper shoulder-ring spine of small to
moderate size, situated beside gill opening; central shoulder¬
ring spine small and tubercle-like, lower shoulder-ring spine
of moderate size and directed laterally outward; superior
trunk and tail ridges with broad thorn-like blunt spines,
enlarged on some rings at regular intervals; superior tail
ridge with tubercle-like spines of moderate size, angled
backward; trunk ridges followed by connecting tail ridges
with spines and tubercles becoming gradually smaller
posteriorly; lateral line with distinct pores, on trunk rings
just above lateral ridge, continuing onto tail to 18th-23rd
tail ring, each pore between raised papillae.
Description. Head moderately long, 70% of trunk length;
snout moderately long, 40% of head length; dorsal fin with
22 rays (23 rays in three of five paratypes), base long,
reaching well over 3 trunk rings and 1 tail ring (2 tail rings
in three of five paratypes); pectoral-fin rays 16 (17 in four
paratypes and 18 in one paratype); anal fin of moderate
size, with 4 rays divided at basal joints (5 rays divided at
basal joints in one of five paratypes); trunk rings 11; tail
rings 36 (36-38); small spine above eyes, its length less
than diameter of eye pupil, directed backward and laterally
outward; a moderately large lateral head spine, curving
backwards; a small upward directed nape spine; small spine
behind eye; small spine below eye; upper shoulder-ring
spine small, situated at gill opening directed laterally
outward, central spine developed as a tubercle at level of
7th-9th last pectoral-fin ray, and lower spine moderately
large and directed laterally outward; coronet slightly raised,
apex star-like with 5 blunt diverging spines; neck-ridge
centrally raised with small thorn-like spine and a small
Figure 15. Hippocampus biocellatus. A, holotype, S , WAM P9398, Shark Bay, WA. B, paratype, 9, WAM P28611-
005, Shark Bay, WA. C, paratype, 9, WAM P30081-002, Shark Bay, WA.
312 Records of the Australian Museum (2001) Vol. 53
Figure 16. Hippocampus biocellatus. Collection sites of specimens
examined.
slender spine at posterior end; superior trunk ridge with
angular tubercles of moderate size, enlarged on 1st, 4th,
and 7th to 11th rings; lateral trunk ridge with similar
tubercles beginning on 2nd ring, increasing in size of
subsequent rings, slanting downwards on last few rings;
ventral trunk ridge with thick membranous keel of skin
along edge; subdorsal spines 4/0,0,1,1 (5/0,0,1,1,1 in one
of five paratypes); superior tail ridge spines well developed
anteriorly, becoming smaller posteriorly, reaching to 14th
ring, with slightly enlarged spines on 5th and 8th rings;
inferior tail ridge a continuation of inferior trunk ridge,
becoming less prominent over first 9 tail rings, subsequent
rings with smooth edge; lateral line pores usually obvious
with dark papillae raised on sides to 21st tail ring (to 23 in
one of five paratypes). Largest specimen 108 mm in height.
Colour in life : blotched patterns of pale cream-grey to pale
brown with blackish scribbles and fine yellowish cream
spots mixed in. Head often with thin black lines around eye
and over operculum. Two distinctive black ocelli surrounded
with yellow laterally on upper sides of the trunk, situated
on enlarged superior ridge spines of 4th and 7th rings. Two
thin longitudinal dark lines along dorsal fin, one marginally
and one centrally. Colour in alcohol : similar to fresh
material, with dark colours reddish brown and black spots
on back remaining distinct.
Distribution (Fig. 16). Appears to be restricted to the Shark
Bay region of Western Australia, where it occurs in shallow
algae or weedy reef habitats from the intertidal zone to a
depth of about 20 m.
Remarks. This species is named biocellatus, from the Latin
bi (two) and ocellus (eye) in reference to the double eye¬
like spots on back, which may serve to distract predators.
Seahorses usually lean forwards and often tuck their head
below their trunk when threatened. In this position the eye¬
like spots would be positioned horizontally on top. Amongst
weed this might look more like a crab or a fish that would
be more of a threat to “seahorse predator” than a potential
meal. Males of this species have a deep keel-like skin
membrane on the ventral trunk ridge that maybe used during
courtship display. This species is very similar to H.
planifrons which, in addition to colour, differs from it in
having a longer and more slender trunk, the trunk length
length more than twice the depth, rather than less than twice
the depth. Hippocampus planifrons also has generally
smaller spines compared to those of H. biocellatus,
especially subdorsally.
Hippocampus breviceps
Short-head Seahorse
Fig. 17
Hippocampus breviceps Peters, 1869. Adelaide, South Australia.
Diagnosis. Dorsal-fin rays 21-22 (usually 22); pectoral-
fin rays 13-14; head and body rather fleshy without obvious
spines, but often with long filaments above eyes, on nape,
coronet and superior trunk ridges anterior to dorsal fin;
coronet distinctly raised with fleshy covering and apex
rounded with up to 5 fleshy filaments. Fin rays : dorsal 21-
22 (usually 22); pectoral 13-14; anal 4. Rings: subdorsal 3
+ 1; trunk 11; tail 38-42. Spines or tubercles: subdorsal 3-
4/0,0,1,1 or 3-4/0,0,1,0; no spines evident on head, covered
by thick fleshy skin and usually with small to long dermal
appendages above eyes, at nape and on superior trunk
ridges; low rounded tubercles intermittently on lateral and
inferior trunk ridges, and on superior tail ridge, the most
prominent beside dorsal-fin base. Lateral line: distinct with
large pores on rings just above lateral ridge on trunk,
continuing onto tail to about 20th tail ring; each pore usually
with several papillae. Coronet: distinctly raised with fleshy
covering; apex rounded with up to 5 fleshy filaments, one
usually at centre. Colour in life: drab grey to bright yellow-
orange with mix of numerous small black spots and black-
ringed white ocelli over much of trunk and anterior part of
tail. Colour in alcohol: mainly cream to pale brown with
dark spots on head and ocelli in the form of tiny dark circles
mixed with dark spots on head and trunk.
Measurements. Largest specimen examined with a height
of 62 mm, but much larger individuals known, estimated to
10 cm in height. Length of trunk is 32-35% of height; head
large, about 78-85% of trunk length; snout short in adults,
about 33% of head, but proportionally longer in juveniles,
up to 50% of head when small.
Distribution (Fig. 18). South Australian gulfs to the Bass
Strait region of Victoria and Tasmania, but apparently absent
from eastern Victorian waters. Occurs in protected bays or
estuaries. Associates with brown algae Sargassum spp. on
low, shallow reefs.
Remarks. The holotype of Hippocampus breviceps ZMB
7082 is missing. However, its description and locality can
only apply to this taxon, as there are no other similar species
in the type locality. Adults of this small temperate species
are normally covered with thick fleshy skin and have
appendages on the head and over the back that are usually
long in males (Fig. 17A). The species lacks spines, and
tubercles are few or reduced, usually being best developed
along the dorsal-fin base and over the superior ridge of the
tail, although they are often mostly fleshy.
Material examined. NMV A17236, <S , height 62 mm, Port Phillip
Bay, Vic., 37°59.8'S 145°02.1'E, depth, 3 m, R. Ickeringill & M.
Lockett, 9 Dec 1995. NMV A13008, 9, height 55 mm,
Kuiter: Australian Hippocampus revision 313
Figure 17. Hippocampus breviceps. A, 6, NMV A17236. B, $ NMV A13008, Port Phillip Bay, Vic. C, $ NMV
A19732, Port Phillip Bay, Vic. D, juvenile, NMV A17232, Port Phillip Bay, Vic.
Figure 18. Hippocampus breviceps. Collection sites of specimens
examined and localities of specimens photographed underwater.
Williamstown, Port Phillip Bay, Vic., 37°52'S 144°53'E, R.J. King,
28 Aug 1969. NMV A19732, 9, height 52 mm, Port Arlington,
Port Phillip Bay, Vic., depth 3 m, M. Lockett & T. Bardsley, 9 Apr
1997. NMV A17232, juveniles (3), heights 28-38 mm, Corio Bay,
Port P hilli p Bay, Vic., 38°06. l'S 144°22.6'E, depth 2 m, T. Bardsley
& R. Ickeringill, 2 Dec 1995.
Hippocampus tuberculatus
Knobby Seahorse
Fig. 19
Hippocampus tuberculatus Castelnau, 1875. Swan River, Western,
Australia.
Hippocampus breviceps (non Castelnau) Paxton et al., 1989: in part.
Hippocampus breviceps (non Castelnau) Lourie etal., 1999: in part.
Diagnosis. Dorsal-fin rays 20-21; pectoral-fin rays 14-15;
coronet tall and angled back, square to rounded in cross
section, without spines on corners, but with small spine
centrally on apex; adults with large tubercles on back and
tail. Fin rays: dorsal 20-21; pectoral 14-15; anal 4. Rings:
subdorsal 3 + 1; trunk 11; tail 36-37. Spines or tubercles:
subdorsal spines 4/0,0,1,0; moderately long spine above
eyes, with secondary smaller spine anteriorly at base, similar
in shape to nose ridge; small spine behind eye; large lateral
head spine; nape spine of moderate size, directed upward;
2 separate low angular spines below eye; 3 moderate to
long spines on shoulder ring, uppermost at gill-opening,
central spine at level of last pectoral-fin ray, and lowermost
largest and recurving; neck ridge with 2 small spines;
superior trunk ridge with blunt rounded spines of moderate
size, enlarged on rings 1, 3, 5, and subdorsally; lateral trunk
ridge with enlarged spines on rings 3, 5, and 7; inferior
trunk ridge with last 3 spines enlarged; ventral trunk with
low downward directed spines; superior tail ridge with
enlarged spines up to about eye-size on every 2nd or 3rd
ring, becoming gradually smaller posteriorly; inferior tail
ridge a continuation of trunk ridge, with spines gradually
reducing in size. Lateral line : well developed with pores
on low-tubes, visible to about 28th tail ring. Coronet:
coronet tall and angled back, square to rounded in cross
section, without spines on corners, but with small spine
centrally on apex, angled upward when young, becoming
smooth and rounded in mature adults. Colour in life:
specimens from floating Sargassum dark brown to almost
black with a well defined white area over interorbital
extending to base of coronet and to tip of snout. Adults on
reefs mainly yellow to brown, with white area over front of
head, some black spotting over operculum, and often
tubercles over back bright red. Colour in alcohol: mainly
brown; some with fine black spots on operculum or below
dorsal-fin base.
Measurements. Largest specimen examined 74 mm in height.
Trunk short, 26-32% of height; head moderately large, 75-
80% of trunk length; snout short, about 35-50% of head length
(proportionally longest in young and sub-adults).
Distribution (Fig. 20). Restricted to Western Australia
between Perth and Onslow. Hippocampus tuberculatus
is often found offshore in floating Sargassum during
juvenile and sub-adult stages, some males have fully
314 Records of the Australian Museum (2001) Vol. 53
Figure 19. Hippocampus tuberculatus. A, 6, WAM P28259-001, Swan River, Perth, WA. B, 9, WAM P25808-
001, off Perth, WA. C, 9, WAM P29245-002, Rockingham, WA. D 8, E 9, WAM P24564.001, off Onslow, WA.
Figure 20. Hippocampus tuberculatus . Collection sites of
specimens examined.
developed pouches. Adults settle on sponge reefs in
depths of about 20 m.
Remarks. The name tuberculatus is very appropriate for
this species, with its large tubercles on the back and tail of
adults. The holotype MNHN A4539 was examined by Sara
Lourie, but she placed Hippocampus tuberculatus in
synonymy with H. breviceps, and illustrating latter with a
specimen of H. tuberculatus (Lourie etal., 1999, p. 141). This
species has an angular nose ridge and double spines above
each eye and is generally spinier than the closely related H.
breviceps , from temperate latitudes of southeastern Australia.
The two species treated as synonyms until now.
Material examined. WAM P28259-001, 8, height 74 mm,
Fremantle, WA, 32°03'S 115°44'E, P. Crystal, 25 Jun 1980. WAM
P25808-001, 9, height 54 mm, E 5 Fathom Bank, WA, 32°16'S
115°39'E, Flinders , 20 m, 29 Nov 1977. WAM P29245-002, 9,
height 50 mm, Rockingham, WA, 32° 17'S 115°42'E, J.B. Hutchins,
Feb 1976. WAM P24564-001 (9), heights 44-51 mm, Onslow
area, WA, 21°38'S 115°07'E, G. Lang, Nov 1973.
Hippocampus taeniopterus
Common Seahorse
Fig. 21
Hippocampus taeniopterus Bleeker, 1852b. Ambon, Indonesia.
Hippocampus melanospilos Bleeker, 1854a. Ambon, Indonesia.
Hippocampus kuda (non Bleeker) Paxton et al., 1989.
Hippocampus kuda (non Bleeker) Randall et al., 1990.
Hippocampus kuda (non Bleeker) Lourie et al., 1999: in part.
Diagnosis. Dorsal-fin rays 17-18; pectoral-fin rays 16;
subdorsal rings 2 + 1-2; tail rings 34-35; spines low and blunt
in juveniles, reducing or becoming rounded and knob-like in
adults; small nose-spine present; shoulder-ring spines small,
upper at gill-opening, centre at level of last pectoral-fin ray,
lower single and not enlarged; large specimens often with many
papillae on nape and interorbital and body. Fin rays’, dorsal
17-18; pectoral 16 (one specimen with 18 on one side); anal
4. Rings’, subdorsal 2 + 1-2; trunk 11; tail 34-35. Spines or
tubercles’, subdorsal 3-4/0,1,1-0,1,1,1. Spines low and blunt
when juvenile, reduced or rounded and knob-like in adults.
Small spine above eye; nose spine small; nape spine small;
small spine behind eye; double low spines below eye; shoulder¬
ring spines small, uppermost at gill-opening, central spine at
level of last pectoral-fin ray, lowermost single and not enlarged;
trunk ridges with enlarged spines or tubercles on rings 1, 4, 7
or 8 and 11, and on tail on about 5th, 9th, 12th, and 15th ring,
though varying in position by up to one ring, largest near pouch
area of male, becoming gradually smaller posteriorly on tail.
Lateral line’, pores small, more or less distinct, becoming
intermittent on tail, ranging to 20th ring. Pores often positioned
between 2 large papillae, although large specimens usually
have many additional papillae, on nape and interorbital, as
well as body. Coronet: small but well developed with 5
diverging sharp spines on apex in young, becoming
proportionally smaller with large adults, having smooth apex
and 3 rounded points angled backwards. Colour in life: (based
on underwater photographs taken in Milne Bay, PNG) grey to
brown, often with fine dark spots and pale scribbles and
striations; females sometimes yellow with pupil-sized black
spots that may elongate on trunk over rings. Colour in alcohol:
pale to dark brown, usually covered with small dark spots.
Kuiter: Australian Hippocampus revision 315
Figure 21. Hippocampus taeniopterus . A, S, AMS 19309, Darwin Harbour, NT. B, $, AMS 19203, SE coast, New
Guinea. C, 6, QM 120093, Russell River, Qld.
Measurements. Largest specimen examined 20 cm in
height. Snout thick in adults, its length about equal to
distance between eye and gill-opening.
Distribution (Fig. 22). Occurring in the Moluccan seas and
ranging south to Papua New Guinea and tropical eastern
Australia. A shallow water species, found mainly in coastal
areas along edges of seagrass beds or in mangroves to about
15 m depth. Juveniles as well as adults sometimes found in
floating weeds well offshore, especially during the wet season.
Remarks. No types are known and Hippocampus
taeniopterus has been treated by recent authors as a
Figure 22. Hippocampus taeniopterus. Collection sites of
specimens examined.
synonym of H. kuda Bleeker, 1852a. I follow Bleeker in
recognising it as a valid species. Hippocampus taeniopterus
is common in most parts of its range, but it appears to be
less so in Australian waters. Australian specimens are rare
in collections, but this may be due to a shortage of collecting
in the remote northern region and inshore habitats along
the Queensland coast. As adults regularly occur in floating
weeds they may travel a long way south from areas like
southeastern Papua New Guinea where the species is known
to be abundant. Some Australian specimens like a Moreton
Bay specimen (AMS 112555), may represent expatriates.
In addition to slight meristic and morphological
differences, H. kuda is distinguishable from H. taeniopterus
in usually having dermal appendages on the head spines,
coronet, and on the enlarged spines or tubercles of superior
ridges. Bleeker described H. kuda from a specimen from
Singapore and in his remarks following his descriptions of
H. moluccensis (1852b) suggested that the dermal
appendages are diagnostic. This is consistent with specimens
from Sri Lanka observed in a Melbourne aquarium, which
have filaments below the snout and to various degrees on
nape and coronet. Juveniles are somewhat spiny over
superior ridges and have spiny coronets. Some have
distinctive double white spots on trunk, especially sub-
dorsally. An illustration in Bleeker (1983, fig. 5, pi. 449),
captioned “ Hippocampus guttulatus Cuv. = kuda Blkr.”
matches the description of kuda, Bleeker’s further remarks
in 1852b, and additional specimens from Singapore (pers.
obs.). The use in the Atlas of the name H. guttulatus, a
European species which often has long appendages similar
to H. kuda seems incorrect. Hippocampus guttulatus is
readily distinguished from H. kuda by a much taller dorsal
316 Records of the Australian Museum (2001) Vol. 53
fin and colour patterns. Some synonyms may have been
created by ichthyologists who continued Bleeker’s work
after his death, possibly causing errors. Photographs of H.
kuda in the literature were taken in the Ryukyu Is, southern
Japan (Kawanabe & Mizuno, 1989), Cebu, Philippines
(Allen & Steene, 1987): the latter photograph was used in
Randall et al., 1990, but replaced by a photograph of H.
taeniopterus from Papua New Guinea in the 2nd edition
1997 (Steene, pers. com.). On the basis of published
photographs and my own observations H. kuda ranges from
Andaman Sea to southern Japan, but there are no records east
of Wallace’s Line; H. taeniopterus is its replacement there.
Material examined. QM 120093, S, height 115 mm, Russell
River, near mouth among mangrove roots, NE Qld, J. Johnson,
Nov 1982. NMV A8952, 6, height 95 mm, Cooktown, Qld,
15°28'S 145°15'E, 1943. AMS 19309, S, height 20 cm, Port
Darwin, NT, 12°27'S 130°48'E, S.W. Cristie, 1908. AMS 19203,
$, height 135 mm, SE coast New Guinea, Macleay, 1907. AMS
112555, 5 , height 115 mm, Moreton Bay, Qld, 27°S 153°E,
Amateur Fishermans Association of Qld, 1912. AMS 138392-001,
juvenile, height 21 mm, Lizard I., Qld, 14°40'S 145°28'E, surface
capture over 30 m depth, M. Emslie, 31 Jan 1995.
Hippocampus tristis
Sad Seahorse
Fig. 23
Hippocampus tristis Castelnau, 1872. Melbourne fish market.
Hippocampus punctulatus (non Kaup) Ogilby, 1889: 732.
Hippocampus sp. A, Allen et al., 1976. Lord Howe I.
Hippocampus whitei (non Bleeker) Paxton et al., 1989: in part.
Hippocampus kuda (non Bleeker) Lourie et al., 1999: in part.
Hippocampus kelloggi (non Jordan & Snyder) Lourie etal., 1999:
in part.
Diagnosis. Dorsal-fin rays 18-19; pectoral-fin rays 18-19;
subdorsal rings 2 + 1-2; tail rings 35-37; nose profile
straight, small spine present; head and body spines small
and blunt, proportionally smallest in largest specimens, most
trunk ridges with low tubercles; upper shoulder-ring spine
small, situated just below gill opening, and lower shoulder¬
ring spine thick, knob-like, single or double with less
developed secondary spine anteriorly; neck ridge short and
distinctly raised above gill opening. Fin rays: dorsal 18-
19; pectoral 18-19; anal 4. Rings: subdorsal 2+1-2; trunk
11; tail 35-37. Spines or tubercles: subdorsal 3/0,1,0. Head
and body spines small and blunt, becoming relatively
smaller with age, most trunk ridges with low tubercles;
tubercles on males mainly on inferior trunk ridges and on
tail along pouch region; nose spine small and greatly
reduced in large individuals; spine above eye small, directed
upward, lateral head spine moderately long; shoulder-ring
spines variable from poorly to moderately developed,
uppermost small just below gill-opening, central spine at
level of last pectoral-fin ray, and lowermost thick, knob¬
like, single or double with less developed secondary spine
anteriorly; neck ridge short and distinctly raised above gill
opening. Lateral line: pores small without papillae, more
or less distinct, becoming intermittent on tail, ranging to
17th ring. Coronet: small but well developed with 5
Figure 24. Hippocampus tristis. Collection sites of specimens
examined.
Figure 23. Hippocampus tristis. A, 6, QM 11008, Lord Howe I., NSW. B, 6, AMS 138517-001, Clarence River
area, NSW. C & D, $ and juvenile, QM 130574, Stradbroke I., Qld. E, syntype, MNHN A4538, unknown locality,
photo by Sara Lourie.
Kuiter: Australian Hippocampus revision 317
diverging short spines on apex in young, becoming less
prominent with age, almost smooth in large adults. Colour
in life : drab grey to pale brown with dusky head and trunk,
sometimes with numerous tiny white spots in longitudinal
lines. Juveniles occasionally pale yellow; large juveniles
occasionally with prominent saddle-like blotches,
contrasting with general colour, at 1st, 4th and 8th trunk
rings, and along tail. Colour in alcohol : cream to pale brown,
some dark spotting near eyes on recently collected material.
Measurements. Largest specimen examined 222 mm in height.
Snout thick in adults, its length about equal to postorbital length.
Distribution (Fig. 24). The extent of the range of
Hippocampus tristis is uncertain. It is known from off
Brunswick in northern New South Wales, from Lord Howe
I. and from southern Queensland. All specimens from
Queensland except one lot from Stradbroke I. have as the
locality “South Queensland coast”. Most specimens came
from trawls between 18-53 m depth and some specimens
washed up on beaches (probably trawl-casualties).
Remarks. Type specimens of Hippocampus tristis (MNHN
A4537 & A4538), both adult males, were probably taken as
bycatch by commercial fishermen in northern New South
Wales waters. Photographs of the specimens, provided by Sara
Lourie, show specimens that are virtually identical to specimens
examined of similar size, including in the shape of the head,
the high neck-ridge and the proportional features of the head
and trunk. This species has been misidentified as H. whitei, H.
kuda, and H. kelloggi. Juveniles of H. tristis may resemble
females of H whitei , small adults could by mistaken for H.
taeniopterus, and very large individuals for H. kelloggi.
Hippocampus tristis is readily distinguished from H. whitei
by the smaller coronet and position of the uppermost shoulder¬
ring spine, at gill opening in H. tristis and near pectoral fin
base in H. whitei. Morphologically, H. tristis and H.
taeniopterus are very similar and the two are best distinguished
by the lowermost shoulder-ring spine that is single in H.
taeniopterus and usually double (anterior one sometimes small
or not obvious) in H. tristis. The fin-ray counts are modally
higher in H. tristis than in H. taeniopterus (D 18-19, P 18-19
versus D 17-18, P 16 respectively) and the coronet is more
distinct in H. tristis with 5 diverging spines on apex, compared
to H. taeniopterus with 5 small diverging spines on apex in
juveniles, reducing to 3 posteriorly directed spines in adults,
and in addition the coronet leans back more in H. taeniopterus
compared to H. tristis. Hippocampus kelloggi is a large species,
reaching 28 cm in height, occurring in southern Japanese waters
and sub-tropical regions of the northern China Seas, but the
name is often applied to other Indo-Pacific species that reach
a large size.
Material examined. MNHN A4537 & A4538, syntypes,
photographs, Melbourne Market, Vic., Australia, 1872. QM 11008,
6 , height 222 mm, Lord Howe I., J.D. Ogilby, 1889. AMS 11959, 6 ,
height 220 mm, Lord Howe I., E. Saunders, no date. AMS 138517-
001, <J, height 185 mm, off Iluka, NSW, 29°21'S 153°23'E, depth
18-28 m, K. Graham, FRV Kapala, 14Nov 1995. AMS 133593-001,
S, height 105 mm, off Bmnswick, NSW, 28°21'S 153°40'E, depth
53 m, K. Graham, FRV Kapala, 10 Nov 1991. QM 13402, 9 , height
170 mm, South Queensland Coast, Qld Fisheries Department, 18 Jun
1919. QM 19726-7(2), 9 9 , heights 135 and 145 mm, SE Qld, R.
Elks, 6 May 1969. QM 130574 (3), 2 9 9 & 6, heights 180-75 mm,
Stradbroke I., Qld, 27°29'S 153°31'E, beach washed, M. Mathieson,
3 May 1996.
Hippocampus alatus n.sp.
Winged Seahorse
Fig. 25
Hippocampus sp. 3 Kuiter, 2000, p. 38.
Type material. Holotype: AMS 120771-102, 9 , height
119 mm, east coast of Cape York, Queensland, 11°37'S
142°56'E, prawn trawl, depth 16-18 m, AIMS-AMS party,
18 Feb 1979. Paratypes: AMS 115557-077, S, height 117
mm, Gulf of Carpentaria, Qld, 17°25'S 140°10'E, prawn
trawl, depth 10 m, I.S.R. Munro, 27 Nov 1963. QM 130556,
juvenile, height 69.5 mm, Gulf of Carpentaria, NT,
10°03.2'S 137°11.2'E, dredge, depth 42 m, S. Cook & J.
Johnson, 21 Nov 1990. QM 127965, juvenile, height 78
mm, Gulf of Carpentaria, Qld, 15°57.6’S 138°41.8'E,
dredge, depth 25 m, S. Cook & J. Johnson, 11 Dec 1990.
NTM S13267-004, 9, height 70 mm, Gulf of Carpentaria,
Qld, 11°18'S 141°38'E, depth 17 m, R. Williams, 27 Nov
1991. NTM S11580-003, juvenile, height 71 mm, off
Dampier Archipelago, WA, 20°03’S 115°48’E, depth 50-
53 m, L. Bullard, 8 May 1983. NTM S10959-001, 9, height
136 mm, off Port Hedland, WA, 18°55’S 119°37’E, depth
80 m, R. Williams, 18 Apr 1983. WAM P28003-001, 9 ,
height 81 mm, off Monte Bello Is, WA, 20°17’S 116 o 0FE,
depth 55-64 m, Soela, 2 Dec 1979.
Diagnosis. Dorsal-fin rays 15-18 (usually 17); pectoral-
fin rays 16-18; trunk rings 11; tail rings 34-36; subdorsal
spines 2-3/0-0.5,l,0-0.5, usually enlarged on 11th trunk
ring; nose spine absent; spine above eyes of moderate
length, length reaching pupil diameter; lateral head spine
large, usually larger than spine above eye; coronet well
developed with 5-7 blunt spines, apex rough and rugose;
upper shoulder-ring spine at gill opening; lower shoulder¬
ring spine low and thick, very broad when single or divided
into two rounded tips; superior trunk and tail ridges with
enlarged spines, forming laterally directed pairs at regular
intervals with 2 or 3 on trunk and 3 or 4 on tail, greatly
produced in young and least prominent in males; long and
sometimes broad dermal flaps, often frilled along edges,
on nape, lateral head spines, and enlarged body spines,
usually attached posteriorly and just below tip; lateral line
with pores on each trunk ring, running just above inferior
ridge, continuing on tail to 12th-15th ring.
Description. Head large, 85% of trunk length, snout
reaching 11th ring when head against trunk; snout deep,
least depth 25% of snout length; dorsal fin with 17 rays
(only male paratype with 15, range otherwise 17-18), base
short, its length twice its height; pectoral-fin rays 17 (16-
18); anal-fin rays 4; trunk rings 11; tail rings 36 (34-36);
subdorsal rings 2+1; moderately long spine above eyes, its
length about equal to eye pupil, perpendicular to snout;
slightly larger lateral head spine, directed laterally, with large
dermal flap; moderately large nape spine with dermal flap
of similar size; small spine behind eye; low spine below
eye; 3 moderately large spines on shoulder ring, uppermost
just below gill opening, central spine largest and at level of
last pectoral-fin ray, and lowermost double with blunt and
rounded tips; coronet distinct with an irregular star
arrangement of rugose tipped spines on apex; neck ridge
raised centrally with rugose tips and with tubercle at
posterior end; superior trunk ridge with enlarged blunt
318 Records of the Australian Museum (2001) Vol. 53
Figure 25. Hippocampus alatus. A, holotype, 9, AMS 120771-102, east coast Cape York, Qld. B, paratype, 6,
AMS 115557-077, Gulf of Carpentaria, Qld. C, paratype, $, WAM P28003-001, off Monte Bello Is, WA. D,
paratype, juvenile, NTM SI3267-004, Gulf of Carpentaria, Qld.
spines on 1st, 4th, 7th and 11th rings, 1st and 4th with dermal
flaps; lateral trunk ridge with series of blunt spines
beginning on 2nd trunk ring, usually spine on 7th enlarged
and with dermal flap; inferior trunk ridge with blunt spines,
most prominent on 7-10th ring; ventral trunk spines poorly
developed on last few rings; subdorsal spines 3/0,1,0 with
central spine on trunk ridge and spine on tail ridge of
moderate size (some paratypes with only 2 spines on trunk
ridge and one paratype with small spines on nodes beside
spine on tail ridge); superior tail ridge with spines of
moderate size beginning on 2nd tail ring, gradually
becoming progressively smaller to 17th ring, except for
enlarged spines bearing dermal flaps on 5th and 8th rings
(spines greatly reduced in largest female); inferior tail ridge
continues with inferior trunk ridge, with spines becoming
progressively smaller posteriorly to 9th ring; lateral line
present although pores small and only detectable
intermittently to about 12th tail ring (pores distinct in
Figure 26. Hippocampus alatus. Collection sites of specimens
examined.
paratypes). Largest specimen, female, 136 mm in height.
Largest male, with fully developed pouch, 117 mm in height.
Colour in life : (based on colour transparency, taken by Bob
Halstead in Milne Bay, Papua New Guinea, of two females).
Pale pinkish white with numerous minute dark spots and
broad reddish grey saddle-like markings over trunk, on
anterior part of tail and around eyes. Colour in alcohol :
adults pale creamy white with numerous tiny brown spots
and dark saddle-like markings on trunk and anterior part of
tail around enlarged spines on superior ridges. Dermal flaps
dark brown to black.
Distribution (Fig. 26). Northern Australia from the Dampier
Archipelago, Western Australia, throughout the Gulf of
Carpentaria, to the tip of Cape York, and southeastern Papua
New Guinea. Known depth range 10-80 m depth. Soft
bottom habitat.
Remarks. This species is named alatus , derived from the
Latin alatus, meaning winged, in reference to the paired
spines on the superior trunk ridges that are directed outward
and have broad dermal flaps, resembling wings. This feature
is most prominent in juveniles and females, and may be
less developed in individuals living in deep water.
Specimens from Western Australia coming from depths over
50 m have smaller dermal flaps but are located on the same
spines as those from shallower depths in the Gulf of
Carpentaria. One specimen (NTM SI 1580-003) lacks the
nape spine but has dermal flap development where the spine
would be located and may have been damaged at some
stage. The lower shoulder-ring spine is normally single,
blunt and rounded at the tip, but a secondary spine may
develop just in front of it, somet im es only on one side of
the head. Adult males are not as spiny as females, and may
be smooth on many rings. Because of this species’ preferred
habitat, specimens in collections have been taken with
dredges or prawns trawl. In Papua New Guinea, specimens
were found on deep open sand slopes with few sponges
Kuiter: Australian Hippocampus revision 319
and gorgonian corals in an area that is influenced by strong
tidal currents. Hippocampus alatus is most similar to H.
queenslandicus the main differences between the two
involving the development of spines. The tail of H.
queenslandicus is always spiny, superior ridge with a series
of prominent spines present to at least 12th ring, while, in H.
alatus, the spines in comparative positions are greatly reduced
in adults, with prominent spines on the tail spaced at intervals
of several rings, in adult as well as juvenile stages.
Specimens comprising the type series were initially
identified variously as H. kuda, H. barbouri?, H. hystrix or
H. histrix and H. spinosissimus. Semi-smooth adults were
confused with H. kuda and small juveniles having a spiny
appearance were confused with several spiny species.
Though superficially similar, smaller H. alatus can be
distinguished in lacking a prominent nose spine.
Hippocampus queenslandicus
Queensland Seahorse
Fig. 27
Hippocampus whitei (non Bleeker) Grant, 1987.
Hippocampus queenslandicus Horne, 2001. Off Townsville, Qld.
Diagnosis. Dorsal-fin rays 17 (rarely 18); pectoral-fin rays
17-18 (rarely 16 or 19); trunk rings 11; tail rings 35-36;
snout long, its length just over 50% of head length, and
deep, its narrowest depth about 30% of its length; spine
above eye as long as eye-pupil, directed upward and slightly
backwards; nose spine absent and nose profile straight;
shoulder ring spines long, uppermost at gill opening, central
spine at about level of 13th pectoral-fin ray, lowermost with
blunt rounded spine and second less developed spine
directed forward; lateral head spine moderately long;
coronet small with 5 diverging spines at apex, posterior-
most longest; superior tail ridge spines moderately long,
enlarged on 4th, 7th and 10th ring; inferior tail ridge with
spines of moderate length along pouch section of males;
lateral line distinct with small pores, most on tail associated
with double papillae, extending to 16th tail ring. Fin rays :
dorsal 17 (one with 18); pectoral 17-18 (one with 16/17
and one with 18/19); anal 4. Rings : subdorsal 2+1; trunk
11; tail 35-36. Spines or tubercles’, subdorsal 3/0,1,0. Head
and body spines short to moderate sized with blunt to sharp
tips; spine above eye of moderate size, its length as long as
eye-pupil diameter, directed upward and slightly backwards;
nose spine absent and nose profile straight (including in
juveniles); nape spine small but distinct, perpendicular to
nape; spine behind eye small; two spines below eye, low
and blunt; shoulder ring spines long, uppermost at gill¬
opening, central spine at level of about 13th pectoral-fin
ray, lowermost with blunt rounded spine and second smaller
spine directed forward; lateral head spine of moderately
length; neck ridge with a moderate sized spine above gill¬
opening and a small spine at posterior end of ridge; superior
trunk ridge with short but broad-based spines, slightly
enlarged at 1st, 4th, 7th and 11th ring; lateral ridge with
spines from 2nd to 10th ring, enlarged on 4th, and 7-10th
rings; inferior trunk ridge with series of spines on 6th to
11th ring, from short to long respectively; ventral trunk ridge
with low angular spines on last 5 rings; superior tail ridge
spines moderately long anteriorly, becoming progressively
smaller posteriorly to about 16th ring, enlarged on 4th, 7th
and 10th rings; inferior tail ridge with spines of moderate
size along pouch section of males. Lateral line : distinct
with small pores, most associated with double papillae
on tail, ranging to 16th tail ring. Coronet’, small with 5
diverging spines at apex, posterior-most longest. Colour
in life : (based on colour transparencies of living specimens
by Mark Norman at James Cook University’s Aquaculture
Department) Yellow-orange to deep red or dark brown,
Figure 27. Hippocampus queenslandicus. A, 8, NMV A21578, off Townsville, Qld. B, 9 , NMV A21579, off
Townsville, Qld. C, juvenile, AMS 14750, Southport, Qld.
320 Records of the Australian Museum (2001) Vol. 53
Figure 28. Hippocampus queenslandicus. Collection sites of
specimens examined.
snout and front of head usually darker or dusky; pale grey
saddles or bands with dark margins over trunk and tail, one
broadly over first 3 trunk rings, a narrower saddle on 7th ring
usually reaching ventral ridge, and one or two broad and narrow
saddles anteriorly on tail. Colour in alcohol : pale to blackish
brown with dusky or grey saddle-like markings.
Measurements. Largest specimen examined 116 mm in
height (some specimens observed an estimated 125 mm in
height). Head moderately long, 83% of trunk length; snout
long, just over 50% of head length and deep, its narrowest
depth about 30% of its length.
Distribution (Fig. 28). Inner reef waters of Queensland
from Southport north to Princess Charlotte Bay, Cape York.
Usually trawled in depths over 20 m, and as deep as 63 m.
Remarks. Specimens of this species were misidentified in
collections variously as Hippocampus histrix, H. kuda and
H. spinosissimus depending on their size or sex. Hippo¬
campus queenslandicus is one of a number of species in
the West Pacific that have spiny juveniles and nearly smooth
adults, but adult H. queenslandicus usually retain spines
anteriorly on the tail. The absence of a nose spine readily
distinguishes H. queenslandicus from “true” spiny species.
The saddle-like markings in H. queenslandicus are usually
prominent and similar to those of closely related species,
but the pattern is also in certain other unrelated species.
Colour patterns of closely related species are usually similar,
but details may differ between species at certain growth
stages or sex that, however small, may represent diagnostic
markings. Hippocampus queenslandicus appears to be
closely related to the stockier H. alatus, which has short
blunt spines or tubercles on the same rings and ridge nodes
that H. queenslandicus has long sharp spines. The species
differs from H. semispinosus in having a shorter, less slender
snout, and in colour. In southern Queensland H. queens¬
landicus is replaced by the similar H. tristis, a species which
lacks spines completely when adult, has more fin rays,
grows to almost twice the height, and differs considerably
in colour.
Material examined. NMV A21578, 6 , height 116 mm, Britomart
Reef, off Townsville, Qld, depth 30-50 m, M.L. Horne, Feb 1999.
NMV A21579 (4), $ $, height 102-115 mm, Britomart Reef, off
Townsville, Qld, depth 30-50 m, M.L. Horne, Feb 1999. QM
123197, 9, height 93 mm, NE of Townsville, Qld Fisheries,
trawled, 9 Dec 1984. QM 119764, 9, height 95 mm, Swains Reefs,
NE Qld, Qld Fisheries, trawled, Jun 1980. AMS 120959-003, 9 ,
height 112 mm, Princess Charlotte Bay, Cape York, Qld, 14°09'S
144°04'E, depth 22-28 m, AMS-AIMS prawn trawl, 23 Feb 1979.
AMS 14750, juvenile, height 70 mm, off Southport, Qld, 27°S
153°E, depth 27 m, D.F. McMichael, 1960. QM 123060 (2), 9 &
<?, Swains Reefs, Qld, 20°52'S 150°40'E, depth 63 m, Qld
Fisheries, 12 Sep 1986. QM 111770 (2), 9 & 6 , far north Qld,
Challenge Survey, Qld Fisheries, 30 Oct 1957. QM 120492, 9 ,
Cairns inner reef survey, trawled, Qld Fisheries, 25 Apr 1982.
QM 111773 (4), 9 9 , off Cairns, Qld, K. Bryson. AMS 115864-
002, 9 , North Keppel I., Qld, 23°05'S 154°45'E, depth 21 m, W.
Ponder, 25 Jul 1969.
Hippocampus semispinosus n.sp.
Half-spined Seahorse
Fig. 29
Hippocampus kuda (non Bleeker) Gloerfelt-Tarp & Kailola, 1984.
Hippocampus sp. 2 Kuiter, 2000, p. 36.
Type material. Holotype: NTM S10749-006, 9 , height
137 mm, east Alas Strait, Indonesia, 08°20'S 116°50'E,
trawl, depth 40-60 m, T. Gloerfelt-Tarp, Jun 1981.
Paratype: NTM SI 1920-002, S, height 120 mm, Timor
Sea, southern Indonesia or northern Australia (see remarks),
T. Gloerfelt-Tarp.
Diagnosis. Dorsal-fin rays 18; pectoral-fin rays 16-17; trunk
rings 11; tail rings 35-36; subdorsal spines 3/0,1,0-0.5,
usually enlarged on 11th trunk ring; nose ridge with straight
profile and no spine; spine above eyes small, length less
than eye diameter, angled back; lateral head spine large,
about twice in size of eye-spine, and recurving; small spine
behind eye, placed high above eye centre; coronet well
developed with 5-7 diverging blunt spines with rugose tips,
posterior spines enlarged and recurving; upper shoulder ring
spine long, situated at gill opening; lower shoulder-ring
spine moderately large, with a poorly developed second
spine anteriorly; superior trunk and tail ridges with rugose
tubercles, enlarged on 1st and 7th rings as broad spines;
superior tail ridge with spines of moderate length from 2nd
to about 15th ring, somewhat enlarged on 5th and 8th ring,
becoming gradually shorter posteriorly; lateral line with
pores intermittently on trunk rings just above inferior ridge,
extending onto tail to 17th-18th ring.
Description. Head long, 75% of trunk length, snout
reaching 11th ring when head against trunk; snout slender,
the least depth 23% of its length, and long, about 55% of
head length; dorsal-fin with 18 rays, base short over 2 trunk
rings and 1 tail ring; pectoral-fin rays 17 (16-17); anal-fin
rays 4; trunk rings 11; tail rings 35 (35-36); subdorsal rings
2 + 1; spine above eyes small, its length less than eye pupil,
directed slightly backward; lateral head spine moderately
large with a strongly recurving tip; nape spine very small
but distinct; a low tubercle behind eye, placed just above
ridge junction; no spine below eye; 3 moderately long spines
on shoulder ring, uppermost just below gill opening, central
spine at ventral level of pectoral-fin base, and lowermost
ventrally with a short second spine directed anteriorly from
base (on one side only in male paratype); coronet distinct
Kuiter: Australian Hippocampus revision 321
Figure 29. Hippocampus semispinosus. A, holotype, 9 , NTM S10749-006, Sumbawa, Indonesia. B, paratype, 6,
NTM SI 1920-002, southern Indonesia or northern Australia.
with an irregular star arrangement of rugose-tipped spines
on apex, posterior spines recurving and central spine
enlarged; neck ridge broadly raised above gill-opening with
rugose tips, and ending posteriorly with spine-like tubercle;
superior trunk ridge with enlarged blunt spines on 1st, 4th,
and 7th to 11th rings (least developed in male paratype);
lateral trunk ridge with enlarged blunt spines on 2nd, 4th,
and 7th to 10th rings; inferior trunk ridge with moderately
large blunt spines, mainly developed on 7-11th ring; ventral
trunk ridge smooth; subdorsal spines 3/0,1,0.5 with spines
on trunk ridge and one on tail ridge of moderate size,
followed by small tubercle on first tail ring; anterior superior
tail ridge spines, except on first ring, moderately large and
blunt, becoming progressively smaller to 17th ring, spines
on 5th and 8th rings slightly enlarged; inferior tail ridge
continues with inferior trunk ridge, with enlarged tubercles
on the first 5-6 rings which become progressively smaller
posteriorly; lateral line present but pores difficult to detect
and intermittently visible to about 17th tail ring. Largest
specimen, female (holotype), 137 mm in height; paratype,
male with fully developed pouch, 120 mm in height. Colour
in life : (based on photo of female holotype illustrated in
Gloerfelt-Tarp & Kailola, 1984) deep orange with grey
saddle-like blotches over the 1st and 2nd, and 6th to 8th
trunk rings, and over 3rd to 5th tail rings; tip of snout pale.
Underwater photographs taken in Bali, Indonesia of several
females and a male, thought to be this species indicate that
females consistently have these markings and range in
colour from dark-red to yellow, while at least some males
are dark brown with several large creamy-white blotches
laterally on the trunk. Colour in alcohol : holotype retains
pattern described above, with orange having faded to pale
cream. Paratype mostly dark-brown with pale blotches on
trunk, and snout and creamy-white pouch.
Distribution. Type material was trawled during a fishery
project to assess fish stocks of southern Indonesia and
northwestern Australia. The holotype came from a region
referred to as “Area B”, ranging from Bali to Timor in the
Lesser Sunda Is (Gloerfelt-Tarp & Kailola, 1984). The area
was reported as being characterised by narrow and rocky
shelves surrounding each of the islands. Habitats in the
region vary greatly from narrow passages with strong
currents to large areas with muddy substrates. The paratype
came from an assortment of fishes collected from 1980-
1983 for which no data were recorded (Helen Larson, pers.
comm.). A discrepancy exists between data presented in
Gloerfelt-Tarp & Kailola (1984), their station TGT 1813
and that accompanying the holotype, NTM S10749-006,
although longitudes and latitudes are close, Gloerfelt-Tarp
& Kailola (1984) state that this species occurs in Areas B,
C, where C is off northwestern Australia. The registration
information for NTM S10749-006 is considered here as the
correct type locality description. It is possible that the
locality provided by Gloerfelt-Tarp & Kailola (1984) may
be that of the paratype. Still, a description of this species is
presented here as there is a chance that the paratype was
collected on the northwest shelf of Australia.
Remarks. This species is named semispinosus in reference
to its spiny appearance that is intermediate between smooth
322 Records of the Australian Museum (2001) Vol. 53
and long-spined species. The female holotype was originally
identified as Hippocampus kuda, and male paratype as H.
spinosissimus, which are “smooth”, and “spiny” species
respectfully. It is most similar to H. moluccensis (Bleeker,
1852b), a species described from Ambon. Two specimens
NMV 46228 a male and NMV 46227 a female, apparently
Bleeker’s types of H. moluccensis (they match his
description exactly), have saddle-like colour patterns like
H. semispinosus, but the head and body of these specimens
are peppered with small dark spots as in H. alatus.
Hippocampus moluccensis differs morphologically from H.
semispinosus in having fewer dorsal-fin rays, (16-17 versus
18); a shorter snout, (length 2 or more times head versus
I. 8 times head or less) and differs from H. alatus, H.
queenslandicus, H. tristis in having a raised and angled nose
profile versus one that is straight. Hippocampus semi¬
spinosus has a longer and more slender snout than H. alatus
and H. queenslandicus (snout longer than 1/2 head-length,
and snout-depth 5x in its length, versus snout about 1/2
head-length, and snout-depth 3.5-4x in its length). Except
for the saddle-like marking of the female holotype, this
species differs considerably in colouration from H. alatus
and H. queenslandicus.
Hippocampus abdominalis
Eastern Potbelly Seahorse
Fig. 30
Hippocampus abdominalis Lesson, 1827. New Zealand.
Diagnosis. Dorsal-fin rays 25-28; pectoral-fin rays 15-16
(usually 15); trunk rings 12-13; tail rings 44-45; subdorsal
spines 4/0,0,1,1 or 5/0,0,1,1,1; head small, 62-77% of trunk
(adult-juvenile respectively); snout usually short, 31-37%
of head (adult-juvenile respectively). Fin rays : dorsal 25-
28; pectoral 15-16 (usually 15); anal 4. Rings: subdorsal
3-4+1-2; trunk 12-13; tail 44-45. Spines or tubercles:
subdorsal 4/0,0,1,1 or 5/0,0,1,1,1. Pelagic young with low
angular spines along trunk and anterior tail ridges. Spines
becoming progressively smaller in post-pelagic young and
remain as low rugose tubercles in large adults. Tubercles
enlarged above eyes, on trunk mainly below the dorsal fin,
and along shoulder ring at gill-opening with one of moderate
size ventrally. Lateral line : small pores with low papillae,
usually indistinct and only detectable intermittently
posteriorly to about 30th tail ring (in one female they are
clearly visible on nearly all rings to 36th tail ring). Coronet:
very low with rugose apex in adults. Colour in life : bony-
white, grey, yellow or brown, usually with black spots on
head and trunk. Those in shallow bays usually brown with
many dark spots, whilst those from deep water plain but
with brighter colours similar to living sponges. Colour in
alcohol: pale brown, plain to spotted on head and trunk
and sometimes with indistinct banding on tail.
Measurements. Largest specimen examined 165 mm in
height, but probably reaching 18 cm. Head small, ranging
from 62 (adult) to 77% (juvenile) of trunk. Snout usually
short, ranging 31-37% of head.
Distribution (Fig. 31). An east coast species, the most
northern record, based on photographs, is off Newcastle,
NSW (about 32°S), and the southern extent of the range is
Eden, NSW, but with pelagic young Hippocampus
abdominalis can be expected further south and possibly
overlaps in range with the southern form, H. bleekeri. Adults
occur in shallow estuaries, usually on low reef habitats with
kelp Ecklonia radiata or in deep tidal channels where they
usually cling to sponges, and coastal reefs where they are
usually in depths over 20 m.
Figure 30. Hippocampus abdominalis. A, 8 , AMS 115657-001, Ship Rock, Sydney, NSW. B, 9 , AMS 116791-
002, Long Bay, Sydney, NSW. C, juvenile, AMS 117670-001, Long Bay, Sydney, NSW.
Kuiter: Australian Hippocampus revision 323
Figure 31. Hippocampus abdominalis. Collection sites of
specimens examined and localities of specimens photographed
underwater.
Remarks. This species, featuring small head and short
snout, is tentatively identified as Hippocampus abdominalis,
a species that was originally described from New Zealand.
Hippocampus abdominalis differs from the very closely
related H. bleekeri in having a smaller head and much
shorter snout. It is also less spiny and lacks a nape spine.
The pectoral fin count in H. abdominalis is lower than in
specimens H. bleekeri from South Australia found at similar
latitudes. Adults usually lack dermal appendages on head
that are nearly always present in large H. bleekeri. However,
the absence or presence of such appendages may be habitat
related. To validate the identity of the New South Wales
population, the New Zealand populations of H. abdominalis
need to be studied, as there appears to be more than one
species there.
Material examined. AMS 113696, 9 , height 135 mm, Millers
Point, Sydney, 33°51'S 151°13'E, E. Briggs, 1916. AMS 115657-
001, 6 , height 165 mm, Ship Rock, Port Hacking, Sydney, 34°04'S
151°09'E, depth 10 m, Neville Coleman, 1 Mar 1970. AMS 116791-
002, $, height 150 mm, Long Bay, Sydney, 33°45'S 151°20'E, R.
Kuiter, 25 Nov 1972. AMS 117670-001, 9, height 90 mm, Long
Bay, Sydney, 33°45'S 151°20'E, R. Kuiter, 21 Jan 1974. AMS
121893-001, juvenile, Botany Bay, Sydney, 33°59'S 151°12'E,
NSW Fisheries, 1980. AMS IA5888, juvenile, Maroubra, Sydney,
33°57'S 151° 16'E, G. Bunt, 1933.
Hippocampus bleekeri
Southern Potbelly Seahorse
Fig. 33
Hippocampus (Macleayina) bleekeri Fowler, 1907. Victoria, Australia.
Hippocampus (Macleayina) agnesae Fowler, 1907. Victoria, Australia.
Hippocampus graciliformes McCulloch, 1911. Near Bass Strait,
Victoria, Australia.
Hippocampus abdominalis (non Lesson) Paxton et al., 1989: in part.
Hippocampus abdominalis (non Lesson) Lourie et al., 1999: in part.
Diagnosis. Dorsal-fin rays 27-30; pectoral-fin rays 14-16
(usually 15 in Victoria and 16 in South Australia); trunk
rings 12-13(usually 13); tail rings 44-48; subdorsal spines
5-6/0.0.3-4; head of small to moderate size, ranging from
73% (juvenile) to 80% (adult) of trunk; snout usually short,
40-50% of head; small nape spine present and adults usually
with long dermal filaments on various head spines. Fin rays:
dorsal 27-30; pectoral 14-16 (usually 15 in Victoria and
16 in South Australia). Rings: subdorsal 4-5+1-2; trunk
12-13 (usually 13); tail 44-48. Spines or tubercles:
subdorsal 5/0,0,1,1,1 or 6/0,0,1,1,1,1. Pelagic young with
angular spines of moderate length along trunk and anterior
tail ridges. Spines diminishing in relative size in post-pelagic
young, becoming low to moderately large rugose tubercles
in adults; tubercles raised at various points, above eyes,
along shoulder ring with upper tubercle at gill-opening and
moderately large lower tubercle positioned ventrally, and
those on the trunk mainly enlarged below dorsal fin. Small
nape spine present. Adults usually with long dermal
filaments on various head spines. Lateral line: pores usually
small, but distinct, visible intermittently on tail to about
25th tail ring. Coronet: Very low with rugose ridges and 2
blunt spines posteriorly in some adults. Colour in life:
usually grey or pale brown to yellow, occasionally bright
orange. Tasmanian population with numerous spots on head
and trunk; South Australian population with few black spots,
often mostly absent from head; and Victorian population
with variable number of black spots, usually most numerous
on head. Tail often with pale bands centred on about every
5th ring in dark individuals. Colour in alcohol: cream to
pale brown, retaining dark spots from life, though faded to
pale or dark brown.
Measurements. Largest specimen from Victoria examined,
female, 230 mm height, and South Australian, female, 225
mm height. Snout length 40-50% of head length, and head
length 73-80% of trunk length.
Distribution (Fig. 32). Known from the northern Great
Australian Bight, SA, to off Lake Entrance, Victoria and
the Derwent Estuary, Tasmania. It occurs in shallow
estuaries and reefs in coastal waters to a depth of at least 35
m. Usually seen clinging to holdfast of Ecklonia radiata
and sponges, but also man-made items that provide
anchorage in currents, ranging from robes and shopping
trolleys to shipwrecks.
Remarks. Hippocampus bleekeri is very similar to H.
abdominalis from New South Wales and New Zealand, and
Figure 32. Hippocampus bleekeri. Collections sites of specimens
examined and localities of specimens photographed underwater.
324
Records of the Australian Museum (2001) Vol. 53
Figure 33. Hippocampus bleekeri. A, 6, NMV A17231, Port Phillip Bay, Vic. B, 9 , NMV A2335, Wilsons
Promontory, Vic. C, 6, NMV A8904, Lakes Entrance, Vic. D-F, $ $, SAMA F5335, upper Spencer Gulf, SA.
differs from it in having a larger head and longer snout. In
contrast to H. abdominalis, whose snout shortens
proportionally with age to less than one-third head length,
the snout in H. bleekeri elongates with growth to half head
length. Hippocampus bleekeri has better developed spines
at all stages and commonly has long filaments extending
from its head spines. In the cooler waters of Tasmania this
species grows larger and usually has more fin-elements than
those from the mainland. This taxon is in need of further
study to determine if populations in the areas of South
Australia, Victoria and Tasmania represent a single species.
Material examined. SAMA F6221, $, height 225 mm, Douglas
Bank, Spencer Gulf, SA, D. Graske, 12 Nov 1985. SAMA F5335
(3), 9 9 , height 98-168 mm, Upper Spencer Gulf, SA, trawled
MRV NGRIN, Terry Sim, 8 Nov 1985. SAMA F3885, juvenile,
height 72 mm, northern Great Australian Bight, SA, S A Fisheries,
5 May 1973. NMV A2899, juvenile, height 46 mm, Port Phillip
Bay, Vic., in floating seagrass, R. Kuiter, 28 Apr 1983. NMV
A8904, d, height 122 mm, Lakes Entrance, Vic., 37°53'S
148°00'E, depth 32 m, W. French, September 1948. NMV 17231
(3), d,2 9 9, heights 200 & 250 mm respectively, Geelong, Vic.,
38°05.4'S 144°23.2'E, R. Ickeringill & T. Bardsley, Dec 1995.
NMV 17242 (2), 9 9 , heights 200-230 mm, Geelong, Vic.,
38°06.8'S 144°21.9'E, M. Lockett & M. Gomon, 1 Feb 1996.
Kuiter: Australian Hippocampus revision 325
Hippocampus zebra
Zebra Seahorse
Fig. 34
Hippocampus zebra Whitley, 1964. Swain Reefs, Queensland,
Australia.
Diagnosis. Dorsal-fin rays 17-18; pectoral-fin rays 15-16;
subdorsal rings 2-3+1; body and head covered by
alternating white and brown to black bands; coronet of
moderate height, with 5 spines at apex; upper shoulder-ring
spine near pectoral fin base. Fin rays’, dorsal 17-18; pectoral
15-16; anal 4. Rings : subdorsal 2-3 + 1; trunk 11; tail 37-
39. Spines or tubercles’, subdorsal 3-4/0,1,0-0.5. Spine
above eye of moderate length; nape spine small, directed
upward; nose profile straight; 3 small shoulder-ring
tubercles or spines, upper two as tubercles, situated at level
of either end of pectoral-fin base, none at gill-opening, and
lowermost as small spine; lateral head spine low; trunk
tubercles along dorsal and lateral ridges of small to moderate
size, some enlarged and pointed, especially beside dorsal
fin and on anterior part of superior tail ridges, becoming
progressively smaller posteriorly. Coronet’, of moderate
height, with 5 spines on apex, posterior 3 directed backward.
Colour in life’, (from Whitley, 1964): “yellowish-white and
dark brown bands, most of them transverse, but some
oblique or curved on the head and belly, the whole forming
a very beautiful and strongly contrasting arrangement with
a tendency for broad and narrow bands to alternate. Eye
blue with yellowish-white iris. Chin yellowish-white,
without dark bands. Fins lighter; dorsal and anal fins brown
proximally”. Colour in alcohol: as in life, with alternating
bands white to brownish-white and dark brown to black.
Measurements. Largest specimen examined with height
of 90 mm. Length of tail about 60% of height; trunk length
about 36% of height; head length about 74% of trunk length.
Males with deeper trunk than females, the depth in males is
about 80% and in females about 60% of head length. Length
of snout slightly less than half head length.
Figure 35. Hippocampus zebra. Collection sites of specimens
examined.
Distribution (Fig. 35). Only known from off the
Queensland coast and Milne Bay, southeastern Papua New
Guinea (based on photograph). Hippocampus zebra is
known from a depth range of about 20-60 m depth. Soft
bottom habitat, probably on black coral or gorgonians.
Remarks. Whitley’s paratype (AMS IB2819) of Hippo¬
campus zebra from Moreton Bay is a specimen of the
“zebra” form of H. dahli (identified as H. planifrons by
Johnson, 1999). Although the zebra-like colour pattern of
H. zebra is distinctive, the sympatric H. dahli can have
similar body patterns, though with finer stripes and a plain
snout. Hippocampus zebra also has a taller coronet, and
the upper tubercle on the shoulder ring is near the pectoral-
fin base rather than near the gill-opening as in H. dahli.
Material examined. AMS IB6015, X-ray of holotype, off Gillett
Cay, Swain Reefs, Qld, dredge 38 fm, Australian Museum 1962
Swain Reefs Expedition, 19 Oct 1962. QM 129845, d, height 58
mm, Cape Grenville, NE Qld, 11°55'S 143°55'E, trawl 20 m, S.
Cook, 21 Nov 1993. James Cook University, photographs of
unregistered specimens (2), d & 9, heights c. 70 mm, off
Townsville, trawl, M.L. Horne, 1999.
Figure 34. Hippocampus zebra. A, 9, holotype, AMS IB6015, Swain Reefs, Qld, photo by Anthony Healy, Australian
Museum. B, 9, James Cook University unregistered, off Townsville, Qld. C, 6 , unregistered, off Townsville, Qld.
D, S, QM 129845, Cape Grenville, Qld.
326 Records of the Australian Museum (2001) Vol. 53
Hippocampus montebelloensis n.sp.
Monte Bello Seahorse
Fig. 36
Type material. Holotype: NTM S10804, height 48 mm,
juvenile, off Trimoulle Island, Monte Bello Islands, Western
Australia, 20°24'S 115°34'E, H. Larson and party, 21 Apr
1983. Paratype: WAM P29078-001, height 78 mm, 9,
Exmouth Gulf, WA, 22°00'S 114°20'E, trawl Dorothea, 1
Oct 1962.
Diagnosis. Dorsal-fin rays 18-19; pectoral-fin rays 15-16;
trunk rings 11; tail rings 37; subdorsal spines 3/0,1,0; spine
above eye of moderate size, length about pupil-diameter,
slightly angled back; moderately large and recurving lateral
head spine; long, forward directed nape spine; nose-profile
straight; coronet moderately high, with spines on corners,
posterior 3 largest and directed backward; 3 spines of
moderate size on shoulder ring, uppermost and central
spines at ends of pectoral-fin base, lowermost ventrally;
superior trunk ridge sharp-edged with spines of small to
moderate size on rings 1, 4, 7 and 11, extended by long
dermal flaps at spine tips in holotype; superior tail ridge
similar to superior trunk ridge, with enlarged spines at
regular intervals, becoming progressively smaller
posteriorly.
Description. Head large, length 95% (90%) of trunk length;
snout long, 55% (50%) of head length; dorsal fin with 19
(18) rays, its base over 2 trunk and 1 tail ring; pectoral-fin
rays 15 (16); trunk rings 11; tail rings 37; moderately long
spine above eyes; tiny spine behind eye; large lateral head
spine, slightly recurving; long nape spine with filamented
tip, perpendicular to head profile; nose profile straight,
without spine, rising from snout at 45° angle; 2 low angular
spines below eye; 3 spines of moderate length on shoulder
ring, uppermost and central spines at ends of fin base,
lowermost positioned ventrally; coronet moderately high,
with 5 flat spines at apex, posterior 3 directed backward,
central spine enlarged; neck-ridge straight with 2 angular
spines, one above gill-opening and one at posterior end;
superior trunk ridge with moderately long spines, spines
enlarged with filament at tips on rings 1,4, 7 and 11; lateral
trunk ridge with series of spines from 2nd to 10th rings,
first small, progressively increasing in size to spines that
are of moderate length and blunt; inferior trunk ridge
moderately developed with downward directed blunt
angular spines; ventral trunk with low downward directed
spines; subdorsal spines 3/0,1,0; superior tail ridge with
enlarged spines and filamentous tips on every 3rd ring,
starting with 4th (5th in paratype), becoming progressively
shorter; inferior tail ridge continuous with trunk ridge,
tubercles becoming progressively smaller posteriorly; lateral
line present with pores detectable to about 14th tail ring.
Largest specimen, a female, 78 mm in height. Colour in
life : unknown. Colour in alcohol: mainly pale brown with
fine darker brown striations along the head.
Distribution (Fig. 37). Only known from Monte Bello Is
and Exmouth Gulf in Western Australia. Holotype caught
at surface at night, clinging to Sargassum fragment with
boat anchored in 5 m of water (Helen Larson, pers. comm.).
No other details recorded for paratype.
Remarks. This species is named montebelloensis in
reference to its type locality. Its closest relative appears to
be Hippocampus zebra, with which it shares the same
meristics and morphology. The superficially similar H.
alatus, occurring in the same region, is readily distinguished
from H. montebelloensis in having its upper shoulder-ring
spine near the gill-opening rather than near the pectoral-fin
base. It is likely that the paratype is close to the maximum
size of this species.
Figure 36. Hippocampus montebelloensis. A, juvenile, holotype, Figure 37. Hippocampus montebelloensis. Collection sites of
NTM S10804, Monte Bello Is, WA. B, 9, WAM P29078-001, specimens examined,
paratype, Exmouth Gulf, WA.
Kuiter: Australian Hippocampus revision 327
Hippocampus whitei
White’s Seahorse
Fig. 38
Hippocampus whitei Bleeker, 1855. Sydney, Australia.
Hippocampus novae Hollandiae Steindachner, 1866. Sydney,
Australia.
Diagnosis. Dorsal-fin rays 16-17 (usually 17); pectoral-
fin rays 15-17 (usually 16); tail rings 33-34; coronet height
moderate to tall, angled back, its spines small and blunt in
young and further reduced in large adults, arranged in five
pointed star at apex, with additional 2 or more small spines
anteriorly on base; neck-ridge spines absent. Fin rays : dorsal
16-17 (usually 17); pectoral 15-17 (usually 16); anal 4.
Rings: subdorsal 2 + 1; trunk 11; tail 33-34. Spines or
tubercles : subdorsal 3/0,1,0. Spine of moderate size above
eye; nape spine small; shoulder-ring spines of moderate size
in 3 locations, one near each end of pectoral-fin base but
none at gill-opening, and lowermost double with sub-equal
spines; lateral head spine low; body tubercles of small to
moderate size along dorsal and lateral ridges, some enlarged
and pointed, especially on 8th trunk ring, below dorsal fin
and on following part of superior tail ridges; neck-ridge
spines absent. Coronet : height moderate to tall, angled back,
its spines small and blunt in young and further reduced in
large adults, arranged in five pointed star at apex, with
additional 2 or more small spines anteriorly on base. Colour
in life: mostly pale to dark brown and almost black, with
fine pale spots or striations, and somet im es entirely yellow.
Often saddle-like markings at 1st, 4th and 8th trunk rings,
and on tail rings where spines are enlarged. Top of snout
finely barred with dusky lines, lines broader near eyes. Nose
spine and adjacent area pale. Colour in alcohol’, brown with
blackish-brown scribbles and blotches.
Measurements. Height of largest examined specimen 103
mm. Length of tail about 66% of height; trunk length about
Figure 39. Hippocampus whitei. Distribution based on specimens
examined and localities of specimens photographed underwater.
33% of height; head length about 90% of trunk length.
Length of snout about 42% of head length.
Distribution (Fig. 39). Apparently restricted to New South
Wales, though the northern extent of its range is uncertain.
Common in estuaries and harbours in Sydney and Newcastle
regions.
Remarks. No types of Hippocampus whitei are known.
Bleeker’s description is based on fig. 2 (plate opposite p.
264 in J. White, 1790) from New South Wales. White’s
figure shows a stylized seahorse with a distinct coronet with
at least three blunt spines (J.M. Leis, pers. comm.). Only
two seahorse species are known to occur in the vicinity of
Sydney: H. abdominalis and H. whitei. Hippocampus whitei
has a well-developed coronet with blunt spines whereas H.
abdominalis adults have no coronet. There can be no doubt
that White illustrated the most common seahorse species in
Sydney Harbour and Botany Bay, thus maintaining the
traditional usage of the name.
Figure 38. Hippocampus whitei. A, 8 AMS 119900-002, Sydney Harbour, NSW. B, 9, AMS 119900-002, Sydney
Harbour, NSW. C, juvenile, AMS 118809-001, Sydney Harbour, NSW.
328 Records of the Australian Museum (2001) Vol. 53
Many specimens in the various Australian institutions
registered as H. whitei represent other species: those from
northern New South Wales are H. tristis, those from southern
Queensland are H. tristis and H. procerus, while those from
Victoria and South Australia are juvenile H. breviceps.
Although specimens from South Australia of H. whitei were
not found in collections, the origin of the records from that
state was not determined. There is a slight possibility of its
existence in the upper Spencer Gulf (other Australian east
coast species have a disjunct distribution there). Hippo¬
campus whitei is distinguished from H. tristis and H.
breviceps in having its upper shoulder-ring spine near the
pectoral-fin base rather than near the gill-opening.
Hippocampus whitei is very similar to H. procerus, differing
from it in lower meristics, lower coronet and generally a
less spiny physiogamy.
Material examined. AMS 119900-002 (2), heights 94-96 mm,
Spit Bridge Marina, Sydney Harbour, 33°48'S 151°14'E,
Macquarie University, class, 2 m, 8 Aug 1976. AMS 118809-001,
height 65 mm, Pilot Station, Watsons Bay, Sydney Harbour,
33°50'S 151°16'E, 3 m, R. Kuiter, 20 Mar, 1976. AMS 119901-
027 (3), heights 75-95 mm, Neilsen Park, Sydney Harbour,
33°51'S 151°16'E, 5 m, Macquarie University, class, 8 Aug 1976.
AMS 138991-003, height 60 mm, Georges River, Botany Bay,
34°01'S 151°09'E, Lockett & Upston, 4 Mar 1998. AMS IA151,
height 103 mm, Domain Baths, Wooloomooloo, Sydney Harbour,
33°53'S 151°13'E, R. Gaul, 1921.
Hippocampus procerus n.sp.
High-crown Seahorse
Fig. 40
Hippocampus whitei (non Bleeker) Paxton et al., 1989: in part.
Hippocampus whitei (non Bleeker) Johnson, 1999.
Hippocampus whitei (non Bleeker) Lourie et al., 1999: in part.
Type material. Holotype: AMS E2914, 9 , height 110 mm,
5-11 km NW Hervey Bay, Fairway Buoy, Queensland, 24°S
152°E, FIS Endeavour, 7 Jul 1910. Paratypes: QM12208,
juvenile, height 55 mm, Norman River, Gulf of Carpentaria,
Figure 40. Hippocampus procerus. A, S , paratype, QM ]
Hervey Bay, Qld. C, $, paratype, QM 130772, Moreton
$, paratype, AMS IA4205, Port Curtis, Qld.
Qld, C. Taylor, 3 Nov 1914. QM 18792, <$, height 98 mm,
off Redcliffe, SE Qld, N. Phillips, 24 Jul 1965. QM 111000
(7), heights 56-96 mm, Moreton Bay, Qld, T.C. Marshall,
24 Oct 1950. QM 113012, 6, height 75 mm, off Redcliffe,
SE Qld, trawl 5 fm, R.J. McKay, 17 Mar 1975. QM 129963,
9, height 65 mm, Newport Canal mouth, Deception Bay,
Moreton Bay, Qld, 27°12’S 153°06’E, trawl, I. Halliday, 1991.
QM 130772, $ & S, heights 75 & 95 mm respectively, off
Chain Banks, Moreton Bay, Qld, J. Johnson, depth 3-7 m,
24 Jan 1997. AMS 1492, 9 , height 94 mm, Moreton Bay,
Qld, 27°25'S 153°20'E, J.D. Ogilby, 1886. AMS 112554,
S, height 105 mm, Moreton Bay, Qld, 27°00’S 153°00’E,
Amateur Fishermans Association of Qld, 1912. AMS IA4205,
juvenile, height 57 mm, Port Curtis, Qld, 23°55'S 151°23'E,
dredged, M. Ward & W. Boardman, 14 Dec 1929. AMS
IB 1011, 9 , height 105 mm, east of Burnett River, Qld, 24°S
152°E, dredged, CSIRO, 14 Sep 1938.
Diagnosis. Dorsal-fin rays 17-19 (rarely 17); pectoral-fin
rays 16-18 (rarely 16, usually 18); trunk rings 11; tail rings
34-35; subdorsal spines 3/0-1,1,0-1; spines of moderate
size at nose, above eyes, and laterally on head; coronet tall
and strongly angled back, with 5 large spines at apex and
additional spines on anterior face; neck ridge with low
spines, one centrally and one at posterior end.
Description. Head long, 85% of trunk length; snout long,
47% of head length; dorsal-fin rays 18 (17-19, one paratype
with 17), its base over 2 trunk and 1 tail ring; pectoral-fin
rays 18 (16-18, rarely 16, usually 18); trunk rings 11; tail
rings 35 (34-35); spine with length of pupil diameter above
eyes; lateral head spine of moderate length, directed laterally
outwards; small spine perpendicular to nape; spine of
moderate length behind eye; 2 separate low spines below
eye; shoulder-ring spines of moderate length, uppermost
near top of pectoral fin base, central spine near bottom of
pectoral fin base, and lowermost double and positioned
ventrally, the more anterior of the two small and less
developed (absent in some paratypes on one or both sides);
coronet tall, angled backward with 5 flanged and pointed
spines on apex in star-like arrangement, and small spines
<-
C □ E
8792, off Redcliffe, SE Qld. B, $, holotype, AMS E2914,
Bay, Qld. D young d, QM 130772, Moreton Bay, Qld. E,
Kuiter: Australian Hippocampus revision 329
Figure 41. Hippocampus procerus. Collection sites of specimens
examined.
anteriorly near base; neck ridge with low spines, one
centrally and one at posterior end; superior trunk ridge with
enlarged blunt spines on 1st, and from 7th ring on; lateral
trunk ridge with spines on 2nd to 10th rings, progressively
increasing in size; inferior trunk ridge with angular and
downward directed spines on last few rings; ventral trunk
ridge with few downward directed spines, those on last 2
rings enlarged; subdorsal spines 3/0,1,0; superior tail ridge
spines well developed anteriorly, except on first ring,
progressively diminishing in size posteriorly; inferior tail
ridge continuous with inferior trunk ridge, with 6 spines
progressively diminishing in size posteriorly; lateral line
with pores detectable to about 19th tail ring. Colour in life :
(based on colour transparency taken by B. Cowell of a
female specimen from Moreton Bay, 8 cm in height) trunk
pale yellowish white ventrally, dusky brown with black
scribbles and numerous tiny white spots dorsally above
lateral trunk ridge, and continuing over tail; snout dusky
with irregular thin white barring; brownish black shading
ventrally on eye and snout; dorsal fin with a dusky
longitudinal line at mid-level. Colour in alcohol : plain
brown with fine dark scribbles or pale saddles.
Distribution (Fig. 41). Southern Queensland from Hervey
Bay to Moreton Bay, with a single record from Gulf of
Carpentaria which may be erroneous (Jeff Johnson, pers.
comm.). Occurs on mixed algae reef to depths of about 20 m.
Remarks. This species is named procerus , from the Latin
for “tall” in reference to the tall coronet. Previously confused
with Hippocampus tristis and H. whitei, H. procerus is more
similar to the latter, differing from it in having a taller and
spinier coronet, higher fin-ray counts, and generally a spinier
physiogamy.
Hippocampus elongatus
West Australian Seahorse
Fig. 42
Hippocampus elongatus Castelnau, 1873: 144. Fremantle, WA.
Hippocampus subelongatus Castelnau, 1873: 145. Fremantle, WA.
Hippocampus angustus (non Gunther) Whitley & Allan, 1958:
in part.
Hippocampus angustus (non Gunther) Paxton etal., 1989: in part.
Hippocampus subelongatus Lourie et al., 1999.
Diagnosis. Dorsal-fin rays 18; pectoral-fin rays 17-18; trunk
rings 11; tail rings 33-34; tall coronet, crown with 5 spines
Figure 42. Hippocampus elongatus. A, 6, AMS 120228-004, Woodman Point, Cockburn Sound, WA. B, 9, AMS
120228-004, Woodman Point, Cockburn Sound, WA. C juvenile, AMS 120228-004, Woodman Point, Cockburn
Sound, WA.
330 Records of the Australian Museum (2001) Vol. 53
in star arrangement when young, spines becoming blunt
and forming rounded flanges in large adults; spines on trunk
and tail of moderate length in juveniles, becoming blunt
with growth, appearing as low tubercles in adults; nose spine
angular. Fin rays : dorsal 18; pectoral 17-18; anal 4. Rings :
subdorsal 2+1; trunk 11; tail 33-34. Spines or tubercles :
subdorsal 3/0,1,0-0.5. Spines of moderate length in
juveniles, becoming blunt with growth and appearing as
low tubercles in adults. Enlarged tubercles above eyes,
laterally on head, and sub-dorsally on trunk. Nape spine
long when young. Nose spine angular. Shoulder-ring with
3 blunt spines, uppermost just above top of pectoral-fin base,
central spine at level of last few rays of pectoral-fin base
and lowermost double, in form of short laterally-directed,
diverging spines. Coronet : tall, crown of juvenile with 5
diverging spines in star arrangement, spines becoming blunt
with growth forming rounded flanges in large adults. Lateral
line : indistinct, pores extending to 22nd tail ring. Colour in
life : highly variable from grey, yellow, brown to burgundy-
red, usually with thin dark barring on snout; body plain or
with dusky striations or reticulations. Colour in alcohol :
pale brown, usually retains features of patterns in live
individuals.
Measurements. Height of largest examined, female, 145
mm, but known to reach a height of about 20 cm. Head
long, about 85-90% of trunk length and snout about 1/2
head length. Trunk slender in both sexes, depth 38-46% of
its length.
Distribution (Fig. 43). Recorded between Cape Naturalist
and Kalbarri in Western Australia, where it inhabits shallow
algae and sponge reefs in sheltered bays.
Remarks. Hippocampus elongatus is a common species in
the Perth region, occurring seasonally in the lower reaches
of Swan River. It is replaced by more spiny species from
Shark Bay northwards. Castelnau (1873) described H.
elongatus and H. subelongatus, each on the basis of a
single specimen sent from Fremantle, WA. Castelnau was
not sure about the validity of H. subelongatus thinking
that it might be the same as H. elongatus. Fourie et al.,
1999 chose to recognise the name H. subelongatus for this
species, despite H. elongatus being the first name published
Figure 43. Hippocampus elongatus. Collection sites of specimens
examined and localities of specimens photographed underwater.
and subelongatus being a derivative of elongatus, giving
the presence of 3 “paratypes” of H. subelongatus and the
absence of the type of H. elongatus in the collection of the
Paris museum (Bertin & Esteve, 1950) as the reason. As
Castelnau had only a single type of H. subelongatus, it is
likely that the second largest specimen regarded as a syntype
by Bertin & Esteve is the type of H. elongatus as suggested
by the following evidence. Castelnau stated that the length
of the H. elongatus type is “three inches [76 mm] in a
straight line”. Although this may be interpreted as a total
length measurement, it may just as easily be a “straight-
line” measurement from the highest to the lowest parts of
the dry specimen along its axis, regardless of the body or
the tail shape. The total length of 76 mm is that of an
extremely small specimen, but if the measurement is taken
as a straight-line along the axis from the highest to the lowest
parts of the specimen, it corresponds exactly with one of
the “paratypes” of H. subelongatus (MNHN A4536).
Castelnau’s description of H. elongatus not only agrees with
this specimen but is consistent with other similar-sized
specimens examined. The MNHN A4536 specimen is 76
mm in a straight line measurement and 140 mm in total
length. Castelnau’s description of H. subelongatus agrees
with the largest of the “paratypes” (MNHN A4535) which
is 124 mm in a straight line measurement and 175 mm total
length. The smallest “paratype” of H. subelongatus (MNHN
A4552), 85 mm in a straight line measurement and 108
mm in total length, represents this species but disagrees
with Castelnau’s descriptions. Hippocampus elongatus has
page priority, and from Castelnau’s statement that
subelongatus “may be a more developed age of Elongatu x”
it clearly would have been his choice. Therefore,
Hippocampus elongatus is here reinstated as the senior
synonym for this species.
Material examined. MNHN A4535, probable type of H.
subelongatus, height 132 mm, photograph, Fremantle, WA.
MNHN A4536, probable type of H. elongatus, height 100 mm,
photograph, Fremantle, WA. MNHN A4552, height 93 mm,
photograph, Fremantle, WA. AMS 120228-004 (3), 6, height 135
mm; 9, height 145 mm; juvenile, height 94 mm; all Woodman
Point, Cockburn Sound, WA, 32°08'S 115°45'E, depth 3-8 m, B.C.
Russell & R. Kuiter, 25 Mar 1978. AMS 126903-002, 9, height
145 mm, Palm Beach Cockburn Sound, WA, 32°H'S 115°43'E,
depth 4-12 m, collector not registered, 25 Jul 1971. AMS 120350-
GO 1, juvenile, height 96 mm, Sulphur Rock Cockburn Sound, WA,
32°11'S 115°43'E, depth 18 m, N. Coleman, 16 Jan 1972.
Hippocampus spinosissimus
Hedgehog Seahorse
Fig. 44
The following is my translation of Weber’s (1913) original
description in German (my additions in italics and within
parentheses):
Hippocampus spinosissimus n.sp. fig. 44. Station
49. Sapeh Strait. 70 m; sand and scallops. 2 specimens
c. 70 mm.
Dorsal fin 17; pectoral fin 15. Trunk 11 and tail 34
rings. Head 1.5x trunk-length and 5.8x total length.
The trunk 2 times tail length. The snout as long as the
post-orbital part of the head, and twice as long as the
eye-diameter, and the least depth equal to 3/4 of the
Kuiter: Australian Hippocampus revision 331
Figure 44. Weber’s fig. 44 (in part) of Hippocampus spinosissimus
from Sapeh Strait (d).
eye-diameter. The brood-pouch is situated below the
first 6 tail rings. The dorsal fin is positioned over the
last 2 trunk rings and first 2 tail rings. The coronet is
c. 1/2-2/3 as long as the snout; and considerably
higher and thicker cylindrical with the 5 sharp
diverging spines, of which the frontal 2 are not slanting
backwards like those behind. At its base lies a vertical,
moderate spine ( nape-spine ). Above the eyes, large
sharp spines, equal in length to the eye diameter,
directed upwards and out. In front between the eyes,
a single forward directed spine {nose-spine). Two
postorbital spines in series; two at the bases of the
pectoral fins and 3 others along the ridge below the
eye (1) and throat (2, the double lower shoulder-ring
spine). The spines on the trunk and following half of
the tail, sharp, enlarged and more protruding on the
1st, 4th, 7th, 11th, trunk rings, and the 4th, 7th and
11th tail rings. The colour in alcohol is plain yellow-
brown with white spots on the head; a submarginal
black stripe in the dorsal fin.
The measurements of the specimens: total length
70 mm, head 12 mm, trunk 18 mm, tail c. 40 mm,
snout 5 mm, eye just under 2.5 mm.
This new species is closely related to H. erinaceus
Gunther ( -H. angustus Gunther) and who knows,
perhaps even identical, but I cannot determine this
from the Kiirze von Gunther diagnosis. In any case,
the statement “Breast-spines double on each side”
disagrees with my specimens, and it does not have 17
rays in the dorsal fin. In 1893 I made a similar
conclusion during the work on fish for Semon’s (Max
Weber. Fische von Ambon etc., in Semon’s Zool.
Forschungsreisen in Australien etc., p. 275.). At that
time I listed a specimen from Thursday I., Torres Strait,
#193 as Hippocampus {erinaceus Gthr.?). This
specimen agrees with the above described examples
of H. spinosissimus, only the median ridge on the trunk
is so deeply shaped with rounded edges, it obscures
the diagnostic spines.
Remarks on the Max Weber description
A later description in English of Hippocampus spinosissimus
in Weber & de Beaufort (1922) is short and was modified
considerably from the original. The caption with Weber &
de Beaufort’s figure is incomplete and misleading. They
assumed that the specimen from Thursday I. was conspecific
with the types from Sapeh Strait. In the original description,
a fish from Sapeh Strait was illustrated in fig. 44 (reproduced
here as Fig. 44), but it was accompanied by a second
illustration showing the facial part of the head based on the
specimen from Thursday I. This is clear in the German
caption, but not mentioned in the English version by Weber
& de Beaufort. The Thursday I. specimen has similar spines
on the head but as Weber stated, the specimen does not
agree with the specimens from Sapeh Strait. The specimen
from Thursday I. has cross bands on the snout which were
mentioned in the English description, but no cross-bands
on the snout are mentioned in the German description of H.
spinosissimus. The specimen from Thursday I. appears to
be H. grandiceps. Reports of H. spinosissimus from
elsewhere are based on a variety of species with distinct
spines. No Australian species match the description and
illustration of H. spinosissimus, especially among those
with 17 dorsal-fin rays and 15 pectoral-fin rays. Spines on
the superior trunk ridges are long in males of H.
spinosissimus, which, except for H. multispinus, are short
in males of Australian species. Lourie et al. (1999) report
H. spinosissimus from various regions in the western Pacific,
including Australia. I have examined material from
Indonesia and the Philippines that match the Lourie et al.
(1999) description of H. spinosissimus, and conclude that
these are not H. spinosissimus, but other species such as H.
moluccensis. The types from Sapeh Strait are the only
known specimens of H. spinosissimus, and its distribution
appears to be limited to the Komodo region
Hippocampus multispinus n.sp.
Northern Spiny Seahorse
Fig. 45
Hippocampus angustus (non Gunther) Paxton etal., 1989: in part.
Hippocampus angustus (non Gunther) Lourie et al., 1999: in part.
Type material. Holotype: NTM S12955-009, 9, height
105 mm, Arafura Sea, Northern Territory, 10°49'S 135°07'E,
depth 54 m, H. Larson, 25 Oct 1990. Paratypes: NTM
S12611-012, $, height 140 mm, Arafura Sea, NT, 10°26’S
136°24'E, depth 56-57 m, W. Houston, 8 Mar 1985. NTM
S13917-001, S, height 110 mm, Elizabeth River, Darwin
Harbour, NT, 12°32.0’S 130°56.3’E, depth 12 m, R.
Williams & H. Larson, 23 Nov 1991. NTM S13257-001,
9, height 100 mm, Gulf of Carpentaria, NT, 13°03'S
136°45.'E, depth 22 m, R. Williams, 23 Nov 1991. NTM
S13326-003, 9, height 98 mm, west of Shepparton Shoal,
Timor Sea, NT, 12°13’S 129°43'E, depth 46 m, NT Fisheries,
6 Dec 1990. NTM S13974-006, 9, height 94 mm, N of
Dampier Archipelago, NT, 20°13’S 116°18'E, L. Bullard,
11 May 1983. NTM S13541-001, juvenile, height 50 mm,
Arafura Sea, NT, 10°28.9'S 134°11.EE, depth 59-60 m, R.
Williams, 26 Sep 1992. CSIRO B3594, NW of Dampier
Archipelago, WA, 20°26.7-25.9’S 116°19.1-20.1’E,
demersal trawl, FRY Soela, CSIRO, 15 Oct 1983.
332 Records of the Australian Museum (2001) Vol. 53
o
D
Figure 45. Hippocampus multispinus. A, 9, holotype, NTM S12955-009, Arafura Sea, NT. B, 9, paratype, NTM
S13326-003, Arafura Sea, NT. C, S, paratype, NTM S13917-001, Darwin Harbour, NT. D, juvenile, paratype,
NTM SI3541-001, Arafura Sea, NT.
Diagnosis. Dorsal-fin rays 18; pectoral-fin rays 16-18
(usually 17); trunk rings 11; tail rings 30-35; spines long
and sharp, spine above eye equal in length or longer than
eye-diameter, perpendicular to snout; nose-spine sharp,
angled forward 45° to snout, anterior spines on superior
tail ridge longer than spaces between them.
Description. Head moderately long, 73% trunk length;
snout long, just over half head length; dorsal fin with 18
rays, base over 2 trunk and 1 tail ring; pectoral-fin rays 17
(16-18, usually 17); trunk rings 11; tail rings 35 (30-35).
Spines long and sharp; subdorsal spines 3/0,1,0; length of
spine above eye equal to eye-diameter, perpendicular to
snout; nose spine moderately long, angled forward 45° to
snout; nape spine long, perpendicular to nape; long spine
behind eye; double spines below eye; shoulder-ring with 3
long spines, uppermost just above level of pectoral-fin base,
central spine at level of last few rays of pectoral-fin base,
and lowermost double, in form of laterally directed,
diverging spines; lateral head spine long; coronet with 5
long diverging spines; neck ridge with 2 spines, one
centrally and one at posterior end; superior trunk ridge with
long spines, slightly enlarged at regular intervals from 1st
ring to below dorsal-fin base, length of some about equal
to eye-diameter; lateral ridge with long spines but spines
absent from rings 1, 3, 5 and 11; inferior trunk ridge with
series of spines from 6th to 11th ring, ranging in length
from short to long; ventral trunk ridge with spines on last 5
rings, angled posteriorly on last two; superior tail ring spines
long, those anterior spines much longer than longitudinal
distances between adjacent spines. Lateral line indistinct,
comprising small pores each with a single papilla,
intermittently detectable on tail to 20th ring. Height of
largest specimens 140 mm. Colour in life : (based on a colour
transparency of the paratype CSIRO B3594 from off the
Dampier Archipelago) evenly dark brown from rear of head
to end of tail; head white with few brown spots radiating
from eye; whitish bands on trunk rings 1, 4, 6, 8 and 11;
spines mostly white around bases, the tips black. Helen
Larson sent an aquarium photograph of a live female from
Darwin Harbour that is an orange-gold colour with few
small white spots scattered over the head and faint dark
bars on the snout. Colour in alcohol : pale brown without
markings, sometimes with few faint dusky bars on snout.
Distribution (Fig. 46). Apparently widespread in northern
waters from the Dampier Archipelago to southern Papua
New Guinea. A photograph of this species was taken by
Bob Halstead in Milne Bay, PNG, clinging to soft coral in a
strong current-prone habitat at a depth of 25 m. Most
specimens trawled at depths between 20 and 60 m, although
one paratype came from 12 m in Darwin Harbour.
Figure 46. Hippocampus multispinus. Collection sites of
specimens examined.
Kuiter: Australian Hippocampus revision 333
Remarks. This species is named multispinus from the
Latin multus (numerous) and spinus (thorny or spiny),
in reference to the numerous spines over the head and
body. Hippocampus multispinus has been confused with
other species with prominent spines, including H. histrix
and H. spinosissimus. Hippocampus multispinus is readily
distinguished from H. histrix in having the long double
lower shoulder-ring spines, that diverge from one another
in a V-shape, rather than the single long spine of H. histrix,
and from H. spinosissimus in having a long snout that is
longer than half the head length, in contrast to well short of
half the head length in H. spinosissimus. Hippocampus
multispinus is similar in morphology to H. hendriki, but
the latter has shorter spines in adults, lacks most spines on
the superior trunk ridges in males, and has saddle-like
markings which are missing in H. multispinus.
Hippocampus histrix
Thorny Seahorse
Fig. 47
Hippocampus histrix Kaup, 1856. Japan.
Diagnosis. Dorsal fin rays 17-19; pectoral-fin rays 17-18;
tail rings 33-34; spines long and sharp; nape spine long,
directed upward, placed close to coronet; lower shoulder¬
ring spine single, long, directed laterally and slightly
forward. Fin rays: (after Nakabo, 1993) dorsal 17-19;
pectoral 17-18; anal 4. Rings: subdorsal 2 + 1; trunk 11;
tail 33-34. Spines or tubercles: subdorsal 3/0,1,0. Spines
long and sharp; length of spine above eye about equal to
eye-diameter; nose spine slightly shorter and nape spine
slightly longer than eye spine; 2 sharp neck spines, similar
to coronet spines in length; trunk ridge spines on nearly all
trunk and tail junctions, only absent on 1st ring of lateral
ridge, starting on 4th inferior trunk ridge, and a single sub¬
dorsal spine on tail ridge. Coronet: of moderate height, but
usually enlarged with 5 long diverging spines on apex. Nape
spine immediately preceding and often reaching as coronet.
Lateral line: not detectable from photographs. Colour in
life: highly variable from grey, greenish, yellow, or brown
to burgundy-red; usually one with or several thin white bars
on snout; body plain or with pale saddle-like markings; tips
of spines often black. Colour in alcohol: Not examined.
Measurements. Height of largest specimen reported 15 cm.
Distribution. Ranging from Japan to Bali, Indonesia, along
Wallace’s Line, and to New Caledonia in the Coral Sea
(based on photographs taken by the author and others).
Occurs on deep coastal slopes, primarily in depths over 15
m on soft bottom, in areas exposed to strong tidal currents.
Usually found clinging to soft corals, sponges or
occasionally with Halimeda algae.
Remarks. There are no specimens of Hippocampus histrix
in Australian collections and no photographs of specimens
taken in Australian waters were found. The extensive
geographical range suggests that young of this species may
be pelagic. Hippocampus histrix is included here because
of previous (incorrect) reports of Australian occurrence, and
the likelihood that the species will be found in Australia.
Figure 47. Hippocampus histrix. Osezaki, Izu Peninsula, Japan.
Photograph taken by Hiroyuki Uchiyama.
Hippocampus hendriki n.sp.
Eastern Spiny Seahorse
Fig. 48
Hippocampus angustus (non Gunther) Paxton etal., 1989: in part.
Hippocampus sp. 1 Kuiter, 2000, p. 34.
Type material. Holotype: AMS 120923-002, S, height
100 mm, Cape York, 10°39'S 142°30'E, prawn trawl, depth
16-18 m, AIMS AMS QM, 15 Feb 1979. Paratypes: QM
115837, <3, height 95 mm, west of Adolphus Passage, N
Qld, 10°38'S 142°28'E, trawled 9-10 fm, AIMS AM QM,
15 Feb 1979. QM 111772 (2), $ & c3, heights 104 & 96
mm, off Cairns, Qld. CSIRO H3639-03, E Newcastle Bay,
Qld, 10°52.8'S 142°58.6'E, Florida Flyer prawn net, depth
20 m, G. Yearsly & P. Graham (CSIRO), FRV Surveyor, 26
Nov 1993. AMS IA6752, juvenile, height 47 mm, off
Lindeman I., Qld, 20°27'S 149°02'E, trawled, G.P. Whitley,
27 Jul 1935.
Other material examined: NTM S13273-019, juvenile, Gulf of
Carpentaria, Qld.
Diagnosis. Dorsal-fin rays 17-18; pectoral-fin rays 16-17;
trunk rings 11; tail rings 34; subdorsal spines 3/0,1,0; nose
spine of moderate length, angular in shape; spine above
eye long, but shorter than eye-diameter; spine behind eye
of moderate size, followed by longer lateral head spine;
coronet moderately high, sloping back in line with head
profile; superior trunk ridge spines reduced to tubercles
before dorsal fin in males, or entire ridge having irregular
rugose parts with small spines on and in between junctions
with rings.
334 Records of the Australian Museum (2001) Vol. 53
Figure 48. Hippocampus hendriki. A, 6 , holotype, AMS 120923-002, Cape York, Qld. B, 6 , paratype, QM115837,
Cape York, Qld. C, 9, paratype, QM 111772, off Cairns, Qld. D, juvenile, paratype, AMS IA6752, Lindeman I.,
Qld. E juvenile, NTM SI3273-019, Gulf of Carpentaria, Qld.
Description. Head large, length 85% of trunk length; snout
moderately long, about equal to postorbital length; dorsal
fin with 18 rays (17 in one of four paratypes), base over 2
trunk and 1 tail ring; pectoral-fin rays 16 on one side and
17 on other side (16-17 in paratypes); trunk rings 11; tail
rings 34; moderately long spine above eyes, length about
70% of eye diameter; spine of similar size behind eye;
slightly larger lateral head spine, directed laterally; nape
spine of moderate length perpendicular to nape; 2 separate
angular spines below eye; 3 spines of moderate length to
long on shoulder ring, uppermost just above pectoral-fin
base, central spine at lower level of pectoral-fin base, and
lowermost as a double spine ventrally, with one portion
directed forward and other directed laterally; coronet raised
and angled back matching angle of head profile, with 5-
spined, star-like crown, and double spine posteriorly below
crown on side; neck ridge even with rugose edge (with 2
spines in small paratypes); superior trunk ridge with blunt
tubercles to 8th ring, followed by spines of moderate length
to end of dorsal fin base; lateral trunk ridge with spines on
2nd, 4th, and 6-10th rings; inferior trunk ridge with spines
to 10th ring, followed by tubercles on remaining rings;
ventral trunk with ridge of skin (with spiny in juvenile and
female paratypes); subdorsal spines 3/0,1,0; superior tail
ridge with enlarged spines on 2nd and 4th to 10th rings,
becoming progressively shorter posteriorly, followed by
smaller tubercles that similarly become progressively
smaller; inferior tail ridge with tubercles that are continuous
with those on trunk ridge to 9th ring, becoming progress¬
ively smaller posteriorly; lateral line with pores detectable
to about 20th tail ring. Height of largest specimen 104 mm.
Colour in life : (based on photographed of female, CSIRO
H3639-03) pale yellow on snout, lower part of head and
front of trunk, darkening to pale orange on top and back
with dark saddle-like markings over back at 1st, 4th, and
8th trunk rings, small saddle below dorsal fin, and others
on tail on about 4th and 7th rings; top of snout with about
10 thin dusky bars; dark blotch on head below gill-opening;
eye with brown vertical bar, extending to spine above it; tips
of spines black; dorsal fin with thin longitudinal lines. Colour
in alcohol : pale to dark brown, sometimes with pale saddle-
like markings at first and eight trunk rings, as well as on tail,
becoming bands posteriorly; snout with thin dark barring.
Distribution (Fig. 49). Apparently restricted to inner Great
Barrier Reef area, from the Capricorn region to Cape York,
Queensland. Habitat unknown. All specimens trawled on
flat substrate at depths of 18 to 25 m.
Figure 49. Hippocampus hendriki. Collection sites of specimens
examined.
Kuiter: Australian Hippocampus revision 335
Remarks. This species is named hendriki for Hendrik Kuiter
in recognition of his keen interest in seahorses that he
successfully conveyed to classmates and teachers.
Hippocampus hendriki has been confused with other species
with prominent spines, including H. angustus, H.
multispinus, and H. grandiceps. Hippocampus angustus has
lateral-ridge spines on the 3rd and 5th rings that are absent
in H. hendriki as well as H. multispinus . Hippocampus
hendriki is distinguished from H. multispinus in having
shorter spines above the eyes, the length obviously shorter
than eye-diameter, rather than longer than the eye-diameter.
Hippocampus grandiceps has a longer head, which is 94%
to over 100% (versus 85% or less) of trunk-length, and holds
its head close to the trunk, whilst the head of H. hendriki is
held at about 90° to the trunk. Males can look superficially
similar to H. queenslandicus which has different fin counts
and a tail with less spine development, but H. hendriki is
readily distinguished from that species by its nose spine
and the barring on its snout. Males of H. hendriki have long
spines on ridges near the pouch region, but lack them on
the superior trunk ridge anteriorly. Small juveniles have
proportionally longer spines and disproportionally
enlarged spines featuring dermal flaps near their tips. In
all specimens examined, the head is maintained at right
angle to the body, which may be an adaptation to living on
open substrates. This is consistent with my observations of
other syngnathids, including seahorses and pipefishes, that
live out in the open and usually position themselves almost
horizontally on the bottom to feed, keeping their head
forward. This contrasts with species that cling to weed and
maintain the body almost vertically while feeding, keeping
their head close to their chest.
Hippocampus angustus
Western Spiny Seahorse
Fig. 50
Hippocampus angustus Gunther, 1870: 200. Shark Bay, Western
Australia.
Hippocampus erinaceus Gunther, 1870: 206. No locality.
Diagnosis. Dorsal-fin rays 18-19; pectoral-fin rays 15-20
(usually 17, rarely 15, 19 or 20); trunk rings 11; tail rings
31-32; trunk slender, its depth equal to or less than snout
length; length of spine above eye about pupil diameter;
double lower shoulder-ring spines; coronet with 5 sharp
spines diverging from apex; snout with thin dusky bars;
dorsal with a thin dusky line at margin. Fin rays: dorsal
18-19; pectoral 15-20 (usually 17, rarely 15, 19 or 20);
anal 4. Rings : subdorsal 2+1; trunk 11; tail 31-32. Spines
or tubercles : Subdorsal 3/0,1,0; length of spine above eye
equal to pupil diameter, perpendicular to snout; nose spine
small, directed forward; nape spine small, directed slightly
forward; small spine behind eye; double spines below eye;
3 shoulder ring spines, upper two at either end of pectoral-
fin base, lowermost double, in form of laterally directed,
diverging spines, anterior one often small; lateral head spine
of moderate size to long; superior trunk ridge with tubercles
of moderate size in males and short but sharp spines in
females; lateral trunk ridge with spines of moderate size on
2nd to 10th rings; inferior trunk ridge with thick downward
directed tubercles in males and spines of moderate length
in females; ventral trunk ridge expanded forward as
scalloped ridge in males, but bearing downward directed
Figure 50. Hippocampus angustus. A, 6 , WAM P27351-001, Shark Bay, WA. B, 9, WAM P8174, Shark Bay, WA.
C, juvenile, WAM P29077-001, Shark Bay, WA.
336 Records of the Australian Museum (2001) Vol. 53
angular spines in females; superior tail ridge with spines of
moderate length, though usually none on 1st ring; inferior
trunk ridge with tubercles of moderate size or small spines,
spines usually enlarged along pouch in males. Lateral line :
small pores, more or less distinct, extending to 20th tail
ring, pores difficult to detect posteriorly. Coronet :
moderately high with 5 diverging sharp spines on apex.
Colour in life : grey to brown, often covered with fine dark
scribbles; snout with 5-6 vertical dark bands; dorsal fin
with thin dusky marginal line. Colour in alcohol : Pale brown
with thin dusky bars on snout and scribble markings on
head and body.
Measurements. Height of largest specimens examined 149
mm. Head length about 80% in trunk length and snout of adults
long, nearly half head length (45.8-48.2%), but short in
juveniles (37.8% in 36 mm specimen). Trunk slender, its depth
usually about equal to or less than snout length in all stages.
Distribution (Fig. 51). Apparently restricted to Western
Australia, from Shark Bay to the Dampier Archipelago,
where it overlaps geographically with H. multispinus.
Remarks. This is one of several similar species with
prominent spines living in the tropics that can be difficult
to distinguish from one and another. It has been confused
with Hippocampus histrix, H. spinosissimus and H.
multispinus. It is readily distinguished from H. histrix in
having a double lowermost spine on the shoulder-ring rather
than a single. In addition, H. histrix has much longer spines
on the head. Hippocampus angustus differs from H.
spinosissimus in having a much longer snout, which is nearly
half the head-length in adults, in contrast to much less than
half the head-length. It can be distinguished from H.
multispinus by its shorter spines, the length of the spine
above the eye about the diameter of the pupil rather than
the diameter of the eye. Hippocampus angustus was
confused with several other species, including H. elongatus
(incorrect synonym), H. grandiceps, H. hendriki, and H.
queenslandicus, for which no names were available until
now. The name was applied loosely to species with
moderately long spines, while H. histrix was used for species
with very long spines, such as H. multispinus and the
juveniles of species that have relatively longer spines than
adults. The inclusion of Hippocampus erinaceus as a
Figure 51. Hippocampus angustus. Collection sites of specimens
examined.
synonym of H. angustus is uncertain. The type locality is
unknown and the specimen appears to be a juvenile H.
angustus. Suggestions that the type locality is “possibly NE
Australia” may have come from Weber’s use of the name
Hippocampus ( erinaceus Gthr.?) when listing a specimen
that originated from Thursday I., Torres Strait, Australia. It
seems that the specimen of H. erinaceus originated from
Shark Bay, since Gunther’s material described in the 1870s
from Australia apparently came from NSW and the Perth
region of WA. Lourie et al. (1999) commented that “The
name angustus is here used in preference to erinaceus
because of its known type locality”. If these names refer to
the same species, Hippocampus angustus is also preferred
as the senior synonym, based on page priority.
Material examined. WAM P5871 (2), 8 8 , heights 132-138 mm,
Shark Bay, WA, Poole Bros, Jul 1963. WAM P6067, 8, height
149 mm, Shark Bay, WA, Poole Bros, 5 Jun 1961. WAM P8174,
9, height 106 mm, Shark Bay, WA, Poole Bros, Bluefin, May
1964. WAM P14580, 9, height 96 mm, Shark Bay, WA, trawl, R.
McKay, 3 Mar 1962. WAM P14590, 8, height 76 mm, Shark
Bay, WA, trawl, B.R. Wilson, 5 Jun 1980. WAM P14738, 8 , height
135 mm, east of Delambre I., Dampier Archipelago, WA, Poole
Bros, September 1963. WAM P26053-001, height 140 mm, about
15 mi WSW Carnarvon, WA, c. 14 fm, L. Marsh & M. Sinclair,
2-3 Jul 1975. WAM P27351-001, 8, height 134 mm, N Monte
Bello I., WA, 20°05'S 115°45'E, Taiwanese Pair Trawl, J. Marek,
8 Nov 1981. WAM P27677-001, 9, height 96 mm, about 15 mi
WSW Carnarvon, WA, depth 24-31 m, L. Marsh & M. Sinclair,
2-3 Jul 1975. WAM P29077-001, juvenile, height 36 mm, Denham,
WA, 25°32'S 113°32'E, depth 1-5 m, L.R. Lenanton, 31 Jul 1965.
Hippocampus grandiceps n.sp.
Big-head Seahorse
Fig. 52
Type material. Holotype: NTM S13273-019, 8 , height
76 mm, W Booby Island, Gulf of Carpentaria, Queensland,
10°44'S 141°53'E, depth 10 m, R. Williams, 29 Nov 1991.
Paratypes: NTM SI3274-007 (3), 8 ,height75 mm; 2 9 9 ,
heights 62 & 69 mm; both N Booby I., Gulf of Carpentaria,
Qld, 10°26'S 141°45'E, depth 10.4 m, R. Williams, 29 Nov
1991. QM 130583, 8 , height 99 mm, Gulf of Carpentaria,
Qld, 11°29'S 141°38'E, dredge, depth 18 m, J. Johnson &
S. Cook, 3 Dec 1990. QM 112287, 9 , height 96 mm, Gulf
of Carpentaria, Qld, 16°40'30"S 140°58'30"E, trawl, depth
6 fm, CSIRO Rama , 6 Oct 1963. QM 126319, 9 , height
105 mm, Gulf of Carpentaria, Qld Fisheries, Jul 1983. AMS
115557-075 (3), 9 9 , heights 95-105 mm, Gulf of
Carpentaria, Qld, 17°24'S 140°09'E, prawn trawl, depth 10
m, CSIRO, 27 Nov 1963.
Diagnosis. Dorsal-fin rays 18; pectoral-fin rays 17-18
(usually 18); trunk rings 11; tail rings 32-33; head long,
94-103% of trunk length; head angled down to near trunk.
Description. Head long, 94% of trunk length (over 100%
in paratypes); snout long, just under half head length (42-
48%); dorsal fin with 18 rays (17-18), base over 2 trunk
and 1 tail ring; pectoral-fin rays 18; trunk rings 11; tail rings
33 (32-33); subdorsal spines 3/0,1,0; spine above eye as
long as pupil diameter, perpendicular to snout; nose spine
angular, directed forward 45° to snout; length of nape spine
equal to pupil diameter, directed somewhat forward; spine
Kuiter: Australian Hippocampus revision 337
Figure 52. Hippocampus grandiceps. A, 6, holotype, NTM SI 3273-019, Gulf of Carpentaria, Qld. B, 6 paratype,
NTM SI 3274-007, Gulf of Carpentaria, Qld. C & D, 9 $, paratypes, NTM SI 3274-007, Gulf of Carpentaria, Qld.
of moderate length behind eye; double spines below eye;
shoulder-ring spines long, uppermost and central spines at
levels with either ends of pectoral-fin base, lowermost
double, anterior barb smaller and directed forward; lateral
head spine moderately long; coronet of moderate height
with 5 short and sharp diverging spines at apex; neck ridge
with 2 short spines, one centrally and one at posterior end;
superior trunk ridge with short spines, slightly enlarged at
regular intervals from 1st ring to below dorsal fin base;
lateral ridge with spines on 2nd to 10th ring, those on rings
2, 4, and 6-10 enlarged; inferior trunk ridge with series of
spines from 4th to 11th ring, progressively from short to
long; ventral trunk ridge with downward angled spines in
females and scalloped edge in males; superior tail ring spines
moderately long from 2nd to 14th ring, becoming
progressively smaller posteriorly. Lateral line with small
indistinct pores, increasingly becoming more difficult to
detect posteriorly, reaching 21st tail ring. Height of largest
specimen examined (female) 105 mm. Colour in life :
unknown. Colour in alcohol : pale brown-grey with pale
saddle-like areas on trunk and tail. Snout with distinct dusky
barring along entire length.
Distribution (Fig. 53). Appears to be restricted to the
Queensland side of the Gulf of Carpentaria. All specimens
trawled or dredged in relatively shallow depths, usually
during prawn surveys.
Remarks. This species is named grandiceps from the Latin
words for large and head in reference to the head in this
species that is proportionally larger than most others in the
genus. Hippocampus grandiceps is very similar to H.
multispinus but has shorter spines and its males lack long
spines over the superior trunk ridge anterior to the dorsal
fin, while males of H. multispinus have long spines in this
position. Where H. grandiceps has spines on the 3rd and
5th rings at the lateral-ridge, they are absent in H.
multispinus. While the head in specimens of H. grandiceps
is held close to the trunk, it is perpendicular to the axis of
Figure 53. Hippocampus grandiceps. Collection sites of specimens
examined.
the body in specimens of H. multispinus. The maximum
size of H. grandiceps is considerably less than that of H.
multispinus and other similar species. Although, H.
spinosissimus also has prominent spines and attains a smaller
maximum size, it has fewer rays in both the dorsal and
pectoral fins. Hippocampus hendriki is also similar to H.
grandiceps , but is closer to H. multispinus , sharing with it
the absence of spines on the lateral-ridge of the 3rd and 5th
trunk ring, and a head which is perpendicular to the body.
The limited geographical range of H. grandiceps may reflect
its preference for a unique habitat, the head-angle, body
patterns, and shallower depth range suggest that it may
inhabit more weedy areas than those populated by most
other species with prominent spines (see remarks, H.
hendriki).
338 Records of the Australian Museum (2001) Vol. 53
Table 1. Selected meristic character set of Australian species of Hippocampus. O indicates value found in material examined,
• indicates usual count.
rings
trunk tail
8 9 10 11 12 13 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48
abdominalis
alatus n.sp.
angustus
bargibanti
biocellatus n.sp.
bleekeri
breviceps
dahli
elongatus
grandiceps n.sp.
hendriki n.sp.
histrix
jugumus n.sp.
kampylotrachelos
minotaur
montebelloensis n.sp.
multispinus n.sp.
planifrons
procerus n.sp.
queenslandicus
semispinosus n.sp.
spinosissimus
taeniopterus
tristis
tuberculatus
whitei
zebra
•a
—
o
o
— —
—
—
_ _ _
—
—
_ _ _ _
■D
-
-
- -
—
—
fS
-
-
_ _ _ _
m
—
—
- o
o
Eg
o
-
- o
—
—
O - -
-
-
_ _ _ _
o
-
-
- -
—
—
irt
-
o
_ _ _ _
_
o
•
o
—
—
— —
—
—
_ _ _
—
o
- - - o
#
-
-
- -
-
-
_ _ _
o
—
• O - -
it
—
—
— —
—
o
O - -
—
—
_ _ _ _
o
-
-
- -
o
o
_ _ _
—
—
_ _ _ _
o
o
o
o
-
- -
-
o
O - -
o
-
- - - -
o
o
- - - - - -------O-----
^ - ____o#-------
v - - - - - % ______
- - - o - - - - - - - o o A ^ « «- —
- - - O - - -------O-----
mm - -
o
fin rays
dorsal pectoral
7... 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 10 11...13 14 15 16 17 18 19 20
abdominalis
alatus n.sp.
angustus
bargibanti
biocellatus n.sp.
bleekeri
breviceps
dahli
elongatus
grandiceps n.sp.
hendriki n.sp.
histrix
jugumus n.sp.
kampylotrachelos
minotaur
montebelloensis n.sp.
multispinus n.sp.
planifrons
procerus n.sp.
queenslandicus
semispinosus n.sp.
spinosissimus
taeniopterus
tristis
tuberculatus
whitei
zebra
m
- - o -
-oo
O - O - - - -
- - - o
O O - -
^ _ _ ._ _ _ _ _ _ _
---------O---
-----O-------
-- ------
____ 0 #-------
-------O-----
— _ - — - --
- - o
Mp 4 - -- -- -- -- --
__________#o------
- - - -«$&'•
- - - - •mf ---------- -
- - - Mb t - -- -- -- -- --
_ _ _ _
- - -
_ _ _ o-m -------------
- - - - mm a* - - -
- - - - ■«■%.- - -
_ _ _ _ • - - O
_ -■ - O - -
__ _*#____
- - oo------
__ ----•O--
_ _ _ - ^ ^ o - -
- - — -
--*4$ tie-
- - - - -^ 4* o - -
_ _ _ _ ^ p-A, _ _ _
- - - - sJp-sJi - - -
- O - -- -- -- -
_ _ _ _ flfp - - - -
- - - - O - -
__ - -- -- OO-
_ _ _ _ - -
- - - - - m • • -
- - - — if-
_ _ _ _ ftp - - -
__
- - - -- -- OO-
- - - o dp**.** - - -
__ _ _ y&m o - - -
- - - -OO----
Kuiter: Australian Hippocampus revision 339
Acknowledgments. M. McGrouther and J. Pogonoski, AMS,
J. Johnson, QM, B. Hutchins, WAM, T. Simms, SAMA, T.
Bardsley, NMV, and H. Larson, NTM, and A. Graham, CSIRO,
facilitated the loan of specimens. M. Horne, James Cook
University, provided specimens of H. queenslandicus and H. dahli.
K. Graham, NSW Fisheries, provided information and photographs
on NSW specimens of H. tristis. S. Lourie, Project Seahorse,
provided photographs of the types of H. tristis and H. planifrons.
R. Steene and T. Wu provided photographs of H. bargibanti. A.
Ogawa provided the photograph of H. kampylotrachelos. H.
Uchiyama provided the photograph of H. histrix. Maurice Pruvost,
MNHN, provided photographs of the types of H. elongatus and
H. subelongatus. Martin Gomon, Museum Victoria, advised on
taxonomy and various aspects of this work. I appreciate the time
taken by Helen Larson, Martin Gomon and John Paxton to read
and comment on the manuscript, improving it in many ways.
References
Allen, G.R., 1998. A new species of cardinalfish (Apogonidae)
from the Komodo Islands, Indonesia. Revue frangaise
Aquariologie 25: 1-2.
Allen, G.R., D.F. Hoese, J.R. Paxton, J.E. Randall, B.C. Russell,
W.A. Starck II, F.H. Talbot & G.P Whitley, 1976. Annotated
checklist of the fishes of Lord Howe Island. Records of the
Australian Museum 30: 365-454.
Allen, G.R., & R.C. Steene, 1987. Reef Fishes of the Indian Ocean.
Plate 18-6. Neptune City, New Jersey: T.F.H. Publication.
Bertin, L., & R. Esteve, 1950. Catalogue des types de poissons du
Museum national d’Histoire naturelle. 6e partie. Imp. Nationale,
Cat. Fish Types, Paris 6e partie: 1-85.
Bleeker, P, 1852a. Bijdrage tot de kennis der ichthyologische
fauna van Singapore ( Memoire sur lafaune ichthyologique
de Singapore ). Natuurkundig Tijdschrift voor Nederlandsch
Indie 3: 82-83.
Bleeker, P, 1852b. Bijdrage tot de kennis der ichthyologische fauna
van de Moluksche eilanden Visschen van Amboina en Ceram.
Memoire sur lafaune ichthyologique des Moluques Poissons
d’Amboine et de Ceram 3: 305-307.
Bleeker, P, 1854a. Vijfde bijdrage tot de kennis der ichthyologische
fauna van Amboina. Cinquieme memoire sur la faune
ichthyologique d’Amboine 3: 505-506.
Bleeker, P, 1854b. Overzigt der ichthyologische fauna van
Sumatra, met beschrijving van eenige nieuwe soorten. Aregu
de lafaune ichthyologique de Sumatra 7: 107-108.
Bleeker, P, 1855. Over eenige visschen van Van Diemensland.
Verhandelingen van het Koninklijke Akademie van Wetenschappen
te Amsterdam 2: 17, 28-31.
Bleeker, P, 1983. Atlas Ichthyologique des Indes Orientales
Neerlandaises. Plates for volumes XI-XIV, pi. 449.
Washington: Smithsonian Institution.
Castelnau, F.L. de, 1872. Contribution to the ichthyology of
Australia. The Melbourne fish market. Proceedings of the
Zoological Acclimatisation Society of Victoria 1: 196-197.
Castelnau, F.L. de, 1873. Contribution to the ichthyology of
Australia. 8. Fishes of Western Australia. Proceedings of the
Zoological Acclimatisation Society of Victoria 2: 144-145.
Castelnau, F.L. de, 1875. Researches on the Fishes of Australia.
Intercolonial Exhibition Essays, Commissioners Melbourne
International Exhibition, Melbourne 2: 3-52.
Dawson, C.E., 1985. Indo-Pacific Pipefishes (Red Sea to
Americas). Ocean Springs (Mississippi): The Gulf Coast
Research Laboratory.
Eschmeyer, W.N., 1998. Catalog of Fishes. Volumes 1-3. San
Francisco: California Academy of Sciences.
Fowler, H.W., 1907. A collection of fishes from Victoria,
Australia. Proceedings of the Academy of Natural Sciences of
Philadelphia 59: 419-444.
Ginsburg, I., 1937. Review of the seahorse ( Hippocampus ) found on
the coasts of the American continents and of Europe. Proceedings
of the United States National Museum 83: 497-594.
Gloerfelt-Tarp, T., & PJ. Kailola, 1984. Trawled Fishes of Southern
Indonesia and Northwestern Australia. Australian Development
Assistance Bureau; Directorate General of Fisheries, Indonesia;
German Agency for Technical Cooperation.
Gomon, M.F., 1997. A remarkable new pygmy seahorse
(Syngnathidae: Hippocampus ) from south-eastern Australia,
with redescription of H. bargibanti Whitley from New
Caledonia. Memoirs of the Museum of Victoria 56(1): 245-
253.
Grant, E.M., 1987. Fishes of Australia. Brisbane: E.M. Grant Pty.
Ltd.
Gunther, A., 1870. Catalogue of the Fishes in the British Museum.
Vol. 8, pp. 198-206. London: British Museum.
Horne, M.L., 2001. A new seahorse species (Syngnathidae:
Hippocampus ) from the Great Barrier Reef. Records of the
Australian Museum 53(2): 243-246.
Johnson, J.W., 1999. Annotated checklist of the fishes of Moreton
Bay, Queensland, Australia. Memoirs of the Queensland
Museum 43(2): 709-762.
Kaup, J.J., 1856. Catalogue of the Lophobranchiate Fish in the
Collection of the British Museum. Pp. 76. London: British Museum.
Kawanabe, H., & N. Mizuno, 1989. Freshwater Fishes of Japan.
Tokyo: Yama-Kei Publishers.
Kuiter, R.H., 2000. Seahorses, Pipefishes & Their Relatives.
England: TMC publications.
Lesson, R.P, 1827. Espece nouvelle d’Hippocampe. Bulletin
Sciences National (Ferussac) II: 127-128.
Lourie, S.A., A.C.J. Vincent & H.J. Hall, 1999. Seahorses. An
Identification Guide to the World’s Species and Their
Conservation. London: Project Seahorse.
McCulloch, A.R., 1911. Report on fishes obtained by the F.I.S.
Endeavour on the coasts of New South Wales, Victoria, South
Australia and Tasmania. Part 1. Zool. Res. Endeavour 1(1): 1-78.
Nakabo, T., 1993. Fishes of Japan with Pictorial Keys to the
Species. Tokyo: Tokai University Press.
Ogilby, J.D., 1889. Notes on some fishes new to the Australian fauna.
Proceedings Zoological Society London 1889 (pt. 2): 151-158.
Ogilby, J.D., 1908. New or little known fishes in the Queensland
Museum. Annuals of the Queensland Museum 9 : 3-41.
Paxton J.R., D.F. Hoese, G.R. Allen & J.E. Hanley, 1989.
Zoological Catalogue of Australia 7 Pisces Petromyzontidae
to Carangidae, pp. 420-422. Canberra: Australian Government
Publishing Service.
Peters, W.C., 1869. Uber neue oder weniger bekannte Fische des
Berliner Zoologischen Museums. Monatsberichte der
Koniglichen Preussischen Akademie Wissenschaften zu Berlin
1869:703-711.
Peters, W.C., 1877. Reise S.S.S. “Gazelle”: uberstanded Fische.
Monatsberichte der Koniglichen Preussischen Akademie
Wissenschaften zu Berlin 1876: 831-854.
Rafinesque, 1810. Carratteri di alcuni nuovi generi e nuove specie
di animali e piante della Sicilia, 18.
Randall, J.E., G.R. Allen & R.C. Steene, 1990, 1997. Fishes of
the Great Barrier Reef and Coral Sea, 1st and 2nd editions.
Crawford House Press.
Steindachner, F., 1866. Zur Fischfauna von Port Jackson of Australien.
Sitzungberichte Akademie Wissenschaften, Wien 53: 424—480.
Weber, M., 1913. Die Fische der Siboga-Expedition. Siboga
Expeditie Reportage 57. Leiden: E.J. Brill.
Weber, M., & L.F. de Beaufort, 1922. The Fishes of the Indo-
Australian Archipelago. Volume 4. Leiden: E.J. Brill.
White, J., (1790). Journal of a voyage to New South Wales with
sixty-five plates of non-descript animals, birds, lizards,
serpents, curious cones of trees and other natural productions.
London: Debrett.
Whitley, G.P, 1931. New names for Australian fishes. Australian
Zoologist 6: 310-334.
340 Records of the Australian Museum (2001) Vol. 53
Whitley, G.R, 1940. Illustrations of some Australian fishes.
Australian Zoologist 9: 397-428.
Whitley, G.R, 1952. Some noteworthy fishes from Eastern
Australia. Proceedings of the Royal Society of New South Wales
1950-1951:27-32.
Whitley, G.R, 1964. Fishes from the Coral Sea and the Swain
Reefs. Records of the Australian Museum 26(5): 145-195.
Whitley, G.P., 1970. Abstract of proceedings. Ordinary general
meeting. 26th November 1969. Proceedings of the Linnaean
Society of New South Wales 94(3): 292-295.
Whitley, G.P., & J. Allan, 1958. The Sea-Horse and its Relatives .
Melbourne: Georgian House.
Manuscript received: 20 July 2000, revised 21 February 2001 and accepted
27 February 2001.
Associate Editor: J.M. Leis.
Index to species
abdominalis, Hippocampus .322
alatus n.sp., Hippocampus .317
angustus, Hippocampus .335
bargibanti, Hippocampus .305
biocellatus n.sp., Hippocampus .311
bleekeri, Hippocampus .323
breviceps, Hippocampus .312
dahli, Hippocampus .308
elongatus, Hippocampus .329
grandiceps n.sp., Hippocampus .336
hendriki n.sp., Hippocampus .333
histrix, Hippocampus .333
jugumus n.sp., Hippocampus .306
kampylotrachelos, Hippocampus .307
minotaur, Hippocampus .304
montebelloensis n.sp., Hippocampus .326
multispinus n.sp., Hippocampus .331
planifrons, Hippocampus .310
procerus n.sp., Hippocampus .328
queenslandicus, Hippocampus .319
semispinosus n.sp., Hippocampus .320
spinosissimus, Hippocampus .330
taeniopterus, Hippocampus .314
tristis, Hippocampus .316
tuberculatus, Hippocampus .313
whitei, Hippocampus .327
zebra, Hippocampus .325
© Copyright Australian Museum, 2001
Records of the Australian Museum (2001) Vol. 53: 341-406. ISSN 0067-1975
Snailfishes (Pisces: Liparidae) of Australia,
Including Descriptions of Thirty New Species
David L. Stein, 1 Natalia V. Chernova 2 and Anatoly P. Andriashev 2
1 NO A A/OAR R/OSS3, 1315 East West Highway Room 11805,
Silver Spring, MD 20910, United States of America
david. stein @ noaa. go v
2 Zoological Institute of St. Petersburg, Russian Academy of Sciences,
St. Petersburg, 199034, Russia
antarct@zin.ru
Abstract. Examination of all available liparid specimens collected in Australian waters revealed more
than 30 new species in three genera: Careproctus Kr0yer, Psednos Barnard, and Paraliparis Collett.
This is the first published description of liparids from Australia. The new species include Careproctus
paxtoni n.sp., Psednos balushkini n.sp., Ps. nataliae n.sp., Ps. whitleyi n.sp., Paraliparis anthracinus
n.sp., P. ater n.sp., P. atrolabiatus n.sp., P. auriculatus n.sp., P. australiensis n.sp., P. avellaneus n.sp., P.
badius n.sp., P. brunneocaudatus n.sp., P. brunneus n.sp., P. coracinus n.sp., P. costatus n.sp., P. csiroi
n.sp., P. delphis n.sp., P. dewitti n.sp., P. eastmani n.sp., P gomoni n.sp., P. hobarti n.sp., P impariporus
n.sp., P. infelicitern.sp., P. labiatus n.sp., P. lasti n.sp., P. obtusirostris n.sp., P. piceus n.sp., P. plagiostomus
n.sp., P. retrodorsalis n.sp., P. tasmaniensis n.sp., and four more unnamed taxa. All are endemic to
Australia. We discuss and describe new characters, describe the new species and provide keys for their
identification. The described Paraliparis species are morphologically very similar to one another, and
form a group that is clearly distinct from the Antarctic species of the genus, suggesting that the two
groups are not closely related and that neither is derived from the other. Two of the unnamed taxa
constitute the second record of liparids from the Indian Ocean.
Stein, David L., Natalia V. Chernova & Anatoly R Andriashev, 2001. Snailfishes (Pisces: Liparidae) of
Australia, including descriptions of thirty new species. Records of the Australian Museum 53(3): 341-406.
The family Liparidae was long considered to be almost
completely confined to the Northern Hemisphere (Burke,
1930), with only a few poorly known species described from
the South Atlantic, the southern tip of South America, and
the Antarctic. In fact, liparids seem to occur worldwide in
marine environments where temperatures are low enough.
In equatorial areas, they exhibit tropical submergence and
occur in cold waters at great depths, but (with one exception,
Liparis fishelsoni Smith, 1967) not in shallower, warmer
waters. Within the last ten years, it has become apparent
that the Southern Hemisphere has an extensive liparid fauna
composed entirely of endemic species. The Southern Ocean
is particularly rich, having well over 100 species
representing seven genera (Andriashev, 1986; Andriashev,
1993; Stein & Andriashev, 1990; Andriashev & Stein, 1998;
Duhamel, 1992; Stein & Tompkins, 1989). There is also a
Chilean group of liparids, including a morphologically
notable endemic genus, Eknomoliparis (Stein etal., 1991).
342 Records of the Australian Museum (2001) Vol. 53
Given this “explosion” of new taxa from a poorly known
region, it is not surprising that other temperate southern
waters also include diverse liparid species.
Recent interest in Australian deep-water fisheries has led
to collections in previously unsampled areas and depths
(Koslow et al., 1994; May & Blaber, 1989). Although
liparids have been reported from Australian waters (Koslow
etal ., 1994; Williams, Last, Gomon, & Paxton, 1996) they
were tentatively identified only to family or genus, and no
species had been described from Australian waters.
Examination of all available material revealed more than
30 new species in three genera of which we describe 30:
Careproctus Krpyer, 1862 (1 new species); Psednos
Barnard, 1927 (3 new species); and Paraliparis Collett,
1878 (26 new species) (Table 1). In this paper we describe
all extant material, name these species where possible, and
provide keys for their identification. In addition, we
describe, but owing to their poor condition, do not name,
three additional species of Paraliparis and one of Psednos.
Because of the diversity and distribution of the liparids
described here, we provide a short description of the south¬
eastern Australian continental slope and its environment.
With two exceptions (the specimens collected off northwest
Australia), all our specimens were collected from about
130°E in the Great Australian Bight, western and
southwestern Tasmania, the eastern end of Bass Strait, and
off Victoria and New South Wales as far north as about
33°S (Fig. 1). Although the surface currents and near shore
(shelf) benthic environment of the region are fairly well
known, until recently those of greater depths were not (Bunt,
1987). To the west, the gently sloping (1:600) heavily
sedimented Cedura Plateau extends between about 130-
134°. Between it and Tasmania (from 134-141°E) the slope
is very steep, cut by many deep canyons with vertical relief
up to 2000 m (Williams & Corliss, 1982). Further east and
south, the continental shelf on both sides of Tasmania is
very narrow and steep so that deep water occurs relatively
close to shore (Exon et al, 1995). Some very large canyons
occur on the eastern side of Bass Strait and on the southern
Victoria coast (Conolly, 1968). Sediments inshore are
primarily CaC0 3 /Si0 2 ; farther offshore on the upper slope
and below, they are CaC0 3 (Williams & Corliss, 1982).
Oceanographically, the southern coast of Australia as far
east as 147°E (the southern tip of Tasmania) is considered
to be part of the Indian Ocean (Rao & Griffiths, 1998);
waters further to the east are part of the western Pacific.
Table 1 . List of new species collected, registration (including type status: H—holotype, P—paratype, sex, museum,
number); standard length (SL, mm); collection location; depth (m); and date of capture.
and specimen
new species collected
registration
SL
collection location
depth
capture date
Careproctuspaxtoni n.sp.
H
$
AMS 129737-003
134
35°29'S 150°55'E
1116-1134
31 Aug 1989
Careproctus paxtoni n.sp.
P
$
AMS 129802-001
144
35°28.5'S 150°53.5'E
1061-1088
30 May 1989
Psednos balushkini n.sp.
H
c?
AMS 124860-002
84
34°48.5'S 151°15.5'E
914-960
4 Oct 1984
Psednos balushkini n.sp.
P
3
AMS 124059-018
82
33°32'S 152°09'E
942-978
23 Aug 1983
Psednos nataliae n.sp.
H
3
CSIRO H2636-04
98.5
42°16'S 144°39'E
1100-1120
18 Mar 1989
Psednos whitleyi n.sp.
H
3
CSIRO H1335-02
91.2
41°18.4'S 144°05'E
900-920
14 May 1986
Paraliparis anthracinus n.sp.
H
$
CSIRO HI576-02
133
41°46.4'S 144°24.4'E
1024-1080
14 May 1986
Paraliparis ater n.sp.
H
$
CSIRO H749-04
124
41°45.8'S 144°24.8'E
1000-992
16 May 1986
Paraliparis atrolabiatus n.sp.
H
3
CSIRO H550-11
114
42°20.45'S 144°40.4'E
1120-1220
17 May 1986
Paraliparis auriculatus n.sp.
H
2
CSIRO H749-06
131
41°45.8'S 144°24.8'E
1000-992
16 May 1986
Paraliparis australiensis n.sp.
H
$
NMVA21497
164
37°01.09'S 137°25.44'E
1090-1160
24 Jan 1988
Paraliparis avellaneus n.sp.
H
3
NMVA5873
132
37°01.09'S 137°25.44'E
1090-1160
24 Jan 1988
Paraliparis badius n.sp.
H
a
CSIRO T1981-01
82
off Tasmania
unknown
20 Oct 1984
Paraliparis brunneocaudatus n.sp.
H
3
CSIRO T1980-01
127
W coast of Tasmania
unknown
Apr 1984
Paraliparis brunneus n.sp.
H
$
CSIRO H749-05
151
41°45.8'S 144°24.8'E
1000-992
16 May 1986
Paraliparis coracinus n.sp.
H
$
CSIRO H1935-02
70+
37°34.53'S 138°57.00'E
1205-1175
1 Feb 1989
Paraliparis costatus n.sp.
H
3
CSIRO H561-02
204
41°51.4'S 144°23.8'E
1366-1370
16 May 1986
Paraliparis costatus n.sp.
P
2
CSIRO H1378-01
235
42°12'S 144°38'E
1042-1080
21 Apr 1988
Paraliparis csiroi n.sp.
H
2
NMVA5874
163
38°37.58'S 141°01.12'E
1080-1110
8 Feb 1988
Paraliparis delphis n.sp.
H
3
CSIRO H749-03
127
41°45.8'S 144°24.8'E
1000-992
16 May 1986
Paraliparis dewitti n.sp.
H
2
CSIRO T889-02
192
34°26.5'S 132°04'E
1175-1118
14 Nov 1984
Paraliparis eastmani n.sp.
H
3
AMS 128900-003
183
33°29.5'S 152°12.5'E
1035-1070
1 Sep 1988
Paraliparis gomoni n.sp.
H
3
NMVA7124
110
39°00.92'S 148°43.71'E
1140-1160
13 May 1988
Paraliparis hobarti n.sp.
H
2
CSIRO H3170-01
124
continental slope of Tasmania 13
Paraliparis impariporus n.sp.
H
2
CSIRO H3168-01
162
41°51.97'S 144°27.16'E
1040-1050
14 Mar 1989
Paraliparis infeliciter n.sp.
H
2
CSIRO H3169-01
153
37°37.9'S 139°00.60'E
1070-1090
30 Jan 1988
Paraliparis labiatus n.sp.
H
2
CSIRO H749-02
150
41°45.8'S 144°24.8'E
1000-992
16 May 1986
Paraliparis lasti n.sp.
H
3
CSIRO T982-02
185
33°45.5'S 129°37.5'E
1152-1000
5 Jun 1983
Paraliparis obtusirostris n.sp.
H
2
NMVA7123
137
38°57.09'S 148°41.95'E
1270-1290
13 May 1988
Paraliparispiceus n.sp.
H
3
CSIRO H805-04
149
41°51.25'S 144°23.1'E
1384-1416
18 May 1986
Paraliparisplagiostomus n.sp.
H
3
CSIRO T488-02
137
42°19.5'S 144°42'E
993-987
9 Jul 1983
Paraliparis retrodorsalis n.sp.
H
2
CSIRO H1935-01
145
37°34.53'S 138°57.00'E
1205-1175
1 Feb 1989
Paraliparis tasmaniensis n.sp.
H
2
CSIRO H2679-02
201
40°26.64'S 143°18.36'E
1000-1100
6 Mar 1989
Paraliparis sp. 1
2
CSIRO H549-05
134+
41°50.4'S 144°23.45'E
1328-1288
25 May 1986
Paraliparis sp. 2
3
CSIRO H555-04
131
42°20.6'S 144°37.25'E
1376-1404
17 May 1986
Paraliparis sp. (cf. copei group)
3
AMS 122809-036
c.178
18°40'S 116°42'E
584-592
4 Apr 1982
Paraliparis sp. (cf. copei group)
3
AMS 122813-018
154+
18°32'S 116°50'E
658-660
6 Apr 1986
juvenile, sex unknown b no other data
130°
135°
140 °
Stein et al.: Australian liparids 343
jW5° 150° 155°
30 °
35°
40°
45°
50 °
, Paraliparis lasti n.sp.
Psednos balushkini n. sp^
Paraliparis australiensis n.sp^
Paraliparis avellaneus n.sp.
pdralifwis obtusirostris n.sp.
Paraliparis coracinus n.sp.
Paraliparis retrodorsalis n.sp.
Paraliparis anthracinus sp.n.
Paraliparis ater n.sp.
Paraliparis auriculatus n.sp,,
Paraliparis costatus n.sp.
Paraliparis delphis n.sp.
Paraliparis impariporus n.sp.
Paraliparis piceus n.sp.
Paraliparis sp.1
araliparis gomoni n.sp.
Psednos nataliae n. sp.
Paraliparis atrolabiatus n.sp.
Paraliparis plagiostomus n.sp.
Paraliparis sp.2
Paraliparis brunneus n.sp.
Paraliparis labiatus n.sp.
30°
35°
40°
45°
50°
130 0 135° 140° 145° 150°
Figure 1 . Chart of southeastern Australia, showing collection locations of the new species where known.
155°
Throughout both areas, slope waters are generally derived
or at least strongly influenced by the cold Antarctic
Intermediate Water (Tchernia, 1980). Temperatures on the
southeastern coast of Victoria range from 7.0° at 720 m to
3.6° at almost 1300 m depth (Huyer et al., 1988). A little
further to the south, during the summer there is an upper
oxygen minimum at 300-500 m, a deep oxygen minimum
at about 1500 m, and a salinity min imum from the Antarctic
Intermediate Water at 700-1100 m (Harris et al., 1987).
The flow in the Bight is from west to east along the slope;
on the western side of Tasmania, surface temperatures are
lower than on the eastern side owing to the influence of the
colder subantarctic waters (Rochford, 1975); on the eastern
side, currents are more complex, becoming northward at
depth but influenced by the warmer, shallower, East
Australian Current and (at least in winter) by deeper water
flowing out of the Bass Strait northwards (Godfrey et al.,
1980; Hamilton, 1990) with the result that “The waters
around Tasmania are a mixture of both subtropical and
subantarctic origin” (Harris et al., 1987).
Materials and methods
We follow Andriashev & Stein (1998) for abbreviations and
methods of counts and measurements with some important
additions, described below. All specimens examined were
preserved in ethanol, and little information was available
regarding colour in life. Collection location coordinates are
averages of the start and finish coordinates of the trawl hauls.
Institutional abbreviations are listed in Leviton et al. (1985).
Abbreviations for counts
A anal-fin rays
C caudal-fin rays
D dorsal-fin rays
f fenestra in pectoral girdle; f 1 is dorsalmost
gr number of gill rakers on the outside of the first
arch
P pectoral-fin rays
r rudimentary pectoral-fin ray
R radial of pectoral girdle; R1 is dorsalmost
pc pyloric caeca
Vert, vertebrae
Abbreviations for pores
Sensory pores of the cephalic series for Careproctus and
Paraliparis are given in the order nasal (n), maxillary (m),
preoperculo-mandibular (pm), and suprabranchial (s)
(Andriashev etal., 1977; Burke, 1930). The maxillary series
includes infraorbital (i.o.^) and first temporal (=
postorbital) (tj) pore. The suprabranchial pores (tsb^) are
the last in the temporal series. For Psednos, the pores are
described in greater detail in the generic and individual
descriptions.
344 Records of the Australian Museum (2001) Vol. 53
Table 2. Ranges and limits of variation of selected proportional measurements, specifying species with morphometry near the extremes,
and providing guidelines for use of relative terminology in Australian Paraliparis only. Minimum or lower (min) values of a range on
left side, maximum or higher (max) values on right side; shading distinguishes each measurement.
measurement and measurement and
min term selected species term max min term selected species term max
17.7
head (HL) as % SL: 17.7-21.4
short.sp. 2
dewitti .
.... long
21.4
head width as % HL: 49-67
49
compressed
. eastmani
dewitti, atrolabiatus .
.... wide
67
head depth as % HL 69-103
69
shallow.
. avellaneus
72
. infeliciter, sp. 2
piceus .
93
impariporus .
.... deep
103
body depth (bd) as % HL 69-132
69
shallow.
. badius
85
.sp. 2
australiensis, csiroi, tasmaniensis .
125
coracinus, impariporus .
.... deep
132
body depth at anal-fin origin (bdA)
as % HL 66-113
66
shallow.
. badius
77-82 .
sp. 2, avellaneus, gomoni
piceus .
108
impariporus .
111
tasmaniensis .
.... deep
113
predorsal length (preD) as % SL 16.1-
28.3
16.1
short.
. eastmani
18.5
.sp. 2
lasti .
26.0
impariporus .
26.8
retrodorsalis .
.... long
28.3
preanal fin length (preA) as % SL 29.0-40.3
29.0
short.
. auriculatus
30.1
. plagiostomus
impariporus .
40.0
costatus .
.... long
40.3
mandible to anus (ma) as % SL 10-15
10.5
short.
. infeliciter
10.9
. plagiostomus
11.4
. lasti
ater, hobarti, impariporus .
. 14.5-
-14.8
anthracinus, brunneocaudatus ....
.... long
15.0
anus to anal fin (aAf) as % SL 15.9-30.0
15.9
short.
. badius
16.7
. auriculatus
retrodorsalis .
25.9
australiensis .
.... long
30.0
upper pectoral-fin lobe length (UPL)
as % HL 60.5-83
60.5
short.
. avellaneus
62
. infeliciter
63
. auriculatus, australiensis, dewitti,
plagiostomus.
brunneocaudatus, lasti .
75
gomoni .
76
piceus .
78
hobarti .
.... long
83
lower pectoral-fin lobe length (LPL)
as % SL 7.2-13.5
7.2
short.
. atrolabiatus
8.5
. infeliciter
8.8
. plagiostomus
gomoni .
12.8
brunneocaudatus .
13.4
lasti .
.... long
13.5
lower pectoral-fin lobe length (LPL)
as % HL 37-85
37 short. atrolabiatus
42 . retrodorsalis
46.5 .sp. 1
csiroi . long 85
61
69
18.8
21.0
LPL as % UPL 61-95
short. atrolabiatus
. impariporus
avellaneus . 93
brunneocaudatus . 94
badius . long 95
eye (E) as % HL 18.8-28.9
small. tasmaniensis
. labiatus.
auriculatus, eastmani, hobarti,
impariporus, obtusirostris . 25.4-26.0
coracinus . 27.0
badius .large 28.9
postorbital length as % HL 45.2-53.5
45.0-45.2 short. hobarti, infeliciter.
coracinus, impariporus, plagiostomus,
tasmaniensis .long 53.0-53.5
snout as % HL 27.8-37.5
. anthracinus
. hobarti
costatus, dewitti . 35.0-36.6
australiensis . long 37.5
27.8 short
30.8 .
30.1
31.3
34.3
interorbital width as % HL 30.1-48.3
narrow. plagiostomus
. csiroi
. avellaneus
labiatus . 44.1
ater . 47.0
auriculatus . broad 48.3
upper jaw as % HL 41.4-54.7
41.4
short
. anthracinus
42.5-42.6 ..
.sp. 1, tasmaniensis
eastmani .
51.3
lasti .
52.0
plagiostomus .
.. long
54.7
lower jaw as % HL 38.3-48.4
38.3
short
. brunneus
39.5
.sp. 1.
atrolabiatus, costatus, plagiostomus .
... 46.0-48.0
eastmani .
.. long
48.4
pyloric caeca as % SL 3.0-9.8
3.0
short
. dewitti
australiensis .
.. long
9.8
gill opening as % HL 15.1-23.6
15.1
short
. badius
15.9
. atrolabiatus
16.4
. auriculatus
obtusirostris .
22.7
brunneocaudatus .
22.8
impariporus .
.. long
25.6
Stein et al .: Australian liparids 345
Abbreviations for measurements
aAf distance from centre of anus to anal-fin origin
bd maximum body depth
bdA body depth at anal-fin origin
disk longitudinal diameter of disk
E eye diameter horizontally
gs length of gill opening
HL head length
io interorbital width
lj lower j aw length
LPL greatest length of lower lobe of pectoral fin
ma length from mandibular symphysis to centre of
anus
md length from mandibular symphysis to anterior
edge of disk
NL length of shortest notch ray
po postorbital head length
preA preanal-fin length
preD predorsal-fin length
sn snout length
so suborbital distance, shortest distance between
margin of eye and horizontal level of oral cleft
SL standard length
uj upper jaw length
UPL greatest length of upper lobe of pectoral fin
All proportions are in percent of SL followed by percent
HL (in parentheses or specified). Paratype measurements
are in square brackets. For pectoral girdle methods and
discussion, see Andriashev & Stein (1998). Cephalic pores
were usually studied by injection, using the method of
Matsubara & Iwai (1954). We use the term “chin pores” for
the symphysiomandibular pore or pores.
Identification of many of the new Paraliparis species is
very difficult because they differ in details that are hard to
quantify; many of these characters have not previously been
used in liparid taxonomy. Therefore, we have expanded or
added to the descriptive terminology used in identifying
them for the snout, eye position, suborbital distance (defined
above), mouth position, lower jaw position, chin pores,
opercular flap, and body shape. Below, for each of these
characters, we provide descriptions of each character and
character state, figures where necessary, and list the new
species as they fit each character state. We provide
guidelines for use of relative terminology in the species
considered herein by listing the extremes for selected
proportions and naming the species displaying them (Table
2). Unfortunately, for some characters, judgement of state
is unavoidably highly subjective. Therefore, if there is any
doubt regarding the existing state in a specimen, it is
absolutely necessary to refer to the appropriate figure before
making a decision about its character state.
Snout (Fig. 2). We follow the traditional method of snout
measurement, i.e., from most anterior point of upper lip or
tip of snout to anterior margin of orbit (Hubbs & Lagler,
1949). Among the Australian Paraliparis, there is a wide
variety of snout shape, length, and depth. They are:
I Mouth horizontal
A Snout deep, blunt, not protruding anterior to
mouth: australiensis, csiroi, gomoni,
obtusirostris
B Snout deep, rounded, slightly projecting:
costatus, dewitti, hobarti, infeliciter,
tasmaniensis, sp. 2
Figure 2. Shape of snout. I, mouth horizontal: A, deep blunt, not protruding anterior to mouth; B, deep rounded,
slightly projecting; C, deep, bluntly rounded; D, deep, slanted, projecting; E, shallow, rounded, projecting; F,
abruptly angled; G, deep, rounded; H, acutely angled. I, deep, rounded, greatly projecting. II, mouth oblique: A,
deep, blunt; B, angled, projecting.
346 Records of the Australian Museum (2001) Vol. 53
C Snout deep, bluntly rounded: anthracinus, ater,
labiatus, piceus
D Snout deep, slanted, projecting: badius
E Snout shallow, rounded, projecting:
atrolabiatus, sp. 1
F Snout abruptly angled: auriculatus
G Snout deep, rounded: brunneus, coracinus,
impariporus
H Snout acutely angled: retrodorsalis,
tasmaniensis
I Snout deep, rounded, greatly projecting:
delphis, lasti, plagiostomus
II Mouth oblique
A Snout deep, blunt: avellaneus, eastmani
B Snout angled, projecting: brunneocaudatus
Sub rostral fold (Fig. 3). The subrostral fold is a fold of
skin (with gelatinous tissue internally) that forms part of
the ventral surface of the snout immediately anterior to the
Figure 3. Subrostral fold. I, absent; II, deep, covering upper lip
entirely; III, present, but not covering upper lip entirely; IV, deep
anteriorly, shallower posteriorly.
upper lip. If present, it may cover the upper jaw entirely or
to various degrees. In our Paraliparis specimens, we
identified four character states:
I Absent: anthracinus, badius, coracinus, csiroi,
hobarti, sp. 1, sp. 2
II Deep, covering upper lip entirely: avellaneus,
brunneocaudatus, plagiostomus
III Present, but not covering upper lip entirely:
A Almost absent, barely covering some of upper
lip: ater
B Covering about half of upper jaw: atrolabiatus,
dewitti, lasti, obtusirostris, retrodorsalis
C Covering about 3 A of upper jaw: costatus,
infeliciter
D Covering upper jaw almost, but clearly not
quite entirely: auriculatus, australiensis,
brunneus, eastmani, gomoni, impariporus,
tasmaniensis
IV Deep, covering most of upper lip anteriorly,
shallower posteriorly: delphis, labiatus, piceus
Eye position and suborbital distance (Fig. 4). The height
of the eye in relation to the profile of the head is a significant
character. It may actually enter the lateral profile and is
located at various distances above the mouth. We found
two categories of variation in our Paraliparis specimens,
each with two subcategories:
I Eye high, nearly touching dorsal contour of head
A Suborbital distance short (0.5-0.6 eye
diameter): australiensis, avellaneus, badius,
sp. 2
B Suborbital distance long (0.7-1.0 eye
diameter): brunneus, coracinus, delphis,
gomoni, obtusirostris, piceus, retrodorsalis,
tasmaniensis , sp. 1
II Eye low, not touching dorsal contour of head
A Suborbital distance short (0.5-0.6 eye
diameter): auriculatus, brunneocaudatus,
csiroi, eastmani, hobarti, impariporus,
infeliciter, lasti
B Suborbital distance long (0.7-0.9 eye
diameter): anthracinus, ater, atrolabiatus,
costatus, dewitti, labiatus, plagiostomus
Mouth position (Fig. 5). Two character states are
represented in our Paraliparis material: horizontal and
oblique. When the mouth is horizontal, the symphysis of
the upper jaw is clearly below eye level; when it is oblique,
the symphysis of the upper jaw is level with or above the
lower margin of the eye. A horizontal mouth may be
terminal, subterminal, or inferior. A terminal mouth is one
in which the symphyses of the upper and lower jaws are
located at the extreme anterior end of the fish, with the snout
not projecting; an inferior mouth is one located below a
greatly projecting snout, on the lower (flat) surface of the
head (similar to that in some sharks); a subterminal mouth
is intermediate between the two states, in which the snout
projects slightly, and the ventral surface of the head is
rounded, not flat, with the lower jaw usually, but not always,
deep below the posterior of the oral cleft.
Stein et al .: Australian liparids 347
Figure 4. Eye position and suborbital distance. I, eye high, nearly
touching dorsal contour of head: A, suborbital distance short (0.5-
0.6 eye diameter); B, suborbital distance long (0.7-1.0 eye
diameter). II, eye low, not touching dorsal contour of head: A,
suborbital distance short (0.5-0.6 eye diameter); B, suborbital
distance long (0.7-0.9 eye diameter).
Figure 5. Mouth position. I, mouth oblique. II, mouth horizontal
A, terminal; B, subterminal; C, inferior.
I Mouth oblique: avellaneus, brunneocaudatus,
eastmani
II Mouth horizontal:
A Terminal: australiensis, costatus, csiroi,
gomoni, labiatus, obtusirostris, piceus
B Sub terminal: anthracinus, ater, atrolabiatus,
auriculatus, brunneus, coracinus, delphis,
dewitti, hobarti, impariporus, infeliciter, lasti,
retrodorsalts, tasmaniensis, sp. 1, sp. 2
C Inferior: badius, plagiostomus
Lower jaw position (Fig. 6). We found five different lower
jaw character states in Paraliparis based on the relationships
of the tooth plates in both the upper and lower jaws when
the jaws are closed and viewed ventrally. We define
“included lower jaw” as one in which the upper tooth plates
can be seen clearly and completely from directly below
when the upper lip fold is pushed out of the way.
“Subterminal lower jaw” is one where the tooth plates of
the upper and lower jaws meet more or less completely, but
the upper jaw extends anterior to the lower jaw. “Terminal
lower jaw” is the case where the upper jaw does not project
anterior to the lower jaw. The “diastema” is the gap between
the left and right tooth plates, present or absent in either or
both jaws.
I Terminal: upper tooth plates exactly mating with
lower plates: labiatus
II Sub terminal: lower tooth plates somewhat behind
upper tooth plates, but clearly overlapping: ater,
atrolabiatus, costatus, hobarti, impariporus,
obtusirostris, piceus, plagiostomus, tasmaniensis
III Subterminal, almost included: lower tooth plates
barely overlapping posterior of upper tooth plates:
auriculatus, australiensis, brunneus, infeliciter , sp. 2
IV Included: lower tooth plates entirely within
348 Records of the Australian Museum (2001) Vol. 53
Figure 6. Lower jaw position. I, terminal: upper tooth plates
exactly mating with lower plates. II, subterminal: lower tooth plates
somewhat behind upper tooth plates, but clearly overlapping. Ill,
subterminal, almost included: lower tooth plates barely overlapping
posterior of upper tooth plates. IV, included: lower tooth plates
entirely within posterior margin of upper tooth plates. V, projecting:
lower tooth plates partially anterior to upper tooth plates.
posterior margin of upper tooth plates:
anthracinus, avellaneus, badius,
brunneocaudatus, coracinus, csiroi, delphis,
dewitti, gomoni, lasti, retrodorsalis, sp. 1
V Projecting: lower tooth plates partially anterior to
upper tooth plates: eastmani
Lower jaw depth (Fig. 7). The lower jaw can be deep or
shallow at the point below the posterior of the oral cleft. In
general, deep jaws are related to a rounded ventral cross-
section of the body, and shallow ones to a flat ventral
surface:
Figure 7. Lower jaw depth. I, jaw relatively deep. II, jaw relatively
shallow.
I Jaw relatively deep: anthracinus, atrolabiatus,
auriculatus, brunneocaudatus, brunneus,
coracinus, delphis, dewitti, eastmani, impariporus,
lasti, retrodorsalis, tasmaniensis, sp. 1, sp. 2
II Jaw relatively shallow: ater, australiensis, badius,
csiroi, gomoni, hobarti, infeliciter, labiatus,
obtusirostris, piceus, plagiostomus
Chin pores (Fig. 8). The mandibular symphyseal (chin)
pores are usually distinctly separated by an interspace. In
one species, P impariporus, the pores are fused to form a
single pore in which the canals enter from each side. In all
other species treated here, two (normal) pores are present.
We have found it necessary to expand the terminology
describing paired chin pores. Three states exist in our
Paraliparis material:
I At the surface: two pores are present, opening
directly and separately on the skin surface, or
opening into a poorly-defined shallow depression
not marked by an edge. Anterior skin fold absent,
anterior and posterior pores similar: avellaneus,
brunneocaudatus, costatus, eastmani, hobarti,
lasti, piceus, plagiostomus, retrodorsalis,
tasmaniensis
II With a crescent-shaped skin fold anterior to the
pores but not extending posterior to them. Pore
depression absent in anthracinus, ater,
atrolabiatus, auriculatus, badius, brunneus,
coracinus, delphis, labiatus ; only in one species
are pores in a poorly-defined, shallow depression
posterior to skin fold: obtusirostris
III In a wide, oval, shallow depression or pit: two
chin pores open into a common clearly-defined
shallow pit, with edges formed by skin folds;
A Interspace equal to about two pore diameters:
dewitti
Stein et al .: Australian liparids 349
Figure 8. Chin pores. I, at the surface: two pores are present,
opening directly and separately on the skin surface, or into a poorly-
defined shallow depression not marked by an edge; anterior skin
fold absent, pores similar in morphology to those located more
posteriorly on the canals. II, with a crescent-shaped skin fold
anterior to the pores but not extending posterior to them. Ill, two
chin pores open into a common wide, oval, shallow pit, with
sharply defined edges: A, interspace equal to about two pore
diameters; B, interspace equal to about one pore diameter or less
(one species with pit on anterior surface of symphysis). IV, a single
pore present.
B Interspace equal to about one pore diameter or
less: australiensis, csiroi (in a pit on anterior
surface of symphysis), gomoni, infeliciter, sp.
1, sp. 2
IV A single pore present: impariporus
Chin shape and chin pore position (Fig. 9). Shape and
structure of the chin (tip of the lower jaw) varies significantly
among the Paraliparis species. There are four states:
I In lateral profile, chin slanted at about 45°: ater,
auriculatus, avellaneus, badius, brunneocaudatus,
brunneus, coracinus, eastmani, impariporus,
infeliciter, lasti, plagiostomus, retrodorsalis,
tasmaniensis, sp. 1, sp. 2
Figure 9. Diagram of chin state and chin pore position. Arrow
points to left-hand chin pore. I, in lateral profile, chin slanted at
about 45°. II, chin profile distinctly right-angled, with sharp firm
edge; chin pores on ventral surface. Ill, chin profile vertical but
with rounded edge; chin pores on ventral surface. IV, chin margin
gelatinous, projecting anteriorly: A, chin pore-pit on anterior
surface, not ventral; B, chin pore pit on ventral surface.
II Chin profile distinctly right-angled, with sharp,
firm edge. Chin pores on ventral surface:
anthracinus, gomoni, labiatus, obtusirostris,
piceus
III Chin profile vertical with rounded edge:
atrolabiatus, australiensis, costatus, delphis,
hobarti
IV Chin margin gelatinous, projecting anteriorly
A Chin pore-pit on anterior surface, not ventral:
csiroi
B Chin pore pit on ventral surface: dewitti
’’Honeycomb” tissue occurs on the lower jaw of some
species but not in others; it looks like a thick subdermal
layer of honey-comb with empty “cells”. We suspect that
in life, these cells are filled with gelatinous material common
in liparids. We do not know the function of this tissue, nor
350 Records of the Australian Museum (2001) Vol. 53
Figure 10. Opercular flap. I, covering gill opening for its entire length: A, triangular with equal sides; B, sharp
tipped, dorsally notched at anterior base; C, tip blunt, dorsal notch absent. II, covering lower % of gill opening: A,
tip rounded; B, tip sharp; C, tip sharp, anterior dorsal notch prominent. Ill, covering lower half or less of gill
opening: A, tip triangular; B, tip rounded. IV, other variants: A, dorsally rounded; B, ear-shaped; C, rounded.
of analogous structures in other fishes. Its occurrence is
unrelated to depth of chin below the rear end of the mouth
cleft. It is clearly present in coracinus, impariporus,
infeliciter, labiatus, retrodorsalis, tasmaniensis , and sp. 1,
and absent in P. gomoni. The last was the only species with
damaged lower jaw skin in which honeycomb tissue was
absent. Because we wished to do as little damage as possible
to the specimens, we did not look for its presence in the
lower jaws of specimens that were unskinned, and therefore,
the condition in all species not listed above is unknown.
Opercular flap (Fig. 10). The opercular flap, which covers
the gill opening, although easily damaged during capture,
provides useful characters for identification. Shape,
presence or absence of a dorsal notch, amount of pectoral-
fin overlap, and position of the flap in relation to the gill
opening can all be useful in identifying species. Position of
the gill opening relative to the eye and orbit may also be
useful. For instance, the ventral end is even with or above
the upper margin of the pupil in P. plagiostomus and P.
badius, but is even with or below the ventral margin of the
eye in P. brunneocaudatus and P. costatus.
I Opercular flap covers gill opening for its entire
length.
A Triangular with equal sides: dewitti, sp. 2
B Sharp tipped, dorsally notched: anthracinus,
impariporus, sp. 1
C Ventral margin rounded: delphis
II Opercular flap covers lower % of gill opening.
A Tip rounded: avellaneus, hobarti, obtusirostris
B Tip blunt, dorsal notch absent: eastmani,
gomoni
C Tip sharp, anterior dorsal notch prominent:
coracinus, infeliciter, retrodorsalis,
tasmaniensis
III Opercular flap covers lower half or less of gill
opening.
A Tip triangular: ater, lasti
B Tip rounded: costatus
IV Other variants
A Dorsally rounded: piceus
B Anthropoid ear-shaped: auriculatus, australi-
ensis, brunneocaudatus, csiroi, labiatus
C Rounded: atrolabiatus, badius, brunneus,
plagiostomus
Stein et al .: Australian liparids 351
Costal ridge (Fig. 11). Several of our new species have a
previously undescribed character state of the epineural and
epipleural ribs (pleural ribs are absent in Paraliparis ).
Although these ribs are present in all species we examined,
they are usually relatively short and slender and do not reach
the surface of the body musculature. In P. costatus, P. dewitti,
P. lasti, sp. 2, and in the damaged specimens of Paraliparis
sp. (cf. copei group), the ribs are long, well developed, and
reach the muscle surface along the anterior dorsal edge of
the body cavity. Their tips can be clearly felt, and form a
serrated ridge. In a number of other species ( ater ;
auriculatus, atrolabiatus, australiensis, brunneus,
coracinus, delphis, hobarti, retrodorsalis), this ridge is
barely present owing to less well-developed ribs. In all other
species, it is absent. Therefore, to determine the state of
development of these ribs in most species, radiographs are
necessary.
Nephrohaemal canal (Fig. 12). In most liparids, the
parapophyses of the abdominal vertebrae are fork-like and
unfused, except for those on the last one or two abdominal
vertebrae, where they are joined to form a short haemal
spine that does not reach the anal-fin pterygiophores
(Andriashev & Stein, 1998). Although we were unable to
dissect specimens owing to scarcity of material, radiographs
clearly showed that in four species (ater, atrolabiatus, csiroi,
delphis ) the parapophyses of abdominal vertebrae 3-11 and
5-11 respectively are joined distally, creating foramina
which in series form a canal. This structure also occurs in
piceus but was only visible on vertebrae 9-11. These species
do not appear to be close relatives, because there are
differences among them in chin pore arrangement and
number of pectoral radials. A similar structure was described
in Pholidae (Andriashev, 1954; Hubbs, 1927: 388;
Makushok, 1958) which included not only blood vessels
but kidney tissue, which they named the “haemonephra-
pophyses” and “nephrohaemal canal”. Because the structure
in these Paraliparis species seems to be the same, we use
the same term for it. Its occurrence in Pholidae is presumably
related to the compression of the body, but in many other
similarly-shaped fishes the parapophyses are normal
(Chabanaud, 1951). Analogous structures are also known
in Scombridae (Kishinouye, 1923: 338), Zeusfaber (Ford,
1937), and some Pleuronectiformes (Chabanaud, 1951).
Figure 11. Epineural and epipleural ribs on vertebrae 3, 5, and 7. I, Paraliparis australiensis, holotype, NMV
A21497, $, 176 mm TL, 164 mm SL. II, Paraliparis costatus, holotype, CSIRO H561-02, S, 224 mm TL, 204 mm SL.
352 Records of the Australian Museum (2001) Vol. 53
Figure 12. Nephrohaemal canal (from Makushok, 1958). I —
vertebral structure of Pholis guntielus\ a, lateral view; b, frontal
view; epp, epipleural; hnph, haemonephrapophysis; ill, iv, xvn,
numbers of vertebrae; paph, parapophysis; przph, praezyga-
pophysis. II —schematic of abdominal cross-section of Pholis
pictus; a.d., aorta dorsalis; cut, skin; epp, epipleural; gon, testes;
in, intestine; m, mesentery; m.corp., abdominal musculature; ren,
kidney; sq, scales; t.n., spinal cord; v.c., vena cava; XIX . 19th
vertebra. Dorsal musculature not shown.
Figure 13. Body shape. I, elliptic, shallow; II, elliptic, deep; III,
dorsal profile straight; IV, ventral profile straight, not dorsally
humpbacked; V, ventral profile straight, dorsally humpbacked; VI,
ventral profile straight, body leaf-like, compressed; VII, dorsal
and ventral contours curved, dorsal much more so, body
humpbacked, not compressed.
Body shape (Fig. 13). Body shape can be useful only in
specimens that have not been badly damaged or contorted
during capture or preservation.
I Elliptic, shallow: ater, brunneus, costatus, gomoni,
hobarti, infeliciter?, retrodorsalis, sp. 2
II Elliptic, deep: anthracinus, australiensis, labiatus,
tasmaniensis
III Dorsal profile straight: avellaneus,
brunneocaudatus, eastmani, sp. 1
IV Ventral profile straight, not dorsally humpbacked:
Stein et al .: Australian liparids 353
auriculatus, atrolabiatus, badius, impariporus
V Ventral profile straight, dorsally humpbacked:
coracinus, delphis, dewitti, lasti, plagiostomus
VI Ventral profile straight, body leaf-like,
compressed: csiroi, piceus
VII Dorsal and ventral contours curved, dorsal much
more so, body humpbacked, not compressed:
obtusirostris
Species groups. To facilitate identification, we have divided
the new Paraliparis species into three groups based on
mouth position and chin pore characters. These groups do
not necessarily represent phylogenetic relationships or real
genetic similarity.
I Mouth oblique: avellaneus, brunneocaudatus,
eastmani
II Mouth horizontal and inferior: badius,
plagiostomus
III Mouth horizontal, terminal or subterminal with
symphyseal chin pores in three states:
a Not in a pit or depression and lacking anterior
skin fold: costatus, hobarti, impariporus, lasti,
piceus, retro dorsalis, tasmaniensis
b In a pit: australiensis, csiroi, dewitti, gomoni,
infeliciter, obtusirostris , sp. 1, sp. 2
c With an anterior skin fold: anthracinus, ater,
atrolabiatus, auriculatus, brunneus, coracinus,
delphis, labiatus
We describe and discuss each species within this context.
Owing to the great similarities of some of these species,
our key is not a full field key. In many cases, it will be
necessary to perform dissections, use a dissecting
microscope, radiograph, or to clear and stain certain
structures.
Key to genera of Australian liparids
1 Ventral sucking disk present. Careproctus Krpyer, 1862
-Ventral sucking disk absent. 2
2 Clearly humpbacked at occiput. Dorsal-fin rays 50 or fewer. Upper
nasal pore (n 0 ) above or posterior to nostril. Mouth always oblique
. Psednos Barnard, 1927
Not humpbacked at occiput, but may be humpbacked more
posteriorly. Dorsal-fin rays 57 or more. Upper nasal pore (n 2 )
anterior to nostril. Mouth usually horizontal. Paraliparis Collett, 1878
Species descriptions
Genus Careproctus Krpyer, 1862
Careproctus Krpyer 1862: 253 (type species Liparis reinhardti
Krpyer, by monotypy).-Garman, 1892: 71; Jordan & Evermann,
1898: 2129; Burke, 1912: 507; 1930: 95; Stein, 1978b: 75;
Kido, 1988: 192; Andriashev, 1998: 258; Andriashev & Stein,
1998: 5.
Diagnosis. One pair of nostrils (a single nostril on each
side of snout). Pseudobranch absent. Ventral sucking disk
present, small to large. Pectoral fins with or without
developed lower lobe, notch variable, typically with fewer
rays than anal fin. Body colour not variegated. About 45
Southern Hemisphere species known, 30 from the Southern
Ocean (Andriashev & Stein, 1998), about 15 from the cool
temperate Pacific and Atlantic coasts of South America
(Andriashev & Stein, 1998), and one from Australia (this
paper).
Comparative notes. The trenchant differences between the
genus Careproctus and other liparid genera having ventral
sucking disks, the most speciose of which is Liparis, are
absence of pseudobranchiae and presence of single nostrils
(v. a pair on each side of the snout). In addition, many
Careproctus have more pectoral-fin rays than dorsal or anal-
fin rays and have simple teeth whereas the other genera
tend to have more longitudinal than pectoral-fin rays, and
have trilobed teeth. Generally, Careproctus occurs at greater
depths than the others, and is distributed worldwide at depths
from the intertidal to over 6000 m, whereas they do not
occur in the Southern Hemisphere. Its morphological
variability, represented by reduction in body structure,
reflects its great depth range (Andriashev & Stein, 1998).
Careproctus paxtoni n.sp.
Fig. 14
Material examined. Holotype AMS 129737-003, 9 , 150
mm TL, 134 mm SL. FRV Kapala, 35°29’S 150°55'E, New
South Wales, E of Brush I., 1116-1134 m, 31 Aug. 1989;
radiograph 29737-1; pectoral girdle 725. Paratype AMS
129802-001, 9, 155 mm TL, 144 mm SL; LRV Kapala,
35°28.5'S 150°53.5'E, New South Wales, SE of Ulladulla,
1061-1088 m, 30 May 1989; radiograph 29737-003;
pectoral girdle 726.
Diagnosis. Vert. 58-59 (10+48-49), D 53-54, P 34-35, C
10-11 (1+4/5+0-1), radials 4 (3+1), notched, three
interradial fenestrae present. Pleural ribs present. Hypural
plate single, unslit. Mouth terminal. Teeth shouldered,
tricuspid, not sharp. Two suprabranchial pores present. Gill
opening entirely above pectoral-fin base. Ventral disk very
small, about 3% SL. Head 23-24%, preanal 45-46%.
Peritoneum black, stomach pale to dusky.
Further description. Counts: D 53 [54], A 46 [47], P 34
[35], C 10 [11], Vert. 58 [59], radials 4 [4] notched,
354 Records of the Australian Museum (2001) Vol. 53
Figure 14. Careproctuspaxtoni n.sp. A, holotype, AMS 129737-003, 9,150 mm TL, 134 mm SL; B, ventral view;
C, teeth; D, P 725, cleared and stained right pectoral girdle.
interradial fenestrae 3 [3], pores 2-6-7-2, gr 8 [9], pc 13.
Ratios: HL 23.7 [22.8], its width 18.3 [—■], its depth 23.7
preD 28.2 [27.8], preA 45.1 [46.2], disk 3.0 [3.1], md
7.9 [9.2], ma 11.5 [—], UPL 14.9 [16.2], NL 9.0 [—], LPL
10.4 [9.9], E 6.2 [5.4], uj 9.9 [9.0], lj 8.6 [8.9], gs 9.0 [—];
in %HL: UPL — [63], disk 12.7 [13.5], E 26.2 [23.7], sn
26.3 [28.0], uj 41.8 [39.2], lj 36.4 [38.9], gs 38.0 [—].
Head moderately large, its dorsal profile sloping straight
to the blunt, high snout, the anterior profile of which is
nearly vertical. Head width about 80% HL. Interorbital
space broad, slightly convex in life, flattened in preserv¬
ation; dorsal margin of orbit not entering dorsal profile.
Mouth terminal, horizontal, snout not protruding beyond
upper lip. Oral cleft reaching to below anterior margin of
orbit, tip of maxilla extending to below mid-eye. Teeth
shouldered and trilobed, middle lobe largest, tips not sharp;
arranged in 12 [14] and 13 [14] irregular oblique rows in
both jaws, 5-7 teeth in a row anteriorly; innermost teeth
largest. No diastema present at junction of upper or lower
jaws. Eye relatively large, about 24-26% HL. Nostril with
distinct short tube. Circumoral pores rather small, round to
oval. Chin pore pair normally spaced, interspace pnq-pnq
nearly equal to pnq-pmo. Suprabranchial pores two, the
second slightly raised. Gill opening of holotype above
pectoral base, extending from level of upper margin of eye
to or just in front of dorsalmost pectoral ray, damaged on
both sides of paratype. Opercular flap very small, rounded.
Upper pectoral lobe rather short, not nearly reaching anal-
fin origin; pectoral rays shortened ventrally to 25 [26]th
ray which is a little shorter than half of upper lobe length.
Pectoral-fin notch shallow but distinct. Lower pectoral lobe
of 9 rays, short, with free tips. Uppermost pectoral ray level
with middle of suborbital space. Lowermost pectoral ray
below anterior margin of orbit. Basal cartilaginous lamina
of pectoral girdle narrow. Radials 4 (3+1), notched. Three
interradial fenestrae present.
Body moderately elongated, greatest depth about equal
to HL. Ventral disk very small, its length about one-half
eye diameter; anterior lobe not developed; disk centre
slightly depressed, surrounded by a very narrow, thin,
delicate, smooth, upturned margin. Anus very close to
(almost immediately behind) disk. Behind anus, a rounded
area of thick, compact tissue present, surrounded by a low
skin fold. Genital opening on anterior part of rounded area,
a small (probably urinary) papilla immediately posterior to
opening. Anterior dorsal rays shortened, not rudimentary.
Interneural of first dorsal ray between fourth and fifth neural
spines; one free (rayless) interneural present anteriorly.
Three thin pleural ribs originate on parapophyses of 8th,
9th and 10th vertebrae, the last two longer, sabre-like, curved
ventrally. Hypural plate single, with a trace of suture. Anal-
fin origin below 12th vertebra. Caudal fin of 10 [11] rays, 9
principal, holotype with one small procurrent ray above
(1+4/5) and paratype with one also below [ 1+4/5+1].
Posterior dorsal- and anal-fin rays apparently overlapping
about one-third caudal-fin length. Skin thin, fragile, prickles
absent. Gelatinous tissue moderately developed. Pyloric
caeca thick, similar in size, length 5-6% SL, located right
Stein et al .: Australian liparids 355
ventrally to stomach and unusually far anterior relative to
its fundus.
Colour. Body dull translucent brown; orobranchial cavity
dusky, peritoneum black. Stomach pale, finely reticulated
with black, appearing dusky.
Distribution. Off New South Wales at 1061-1134 m.
Careproctus paxtoni is the only species of the genus known
from Australia.
Biology. Both specimens are females with eggs close to
being ripe, 3.7 and 3.3 mm diameter respectively, in addition
to very small oocytes. The unusual postgenital area of thick,
compact tissue with an associated skin fold in both
specimens may be related to spawning and may thus be a
temporary phenomenon. Its function is unknown.
Etymology. Named after John R. Paxton, formerly Principal
Research Scientist in the fish collection at the Australian
Museum, Sydney, in honour of his many contributions to
knowledge of Australian and deep-sea fishes.
Comparative notes. Careproctus paxtoni belongs in
Careproctus s. str. (see Andriashev, 1998) because it has 4
(3+1) notched pectoral radials, with three interradial
fenestrae. This and other character states and its distribution
place it in the Careproctus group around the Antarctic,
including species from South Africa, Argentina, New
Zealand, and the South Shetland Is. Of these species it is
most similar to C. novaezelandiae Andriashev but differs
in having two suprabranchial pores (v. one), blunt teeth,
pale to dusky stomach (v. black), fewer pectoral rays (34-
35 v. 37-38), unslit hypural plate (v. divided) and a smaller
disk (3.0-3.1% v. 6.7% SL). In addition, postgenital
morphology like that of C. paxtoni has not been described
for any other species, although that may be a function of
temporary existence only at spawning times or having been
overlooked.
Genus Psednos Barnard, 1927
Dwarf snailfishes
Figs. 15, 16
Psednos Barnard, 1927: 927 (type species micrurus Barnard).-
Andriashev, 1992: 3 (redescription of the genus).
Paraliparis (non Collett) Stein, 1978a; 1986: 493.
Diagnosis. Ventral disk absent. No pseudobranchiae. One
pair of nostrils. Mouth oblique, superior or terminal. Opercle
very long. Infraorbital sensory canal (c.io) widely
interrupted behind eye, consisting usually of six (5+1) or
five (5+0) pores (Fig. 15). Nasal pores widely spaced, the
upper nasal pore (n 2 ) opening higher and behind vertical
through nostril. Coronal commissure (C.) with or without a
coronal pore (cor). Suprabranchial pore (t sb ) very widely
spaced from top of gill slit. Preoperculo-mandibular pores
(pm) 6-7. Pectoral fin of 13-17 rays. Caudal fin usually of
6 rays. Pectoral girdle with 3 radials, rounded or notched.
Interradial fenestrae present or absent. Vertebrae 41-58.
Pleural ribs absent. Hypural plate single, unslit. Body
distinctly hump-backed, high at occiput. Anterior abdominal
vertebrae and base of cranium forming an arch at about 90°
(Fig. 16). At least about 30 poorly known species from
temperate and subtropical seas of Southern and Northern
Hemispheres. Five or six species known from the northern
part of the Southern Ocean. Probably mesopelagic or
epibenthic.
Comparative notes. The genus Psednos differs from
Paraliparis and other diskless genera of liparids in having
a number of distinctive features: the morphology of the
sensory canal system is unique in having nasal pores more
dorsally located and the infraorbital canal interrupted behind
the eye; the distinctly hump-backed body because of the
sharply curved anterior of the vertebral column; and the
pectoral girdle with 3 radials.
tsb
i
N
n 2
cor
Figure 15. Psednos. Diagram of cephalic sensory canal system and pores. I, lateral view. II, dorsal view. C, coronal
commissure; cor, coronal pore; io, infraorbital pores (io 6 = supraorbital pore); n, nasal pore; N, nostril; pm,
preoperculo-mandibular pores; t, temporal pores; t sb , suprabranchial pore (= t 4 ).
356 Records of the Australian Museum (2001) Vol. 53
Figure 16. Psednos micrurus, holotype, BMNH 1930.1.14.7, 9,45 mm SL. Radiograph showing typical “hump¬
backed” vertebral column. Radiograph 17451.
Key to Southern Ocean Psednos
1 Vertebrae 42-43. Anal-fin rays 28-31. Pectoral radials 3, rounded,
equidistant, unnotched. Interradial fenestrae absent... Psednos micrurus Barnard (South Africa)
-Vertebrae 56-58. Anal-fin rays more than 40. Pectoral radials 3,
not equidistant (2+0+1), notched. One or more interradial fenestrae
present. 2
2 Gill opening short, 24% HL, less than eye diameter, entirely above
pectoral base. Eye 6.3% SL (27.6% HL). Psednos sp.
-Gill opening longer, 31-35% HL or 1.6-1.8 eye diameter, reaching
ventrally in front of 2-5th pectoral ray. Eye 4.1-5.2% SL (19-
21.2% HL). 3
3 Peritoneum light brown, stomach dusky (not black). Tongue pale.
Gill opening reaching ventrally to 2nd pectoral ray. Rays of lower
pectoral-fin lobe shorter than those of upper pectoral-fin lobe. Psednos balushkini n.sp.
-Peritoneum black, stomach black. Tongue black-dotted or dusky.
Gill opening reaching to 4-5th pectoral ray. Rays of lower pectoral-
fin lobe slightly longer than those of upper pectoral-fin lobe. 4
4 Opercular lobe very weakly developed, indistinct (Pig. 18). Depth
of head at occiput about 133% HL. Pyloric caeca of different
lengths, 5-9.5% SL. All gill arches dark-dotted. Psednos nataliae n.sp.
-Opercular lobe small but distinct, prominent as an angle (Pig. 19).
Depth of head at occiput about 106% HL. Pyloric caeca of similar
lengths, 3.5-4.5% SL. Gill arches undotted. Psednos whitleyi n.sp.
Psednos balushkini n.sp.
Fig. 17
Material examined. Holotype AMS 124860-002, S , 90
mm TL, 84 mm SL. PRV Kapala, fid. no. K 84-17-05,
34°48.5'S 151°15.5'E, E of Shoalhaven Heads, New South
Wales, 914-960 m, 4 Oct. 1984; radiograph ZISP 20032;
pectoral girdle 728. Paratype AMS 124059-018, S, 92 mm
TL, 82 mm SL. PRV Kapala , Lid No. K 83-09-02, 33°32'S
152°09'E, off Norah Head, New South Wales, 942-978 m,
23 Aug. 1983; radiograph ZISP 20032; pectoral girdle 729.
Diagnosis. Vert. 57 (11-12+45-46), D 49-50, P 17 (9-
10+1-2+6), C 6, radials 3 (2+0+1), notched with 2 fenestrae
(fl, f2). Depth at occiput 111-123% HL. Gill opening
extending ventrally in front of two pectoral rays. Opercular
lobe small but distinct. Rays of lower pectoral-fin lobe
shorter than in the upper. Pyloric caeca of similar length, c.
6% SL, slightly dotted at bases. Pores n 2, i.o. 5+1, t 1+1,
cor. absent. Orobranchial cavity pale scarcely dotted, tongue
pale, peritoneum light brown or paler. Gill arches not dotted.
Further description. Counts: D 49 [50], A 43 [43], P 17
[17], C 6 [6], Vert. 57 [57], radials 3 (2+0+1), two interradial
Stein et al .: Australian liparids 357
Figure 17. Psednos balushkini n.sp. A, holotype, AMS 124860-002, 3, 90 mm TL, 84 mm SL. B, P 728, cleared
and stained right pectoral girdle; fin rays reconstructed from left side.
fenestrae, gr 10 [9], pc 10 [8]. Ratios: HL 23.8 [24.4], its
width 11.9 [12.2], depth at occiput 29.2 [27.1], bdA 18.1
[17.4], preD 31.0 [29.3], preA 37.5 [40.2], ma 17.9 [18.5],
aAf 22.0 [22.0], UPL 15.5 [14.9], NL 6.0 [6.7], LPL 13.1
[12.1], io 10.7 [10.4], E 5.0 [5.2], sn 7.1 [8.5], uj 11.7 [12.1],
gs 8.3 [torn]. In % HL: depth at occiput 123 [111], hd width
50 [50], UPL 65 [61], io 45 [42.5], E 21 [21], sn 30 [35], uj
49 [50], gs 35, pc 6.
Head high, its upper profile forming at occiput an angle
of about 100° with rounded top. Depth at occiput 1.2 [1.1]
HL, 1.6 [1.6] depth at anal-fin origin. Head width 0.41 [0.45]
its depth. Area from upper lip to crest of occiput abrupt,
distance between them equal to HL. Mouth superior,
distinctly oblique, forming an angle of about 40° to the
horizontal. Lower jaw projecting, with a tooth-like knob
on outside of symphysis but absent inside. Tooth plates
narrow, with 19-23 oblique rows, 5-6 teeth per row
anteriorly. Narrow diastema present in upper jaw. Posterior
end of lower jaw forming a distinct ventrally-directed angle.
Nostril level with lower margin of eye, with slightly raised
rim. Eye not large. Gill opening slightly oblique, originating
below horizontal through eye centre, reaching ventrally in
front of second pectoral-fin ray. Opercular lobe not large,
but distinct with rounded tip. Infraorbital canal widely
interrupted behind eye; infraorbital pores 6 (5+1), the last
on a horizontal through upper margin of eye. Coronal pore
absent. Upper nasal pore (n 2 ) opening above and behind
nostril. Preoperculo-mandibular pores six; pm 6 on horizontal
with pupil of eye. Postorbital pore (t 2 ) situated high, above
pore i.o. 6 . Suprabranchial pore single, opening well above
and ahead of dorsal end of gill opening, at distance of about
55% HL from it. Chin pores normally spaced, pmj-pmj
about equal to pmj-pm^
Pectoral fin distinctly notched; upper pectoral lobe not
quite reaching anal-fin origin, consisting of 10 [9] rays;
notch ray single [2], 38% of upper lobe length. Lower
pectoral lobe of 6 rays, slightly shorter than the upper one.
Uppermost pectoral ray level with posterior end of mouth
cleft. Basal cartilaginous lamina of pectoral girdle uniformly
wide, with 3 (2+0+1) radials, two upper radials and scapula
notched, including two rounded interradial fenestrae (fl, f2).
Body distinctly hump-backed, dorsal outline sloping
gently from high occiput caudally; depth at anal-fin origin
62.5% of depth at occiput. Angle of anterior four or five
vertebrae with base of cranium a little less than 90°.
Interneural of first dorsal ray between sixth and seventh
vertebrae (between fifth and sixth in paratype); one free
(not bearing a ray) interneural present anteriorly. Trunk part
of body rather large, more than 2 A SL. Distance from
mandibular symphyseal knob to anus less than interspace
between anus and anal-fin origin. Hypural plate single,
unslit. Caudal fin of 6 (3/3) rays. Skin thin, loose,
subcutaneous gelatinous tissue well developed. Pyloric
caeca about the same length and size. Small urogenital
papilla present.
Colour. Skin white. Orobranchial cavity pale, partially with
small sparse dots. Tongue pale anteriorly, with small
sparsely scattered dots centrally. Gill arches undotted.
Peritoneum light brown (lighter in paratype). Stomach
dusky with reticulated pattern of melanophores extending
over anterior part of intestine, resembling network of black
blood vessels. Pyloric caeca pale, sparsely dotted at bases.
Distribution. Off New South Wales at 914-978 m.
Etymology. The new species is named after Dr Arkady V.
Balushkin of the Zoological Institute, Russian Academy of
Sciences, St. Petersburg, in honour of his many contributions
to the knowledge of Antarctic fishes, especially to the
taxonomy of the suborder Notothenioidei.
Comparative notes. Psednos balushkini differs from other
Australian species of the genus Psednos in having a pale
orobranchial cavity (v. dark or dotted) and a light brown
peritoneum (v. black). Lor more detailed descriptions of
differences, see descriptions of P. nataliae and P. whitleyi.
Psednos nataliae n.sp. Stein & Andriashev
Pig. 18
Material examined. Holotype CSIRO H2636-04, 3 ,
unknown TL, 98.5 mm SL. PRV Soela, stn. So 2/89/80,42° 16'S
144°39'E, SW of Cape Sorell, Tasmania, 1100-1120 m, 18
Mar. 1989; radiograph 20174; pectoral girdle 730.
Diagnosis. Vert. 58 (12+46), D 50, P 16 (8+2+6), radials 3
(2+0+1), notched with single fenestra (f2). Depth of occiput
133% HL. Gill opening extending ventrally in front of about
fourth to fifth pectoral-fin rays. Opercular lobe very weakly
developed, hardly discernible. Lower pectoral-fin rays
110% upper lobe rays. Pores n 2, i.o. 5+1, pm 6,11+1, cor.
358
Records of the Australian Museum (2001) Vol. 53
Figure 18. Psednos nataliae n.sp. A, holotype, CSIRO H2636-
04, 8 , unknown TL, 98.5 mm SL. B, P 730, cleared and stained
right pectoral girdle.
absent. Tongue densely black-dotted. Orobranchial cavity
dark, closely dotted with melanophores, as are gill arches.
Peritoneum, stomach, and anterior part of intestine black.
Pyloric caeca different lengths, 5-9.5% SL, all dark-dotted.
Further description. Counts: D 50, A 42, P 16, C missing,
Vert. 58, radials 3, fenestra single (f2), gr 9, pc 7. Ratios:
HL 21.3, its width 10.2, depth at occiput 28.4, bdA 21.3,
preD 28.9, preA 35.5, ma 17.3, aAf 21.3, UPL 14.7, NL
7.6, LPL 16.2, io 9.1, E 4.1, sn 7.1, uj 11.4, gs 7.3, pc 5-
9.5. As percentage HL: depth of head 133, UPL 69, LPL
76, io 43, E 19.0, sn 33.3, uj 53, gs 34.5.
Dorsal profile of snout and head rising very steeply at
an angle of 70-80°, forming at occiput a high hump forming
an included angle of about 90°, with broadly rounded top.
Head short, compressed, its width 36% of head depth; the
latter exceeds 130% HL. Abrupt distance between upper
lip and top of occiput about equal to HL. Mouth superior,
very oblique, angle of 40-45° to a horizontal midline.
Interorbital space relatively broad, slightly convex.
Symphyseal knob present. Teeth simple, small, slender,
sharp canines, in each jaw forming a narrow band of 20
and 17 irregular oblique rows, uniserial posteriorly, of 5-6
teeth in each anterior row. Wide diastema present at
premaxillary joint. Teeth in lower jaw similar, diastema
absent. Posteroventral corner of lower jaw forming an angle
protruding ventrally below lower outline of head. Eye
relatively small; dorsal margin of orbit near profile of head
but not entering into it. Nostril damaged. Gill opening origin
slightly below horizontal through ventral margin of eye,
Opercular lobe very weakly developed, forming a broad
convex arc, tip of opercle projecting only slightly to form a
shallow rounded protrusion. Infraorbital canal widely
interrupted behind eye; pores 6 (5+1), i.o. 6 above level of
upper margin of eye and far from its posterior margin (at a
distance of two eye diameters). Postorbital pore (tj) above
io 6 , coronal pore absent. Nasal pores widely spaced, the
upper pore (n 2 ) situated above and behind nostril.
Suprabranchial pore single, high above dorsal end of gill
opening and in front of it at a distance equal to 50% HL.
Preoperculo-mandibular pores 6. Chin pores regularly
spaced; all pores small, hardly discernible.
Pectoral-fin notch moderately deep; its upper lobe of 8
rays; two notch rays about half as long as upper pectoral
lobe. Lower lobe of 6 rays, a little longer than upper lobe
rays. Basal cartilaginous lamina of pectoral girdle uniformly
wide, with 3 (2+0+1) radials, two upper notched and
including one oval fenestrae (f2). Lowermost radial largest.
Scapula unnotched, with very small helve. Coracoid with
elongated helve and small opening.
Body strongly hump-backed, dorsal outline rising
abruptly to prominent hump and tapering rapidly posterior
to it to a point above end of abdominal cavity, then evenly
to caudal. Depth of occiput 140% depth at anal-fin origin.
Vertebral column strongly curved, forming nearly right
angle with base of skull. Interneural of first dorsal-fin ray
between sixth and seventh neural spines; one free (rayless)
interneural present anteriorly between 5th and 6th neural
spines. Trunk not large, slightly more than 33% SL. Distance
from mandible to anus 83.3% of distance between anus and
anal-fin origin. Caudal fin unknown; hypural plate and all
rays missing. Skin thin, naked, loose. Subcutaneous
gelatinous tissue obviously well developed.
Colour. Body white. Pale tongue with strongly contrasting
black dots. Orobranchial cavity dark, closely dotted with
melanophores. Peritoneum, stomach and anterior part of
intestine black. Pyloric caeca and all gill arches dark-dotted.
Distribution and biology. West coast of Tasmania at 1100-
1120 m. The specimen is an adult male with ripe testes and
a long tubular urogenital papilla. The anus and genital area,
with the adjoining part of the belly, are directed ventrally
and forward in spawning condition.
Etymology. The new species is named after Dr Natalia V.
Chernova, Zoological Institute, Russian Academy of
Sciences, St. Petersburg, in honour of her contributions to
knowledge of Arctic liparids and other fishes.
Stein et al .: Australian liparids 359
Figure 19. Psednos whitleyi n.sp. A, holotype, CSIRO H1335-
02, 6 , 105.4 mm TL, 91.2 mm SL. B, P 727, cleared and stained
right pectoral girdle.
Comparative notes. Psednos nataliae n.sp. differs from
the other two known species of Australian Psednos in having
a weakly developed opercular lobe and dark-dotted gill
arches. Furthermore, it differs from P. balushkini in having
a black peritoneum (v. light brown), a dark orobranchial
cavity (v. pale, scarcely dotted) and a lower pectoral-fin
lobe slightly longer than the upper. It differs from P whitleyi
by its very high occiput (133% HL v. 106%).
Psednos whitleyi n.sp.
Fig. 19
Material examined. Holotype CSIRO H1335-02, S, 105.4
mm TL, 91.2 mm SL. FRV Soela, stn. So 3/86/28, 41°18.4'S
144°05'E, NW of Kenneth Bay, Tasmania, 900-920 m, 14 May
1986; radiograph 20173; pectoral girdle 727.
Diagnosis. Vert. 56 (11+45), D 48, P16 (8+2+6), C 6, radials
3 (2+0+1), two upper radials notched, a large fenestra (f2)
between them. Depth at occiput 106% HL. Gill opening
extending ventrally in front of 4-5 pectoral-fin rays.
Opercular lobe small but distinct, protruding as a right angle.
Tongue light dusky. Orobranchial cavity dusky, black-
dotted. Peritoneum and stomach black, anterior part of
intestine dark brown to black. Pyloric caeca short, similar
in length, 3.5-4.5% SL, all dark-dotted. Gill arches without
melanophores. Pores n 2, i.o. 5+1, pm 6,10+1, cor. absent.
Further description. Counts: D 48, A 41, P 16, C 6, Vert.
56, radials 3 (2+0+1), notched, one interradial fenestra (f2),
gr 8, pc 8. Ratios: HL 24.8% SL, its width 11.0, its depth at
occiput 26.3, bdA 19.7, preD 30.7, preA41.1, ma 18.6, aAf
23.0, UPL 14.8, LPL 15.6, io 9.9, E 4.7, sn 7.5, uj 12.3, gs
7.7. As percentage HL: depth of head 106, bdA 80, UPL
60, uj 50, io 40, sn 30.1, E 19, gs 31.
Head comparatively large, compressed, its width about
44.2% its length, its depth slightly greater than its length.
Anterior profile forming a right angle with tip of lower jaw
at apex. Interorbital space broad, slightly convex. Mouth
distinctly superior, very oblique, about 45° to horizontal.
Lower jaw protruding before upper lip; prominent knob
present at joint between dentaries. Posteroventral corner of
lower jaw forming a sharp, protruding 90° angle directed
below centre of eye. Teeth small, slender canines arranged
in about 26 and 21 irregular oblique rows in jaws forming
narrow bands 5-6 teeth per row anteriorly. Wide diastema
in premaxilla; almost no dentary diastema. Eye moderately
small, dorsal margin of orbit well below dorsal outline of
head; pupil about half eye. Gill opening distinctly oblique,
slightly exceeding eye, reaching ventrally in front of 4-5
pectoral-fin rays, its dorsal end about level with lower
margin of eye. Opercular lobe not large, well developed in
upper part of gill opening as nearly right angle; supported
dorsally by narrow, sharply pointed opercle which is
noticeably curved ventrally. Infraorbital canal widely
interrupted behind eye, of 6 pores (5+1). Nasal pores widely
spaced, second pore (n 2 ) opening high above and behind
vertical through nostril. Small tube present in place of
coronal pore; pore itself not discernible. Supraorbital pore
single, high above and anterior to upper end of gill opening,
at a distance equal to 38% HL. Postorbital pore (t,) not found
owing to damaged skin.
Preoperculo-mandibular pores 6, anterior pair normally
spaced.
Level of uppermost pectoral-fin ray below orbit, about
even with tip of suborbital stay, midway between orbit and
posterior of maxilla. Pectoral-fin base angled forward
ventrally. Fin divided into two lobes by moderately deep
notch; rudimentary rays absent. Upper lobe of 8 rays not
reaching anal-fin origin. Two notch rays present; lower lobe
slightly longer than upper. Basal cartilaginous lamina short
and wide, with three (2+0+1) radials, the two uppermost
notched with an oval fenestra between them (f2). Scapula
unnotched, one rudimentary fenestra (fl) present ventrally.
Body moderately hump-backed, depth at occiput 133%
depth at origin of anal fin. Maximum depth at occiput about
on vertical through lower end of gill slit. Trunk tapering
evenly behind abdominal cavity to caudal fin. Interneural
360 Records of the Australian Museum (2001) Vol. 53
of first dorsal-fin ray between 5th and 6th neural spines.
Caudal fin of 6 (3/3) rays, overlapped about 40% by dorsal-
and anal-fin rays. Skin very thin, fragile, easily torn;
holotype partly skinned. Pyloric caeca all of similar length,
3.5-4.5% SL.
Colour. Body white except for dark abdominal cavity
showing through wall. Tongue light dusky. Orobranchial
cavity, tongue and pyloric caeca dotted with small
melanophores. Gill arches not dotted. Peritoneum and
stomach black, anterior part of intestine dark brown to black.
Distribution. Off NW Tasmania between 900 and 920 m depth.
Etymology. The new species is named in honour of Gilbert P.
Whitley (1903-1975), well known Australian ichthyologist,
formerly Curator of the fish collection in the Australian
Museum, Sydney.
Comparative notes. Psednos whitleyi n.sp. is similar to P.
nataliae in having a black peritoneum and stomach and the
lower pectoral-fin lobe slightly longer than the upper one.
It differs in the lower hump (depth at occiput 106% HL v.
133%), shorter pyloric caeca (3.5-4.5% SL v. 5-9.5%),
undotted gill arches (v. dark-dotted in P. nataliae ) and the
well-developed angulate opercular lobe.
Psednos sp.
Material examined. AMS 124980-003 (dried), unknown
sex and TL, 63 mm SL. FRV Kapala, fid. no. K84-16-15,
33°43.5'S 152° 00.5'E, off Sydney, New South Wales, 960-
998 m, 27 Sep. 1984.
Description. Vert. 57 (11+46), interneural of the 1st dorsal
ray fits between 6th and 7th neural spines, 1 free interneural
present anteriorly between 5th and 6th neural spines. P 16
(9+1+6). Ratios: HL23, its width 11.1 (48), depth at occiput
c. 26.2 (114), E 6.3 (27.6), uj 12.7 (55), gs 5.5 (24.1).
Coronal pore absent. Teeth in 17 and 20 irregular rows,
uniserial posteriorly, about 5 teeth in a row near symphysis.
Opercular lobe distinct. Gill opening less than eye (0.8 eye)
and entirely above pectoral base.
Colour. Body light, peritoneum black. Orobranchial cavity
and tongue light.
Comparative notes. Differs from all other Australian
Psednos in the large eye (6.3 v. 4.1-5.2% SL and 27.6 v.
19-21% HL) and short gill opening (24 v. 31-35% HL)
less than eye (0.8 v. 1.6-1.8 eye) and entirely above the
pectoral base (v. reaching ventrally to 2nd-5th pectoral
rays). Because of its poor condition and our inability to
provide a complete description, we do not name this species.
Genus Paraliparis Collett, 1878
Paraliparis Collett, 1878: 34 (type species Paraliparis bathybii
Collett, 1878 by monotypy). Burke, 1930: 154; Andriashev,
1954: 464; Cohen, 1968: 385; Stein, 1978a: 5, 1978b: 37;
Andriashev, 1986: 14.
Diagnosis. One pair of nostrils (nostrils single). Ventral
sucking disk absent. A single terminal (sensu Andriashev,
1986) or suprabranchial (sensu Burke, 1930) pore present
in temporal canal. Pectoral fin divided into two lobes or
not; if present, lower lobe not forming a single filament.
Pseudobranch absent. Coronal pore absent. Barbels or skin
flaps on head absent.
Key to Australian Paraliparis
1 Chin pore one (Fig. 8-IV). Paraliparis impariporus
-Chin pores two (Fig. 8-1, II, III). 2
2 Mouth oblique, symphysis of upper jaw level with lower margin
of eye (Fig. 5-1). 3
-Mouth horizontal, symphysis of upper jaw below eye level (Fig.
5-II). 5
3 Lower jaw projecting (Fig. 6-V). Body ink-black, skin dense, not
transparent. Secondary caudal rays present, C 1+3/3+1. P. eastmani
-Lower jaw included (Fig. 6-IV). Body nut-brown or reddish-
brown, skin thin and semitransparent. Secondary caudal rays
absent, C 4/4.4
4 Teeth tiny, tooth plates look smooth; diastemae absent. Eye close
to dorsal contour of head. Body uniform nut-brown. Lips,
subrostral fold inside, and chin same colour as head. D 58, first
dorsal-ray interneural between neural spines 5 and 6. Radials 3+0,
fenestrae in cartilaginous basal lamina absent. Pectoral fin short,
60.5% HL. P. avellaneus
-Teeth normal, quite large; diastemae present. Eye far below dorsal
contour of head. Body reddish-brown, caudal darker. Lips and
Stein et al .: Australian liparids
chin blackish, subrostral fold inside brightly black-dotted. D 62,
first dorsal-ray interneural between neural spines 3 and 4. Radials
2+0, fenestrae two. Pectoral fin long, 75% HL.
. P. brunneocaudatus
5 Mouth inferior (Fig. 5-IIC). Gill opening dorsally located, ventral
end level with upper margin of pupil or above (Fig. 26 or 43).
. 6
—— Mouth terminal or subterminal (Fig. 5-IIA,B). Gill opening more
ventrally located, ventral end level with eye centre or below.
. 7
6 Body very dark brownish-black, tooth plates dark grey. Subrostral
fold present, very deep, entirely covering upper lip (Fig. 3-II). Lower
jaw subterminal. Teeth tiny, tooth plates look smooth. Pores distinctly
contoured. Eye 23% HL, gill opening almost equal to eye.
. P. plagiostomus
—— Body colour brown, tooth plates pale. Subrostral fold absent, upper
lip entirely visible (Fig. 3-1). Lower jaw included. Teeth normal,
prominent. Pores not contoured. Eye large, 29% HL. Gill opening
half of eye.
. P. badius
7 Chin pores without common skin fold anteriorly (Lig. 8-1, III, II
obtusirostris only).
. 15
-Chin pores with common skin fold anteriorly (Lig. 8-II). (Teeth
small. Colour light to dark brown or black).
. 8
8 Eye 27% HL. Gill opening reaching to pectoral ray 4. Pectoral
notch rays rudimentary, P 16/17+2r+3 (Pig. 29). Radials 3+1,
lowermost radial half-moon shaped, at posterior margin of
cartilaginous basal lamina (subrostral fold absent. Body jet-
black. Anus below rear of postorbital space).
. P. cor acinus
-Eye 21-25% HL. Gill opening usually entirely above base of
pectoral fin, only in P. anthracinus reaching to second pectoral
ray (Pig. 20). Pectoral notch rays not rudimentary, P 15/18+1/
2+3/4, radials 2-4, all round.
. 9
9 Opercular lobe ear-shaped (Pig. 10-IVB). Snout abruptly angled,
almost absent in lateral view (Pig. 2-IP). Anterior 4 dorsal
rays short. Anus far posterior, below gill opening.
. P. auriculatus
-Opercular flap of other shape (except in P. labiatus). Snout
normal, well developed. Anteriormost dorsal rays usually not
short, if so, not more than 2. Anus below first or second third of
postorbital space (except P. ater, P. labiatus ).
.10
10 Subrostral fold entirely absent. Gill opening reaching ventrally in
front of second pectoral ray. Lower jaw included. Body elliptic,
deep (Pig. 13-11). Opercular flap sharp-tipped, dorsally notched
(Pig. 10-IB).
. P anthracinus
-Subrostral fold present, usually deep (if shallow, lower jaw
sub terminal, nephrohaemal canal present and radials 3+0 or 2+0
P. ater and P. atrolabiatus). Gill opening above or reaching
ventrally in front of uppermost pectoral ray. Body elliptic, low or
ventrally straight. Opercular flap of other shape.
. 11
11 Opercular flap small, triangular, covering lower half of gill opening
(Pig. 10-IIIA). Mouth short, reaching to below anterior margin of
eye. First dorsal-ray interneural between neural spines 3 and 4,
free interneurals absent. Radials 3+0, round. (Fenestrae absent.
Uniformly black, including lips)..
. P. ater
-Opercular flap larger, rounded, covering upper half of gill
opening. Mouth short or longer, reaching to below pupil. First
dorsal-ray interneural posterior to neural spine 4 (4/5 to 6/7), free
interneurals present. Radials 2 or 4.
. 12
12
362 Records of the Australian Museum (2001) Vol. 53
12 Eye not large, 21% HL. Opercular flap ear-shaped (Fig. 10-IVB).
Chin margin right-angled, with sharp firm edge (Fig. 9-II). Radials
3+1, round; fenestrae absent. Snout deep, bluntly rounded (Fig. 2-
IC). (Fip wide). P. labiatus
-Eye larger, 23-24% HE. Opercular flap of other shape. Chin slanted
or with rounded edge (Fig. 9-1 or 9-III). Radials 2 or, if 4, notched,
3 fenestrae present. Snout not blunt, rounded. 13
13 Body light brown, head much paler, yellowish, lips distinctly darker,
blackish. First dorsal ray interneural between neural spines 6 and
7. Anus below first third of postorbital space. Snout shallow,
rounded, projecting (Fig. 2-IE). P. atrolabiatus
Head and body colour uniform dark. First dorsal ray interneural
between neural spines 4 and 5 or 5 and 6. Anus below second third
of postorbital space. Snout deep (Fig. 2-IG or I). 14
14 Body elliptic, low, not humpbacked (Fig. 13-1). Anterior 2 dorsal
rays short. Anus below second third of postorbital space.
Nephrohaemal canal absent. P 18+1+4, radials 4 (3+1), notched, 3
fenestrae present. Chin pore interspace 1 pore diameter, unpigmented.
Eye nearly touching dorsal contour of head. Fips paler than head. P. brunneus
—— Body ventrally straight, humpbacked (Fig. 13-V). Anterior dorsal
rays not short. Anus below middle of postorbital space.
Nephrohaemal canal on vertebrae 5-11. P 15+2+4, radials 2+0,
round, fenestrae absent. Chin pores almost touching, interspace
unpigmented. Eye far below dorsal contour of head. Fips as dark
as head. P. delphis
15 Chin pores not in a common depression (Fig. 8-1). 23
-Chin pores in a common shallow oval depression (Fig. 8-II,
IIIA,B). 16
16 Pectoral notch rays one or two, not rudimentary, about l A-Vi upper
lobe length. 17
-Pectoral notch rays absent (P 16+0+4), or rudimentary (15+lr+4
=3+lr) and visible only in preparations. 18
17 Prominent costal ridge present on each side of body above anterior
half of pectoral fin. Chin pore interspace pigmented. Opercular flap
triangular, with equal sides (Fig. 10-IA), covering gill opening
entirely. 22
-Costal ridge weak or absent. Chin pore interspace unpigmented
with exception of P. gomoni. Opercular flap not as above (Fig. 10-
IB, IIA,B,C) (Parapophyses of posterior abdominal vertebrae not
joined). 19
18 Chin pore pit on lower surface of chin (Fig. 9-III). Subrostral fold
present. Eye close to dorsal contour of head. Fips as dark as head.
Nephrohaemal canal absent. P. 16+0+4. P. australiensis
—— Chin pore pit on anterior surface of chin (Fig. 9-IVA). Subrostral
fold absent. Eye well below dorsal contour of head. Fips paler than
head. Nephrohaemal canal present. P. 15+lr+4. P. csiroi
19 C 9 (1+4/4). Fower jaw subterminal. Radials 2+0. Body ventrally
straight, leaf-like (Fig. 13-VII). (Mouth reaching to below anterior
half of pupil. P 14+2/3+3/4. Anterior two dorsal rays short. Snout
deep, blunt. Mouth terminal). P. obtusirostris
C 8 (4/4). Fower jaw included or almost included. Radials 3+0.
Body of other shape (Fig. 13-1, III). 20
Stein et al .: Australian liparids 363
20 Mouth terminal. Chin margin vertical, right-angled (Fig. 9-II),
honeycomb tissue on chin absent. Snout deep, blunt (Fig. 2-IA).
Chin pore interspace pigmented. Opercular flap rounded ventrally
(Fig. 10-IIB). Radials unnotched, round, fenestrae absent. P. gomoni
-Mouth subterminal. Chin margin slanted or rounded, honeycomb
tissue on chin present. Snout rounded (Fig. 2-IB,E). Chin pore
interspace unpigmented. Opercular flap sharp-tipped (Fig. 10-IB,
IIC). Radials notched, fenestrae present. 21
21 Upper pectoral-fin lobe rays 17. Body hump-backed. Mouth cleft
reaching to below anterior margin of eye. Subrostral fold present,
anus below first third of postorbital distance. P. infeliciter
-Upper pectoral-fin lobe rays 14/15. Body straight-backed, ventral
contour much more curved than dorsal. Mouth cleft reaching to
below anterior margin of pupil. Subrostral fold absent, anus almost
below gill opening. Paraliparis sp. 1
22 Chin pore interspace equals 2 pore diameters. Oral cleft reaching
to below pupil. Anterior 3 dorsal rays not short. Head 21.4% SL.
Pyloric caeca grey. Radials 4 (3+1). Subrostral fold present. Lower
jaw included. Body ventrally straight, hump-backed. P dewitti
-Chin pore interspace equals 1 pore diameter. Oral cleft reaching
to below anterior margin of eye. Anterior 3 dorsal rays short. Head
small, 17.7% SL. Pyloric caeca pale. Radials 2+0. Subrostral fold
absent. Lower jaw subterminal. Body elliptic, low. Paraliparis sp. 2
23 Subrostral fold absent. Radials 2+0. Opercular flap covers lower
2 A of gill opening, rounded (Lig. 10-IIA). Preoperculo-mandibular
sensory pore series unusually close to infraorbital series, pm 7 close
to eye (distance c. Vi postorbital space). (Colour brown, with
whitish blotches but without reddish tint). P hobarti
-Subrostral fold present. Radials 3 or 4. Opercular flap of other
shape (Lig. 10-IIC, IIIA,B, IVA). Preoperculo-mandibular sensory
pore series not close to infraorbital series, pore pm 7 normally
located (distance c. Vi postorbital space). 24
24 Body reddish-brown. Teeth large, stout. Opercular flap small,
triangular, covering lower half of gill opening (Pig. 10-IIIA).
(Radials 3+0. Lower jaw included). P. lasti
—— Body black or dark blackish-brown. Teeth tiny or large (P.
retrodorsalis ). Opercular flap not as above. (Radials 3 or 4) . 25
25 Pectoral notch ray single, rudimentary, two or more pectoral girdle
fenestrae present (Pig. 45). P. tasmaniensis
-Pectoral notch rays two or more, not rudimentary, fenestrae
absent. 26
26 Lower jaw included. Tooth plates dark. Mouth cleft reaching
to below eye centre. Snout acutely angled. Opercular flap as in
Pig. 10-IIC. P. retrodorsalis
-Lower jaw subterminal. Tooth plates pale. Mouth cleft reaching
to below anterior margin of eye. Snout deep, blunt, vertical.
Opercular flap as in Pig. 10-IIIB or IVA. 27
27 D 62-66. Costal keels distinct. Procurrent caudal rays present, C
1+3/3+1. Radials 3+1. Opercular flap covers lower half of gill
opening (Pig. 10-IIIB). Chin slanted. P. costatus
-D 59. Costal keels absent. Procurrent caudal rays absent, C 4/4.
Radials 3+0. Opercular flap dorsally rounded (Pig. 10-IVA). Chin
right-angled (Pig. 9-II). P piceus
364 Records of the Australian Museum (2001) Vol. 53
Figure 20. Paraliparis anthracinus n.sp. A, holotype, CSIRO H1576-02, ripe !, 150 mm TL, 133 mm SL. B,
ventral view of mouth and upper jaw tooth plate. C, P 793, cleared and stained right pectoral girdle.
Paraliparis anthracinus n.sp.
Fig. 20
Material examined. Holotype CSIRO HI576-02, ripe $,
150 mm TL, 133 mm SL. FRV Soela, stn. So 3/86/27,
41°46.4'S 144°24.4'E, W coast of Tasmania, W of Granville
Harbour, 1024-1080 m, 14 May 1986; radiograph 680 D;
pectoral girdle 793.
Diagnosis. Vert. 68, D 60, C 8, radials 4, rounded. Snout
short, 27.8% HL. Mouth horizontal, subterminal, lower jaw
included. Subrostral fold absent. Teeth simple, small. Chin
pores with common skin fold anteriorly, located in a small
depression undefined posteriorly. Ventral end of gill opening
reaching in front of 2nd pectoral ray, dorsal pectoral ray
level with eye centre. P 16+2+4, rudimentary notch rays
absent. Distance from chin to anus long, 15% SL. Anus
below second third of postorbital space. Head 20.3% SL,
preanal distance large, 38%. Body black, peritoneum black.
Further description. Counts: D 60, A 55, P 22, C 8 (4/4),
Vert. 68 (11+57), radials 4 (3+1), fenestrae absent, gr 8, pc
6, pores 2-6-7-1. Ratios: HL 20.3, its width 12.9 (59), and
depth 17.3 (85), bd 21.8 (107), bdA 18.8 (93), preD 25.6,
preA 38.3, ma 15.0, aAf 22.6, UPL 13.2 (65), LPL —, NL
—, E c. 5.1 (25.0), gs 4.1 (20.0), sn 5.6 (27.8), po 10.5
(52), io 8.3 (40.7), so 3.8 (18.5), uj 8.3 (41.4), lj 8.6 (42.5),
pc 6.2.
Head not small, 4.9 in SL, dorsal profile evenly sloping
anteriorly to deep snout. Head depth 1.4 its width. Eye quite
large, 4 in head, not touching dorsal contour of head;
suborbital distance long, 0.7 eye; pupil % eye. Interorbital
space 1.6 eye. Snout short, deep, bluntly rounded, 1.1 eye,
slightly projecting anterior to upper jaw, its highest point
level with upper margin of pupil. Subrostral fold absent.
Nostril level with mid eye. Mouth horizontal, subterminal,
cleft reaching to below anterior fourth of eye; lower jaw
included. Teeth simple, small, slightly prominent, 26 rows
in upper jaw, anteriorly 11-12 teeth in length. Lower tooth
plates damaged. Diastema of upper jaw narrow, absent in
lower jaw. Chin right angled, lower jaw below posterior of
oral cleft deep. Circumoral pores with thickened rims; chin
pore interspace equal to pore diameter, pigmented, common
anterior skin fold present with pores in a shallow depression
undefined posteriorly. Gill opening short, 0.8 eye, ventral
end level with eye centre, reaching in front of 2nd pectoral
ray. Opercular flap triangular, dorsally notched, sharp tip
level with upper margin of eye.
Uppermost pectoral ray level with mid-pupil. Pectoral
rays broken, fin membrane missing. P 16+2+4, rudimentary
rays absent. Radials 3+1, round, of similar sizes. Fenestrae
absent. Coracoid helve thin, comparatively short.
Body deep, deepest between dorsal and anal-fin origins,
4.6 in SL; dorsal contour broadly rounded anteriorly and
posteriorly from this point. General body shape elliptic,
dorsal and ventral contours similar. Horizontal midline
touching lower margin of eye. Preanal length large, 38%
SL. First dorsal ray interneural between neural spines 5 and
6, two free anterior interneurals present. Anterior dorsal rays
embedded in gelatinous tissue. Parapophyses of vertebra
11 joined together, forming short haemal spine. Costal ridges
absent. Epineural ribs on vertebrae 2-13, epipleural ribs on
3-23, anterior ribs of both series not long, shorter than 3
vertebrae. Anus below second third of postorbital space.
Vertical fins overlap about half of caudal fin. Skin opaque,
prickles absent. Gelatinous tissue moderately developed.
Stein et al .: Australian liparids 365
Genital papilla absent. Ovarian eggs up to 3.5 mm. Pyloric
caeca sharply pointed.
Colour. Head and body, lips, subrostral fold and chin
uniformly black. Mouth grey, tongue slightly lighter, grey-
dotted, tooth plates pale. Branchial cavity black, gill arches
dark grey. Peritoneum black, pyloric caeca and stomach
pale.
Distribution. West coast of Tasmania, 1024-1080 m.
Etymology. Anthracinus from the Latin, coal-coloured.
Comparative notes. A member of group IIIc, P. anthracinus
is distinguished by absence of a subrostral fold, gill opening
reaching ventrally to the 2nd pectoral ray, included lower
jaw, right-angled chin, short snout, 3+1 round radials,
elliptical body, uniform black colour, and normally
developed anterior dorsal-fin rays. Most similar to P.
labiatus but differs in the absence of the subrostral fold (v.
present and deep anteriorly), in having lip posteriorly
narrowed (v. distinctly wide), mouth subterminal (v.
terminal), lower jaw deep below posterior end of oral cleft
(v. shallow), opercular flap sharp-tipped and dorsally
notched (v. ear-shaped).
Paraliparis ater n.sp.
Fig. 21
Material examined. Holotype CSIRO H749-04, ripe 9 ,
140 mm TL, 124 mm SL. FRV Soela, stn. So 3/86/32,
41°45.8'S 144°24.8'E, W coast of Tasmania, W of Granville
Harbour, 16 May 1986, 1000-992 m; radiograph 680 F 2;
pectoral girdle 794.
Diagnosis. Vert. 67, D 62, C 8, radials 3. First dorsal
interneural between neural spines 3 and 4. Mouth horizontal,
subterminal, small; teeth simple, small. Lower jaw
subterminal. Interorbital wide, 2.1 eye. Chin pores with
anterior common skin fold, interspace less than diameter.
Ventral end of gill opening and dorsal pectoral ray horizontal
with lower half of eye. P 16+1+3, rudimentary notch rays
absent. Head 18.3% SL, ma large, 14.6%. Nephrohaemal
canal present. Colour black, peritoneum black.
Further description. Counts: D 62, A 53, P 20, C 8 (4/4),
Vert. 67 (11+56), radials 3+0, gr 7, pc 6, pores 2-6-7-1.
Ratios: HL 18.3, its width 10.9 (60), and depth 15.4 (84),
bd 17.6 (96), bdA 16.2 (89), preD 20.6, preA 36.7, ma 14.6,
aAf 20.0, UPL 12.7 (70), LPL 10.7 (59% HL, 84% UPL),
NL —, E 4.0 (22.2), gs 3.8 (20.9), sn 5.7 (31.3), po 8.9
(49), io 8.6 (47.0), so 3.0 (16.5), uj 8.1 (44), lj 7.8 (42.6),
pc 4.7.
Head small, 5.5 in SL, evenly sloping anteriorly to
rounded snout, depth 1.4 width. Eye small, not touching
dorsal contour of head, suborbital space about Vs eye
diameter. Pupil about Vs eye. Interorbital space wide, 2.1
eye. Snout 1.4 eye, bluntly rounded, slightly projecting
beyond upper jaw, its highest point level with upper margin
of pupil. Subrostral fold present, shallow and almost absent,
not covering upper lip. Nostril pore-like, horizontal with
mid-pupil. Mouth subterminal, horizontal and small, oral
cleft reaching only to below anterior margin of eye. Lower
jaw subterminal, teeth simple, very small, only slightly
prominent, in 21-23 rows of up to 8-9 teeth anteriorly.
Diastemae narrow, almost absent. Lower jaw tapering
anteriorly, chin rounded in ventral view. Circumoral pore
rims slightly thickened. Chin pores slit-like, interspace
pigmented, less than their diameter, with thin co mm on skin
fold anteriorly. Chin symphysis slanted posteroventrally in
lateral view, not right-angled. Gill opening small, 0.9 eye,
Figure 21. Paraliparis ater n.sp. A, holotype, CSIRO H749-04, ripe $, 140 mm TL, 124 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 794, cleared and stained right pectoral girdle.
366 Records of the Australian Museum (2001) Vol. 53
its lower end level with eye centre. Opercular flap small,
triangular, tip rounded, horizontal with upper margin of
pupil, covering lower half of gill opening.
Uppermost pectoral ray level with lower margin of pupil,
ventralmost ray below anterior third of postorbital space.
Pectoral upper lobe not reaching to anal-fin origin, lower
lobe c. 84% UPL. P 16+1+3, rudimentary rays absent. Skin
of pectoral fin damaged, notch membrane apparently
attached to body almost at full ray length. Radials 3+0,
round, fenestrae absent. Coracoid with thin, comparatively
short helve.
Body elliptic, not deep, 5.7 in SL (96% HL), deepest
between dorsal and anal-fin origins; dorsal contour broadly
rounded anteriorly and posteriorly from this point, dorsal
and ventral contours similar. Horizontal midline touching
lower margin of pupil. Preanal distance long, 37% SL.
Interneural of first dorsal ray between neural spines 3 and
4, free interneural absent. Anterior dorsal rays embedded in
gelatinous tissue. Parapophyses of vertebrae 5-11 obviously
joined, forming nephrohaemal canal. Epineural ribs on
vertebrae 2-9, up to 3 vertebrae in length; epipleural ribs on
vertebrae 4—16, slim, not clearly visible on radiograph, up to
1.5 vertebrae in length; costal keel detectable, but weak. Anus
below posterior third of postorbital space. Vertical fins
overlapping caudal fin slightly more than half. Skin thick,
opaque, prickles absent. Gelatinous tissue moderately well
developed. Genital papilla absent. Individual mature, a few
ripe eggs in oviduct up to 2.4 mm, ovarian eggs much
smaller. Sexually mature at a short length, about 126 mm
SL. Pyloric caeca, thick, with sharp tips.
Colour. Body and head uniformly black; lips and inner
surface of subrostral fold almost as black as head; chin
slightly paler, dark grey. Pores pale inside. Mouth grey, tooth
plates dark, tongue slightly lighter, grey-dotted. Branchial
cavity brownish-black, gill arches grey. Peritoneum black,
pyloric caeca and stomach pale.
Distribution. West coast of Tasmania, 1000-992 m.
Etymology. Ater in Latin, meaning black as soot.
Comparative notes. Paraliparis ater belongs to group IIIc.
It is distinguished by a nephrohaemal canal, anterior dorsal-
fin origin (between neural spines 3 and 4, free interneurals
absent), wide interorbital 47% HL, short oral cleft
(extending posteriorly to below the anterior margin of the
eye), pectoral girdle with radials 3+0, round, fenestrae
absent; by its uniformly dark black colour (lips as dark as
the head), and in the shape of the opercular flap.
Paraliparis atrolabiatus n.sp.
Fig. 22
Material examined. Holotype CSIRO H550-11, mature
6, 127 mm TL, 114 mm SL. FRV Soela, stn. So 3/86/36,
42°20.45'S 144°40.4'E, W coast of Tasmania, W of Cape
Sorell, 1120-1220 m, 17 May 1986; radiograph 680 G;
pectoral girdle 788.
Diagnosis. Vert. 68, D 60, C 8, radials 2. Mouth subterminal,
teeth small. Chin pore interspace equals twice their diameter;
common skin fold present anteriorly. Gill opening short,
lower end horizontal with eye centre. Pectoral fin 16+2+3,
short, only posterior part of upper and lower lobes
protruding from skin; notch shallow. Anus below first third
of postorbital space. HE 19.3% SL, aAf short, 18.8% SL,
preA 32% SL. Nephrohaemal canal present. Body and head
light brown, peritoneum black, mouth dark grey, tongue
black-dotted.
Further description. Counts: D 60, A 55, P 21, C 8 (4/4),
Vert. 68 (11+57), radials 2 (2+0+0), gr 7, pc 4, pores 2-6-7-
1. Ratios: HL 19.3, its width 13.0 (67), and depth 15.8 (82),
bd 18.4 (95), bdA 17.5 (91), preD 21.9, preA 31.6, ma 12.2,
aAf 18.8, UPL 11.8 (61), LPL 7.2 (37% HL, 61% UPL),
NL 1.9 (16.4% UPL), sn 7.1 (36.8), E 4.5 (23.2), gs 3.1
(15.9), po 9.6 (50), io 7.9 (41), so 3.9 (20.5), uj 8.8 (45.5),
lj 8.9 (46), pc 6.1.
Head short, 5.2 in SL, and wide, its width 67% HL, its
depth 1.2 its width. Dorsal contour of head deep at occiput,
gradually sloping in anterior profile. Snout projecting,
extending above upper jaw, its most anterior point below
horizontal through eye centre; snout length 1.6 eye diameter.
Subrostral fold present, not deep, covering upper lip about
half. Eye not large, 4.3 in HL, its contour not touching dorsal
profile of head, suborbital space almost equal to eye. Pupil
c. half eye diameter. Interorbital space 1.8 eye diameter.
Nostril small, pore-like, horizontal with lower half of pupil.
Mouth subterminal, its cleft reaching to below anterior third
of eye. Lower jaw subterminal (shorter than upper, but not
included). Depth of lower jaw below end of mouth cleft
quite deep. Teeth small, slightly prominent, in about 20 rows
of up to 8-9 teeth anteriorly. Diastemae absent from both
jaws. Circumoral pores small, not contoured (rims not
thickened), at surface, not in pits. Chin pore interspace
pigmented, twice their diameter, chin pores small,
longitudinally oval, anterior shallow skin fold present. Gill
opening short, 0.7 eye diameter, vertical, its upper end above
eye level, lower end on horizontal through pupil. Opercular
flap small, rounded, covering 3 A of gill opening, its tip level
with upper margin of eye.
Dorsalmost pectoral ray on horizontal through upper half
of pupil, base of ventralmost on vertical through middle of
postorbital space. P 16+2+3, notch rays short (16% UPL),
rudimentary rays absent. Pectoral lobes not reaching anal-
fin origin; lower lobe short, only 37% SL, 7.2% SL. Skin
on body covering proximal half of upper lobe, only its
posterior half free and movable; lower fin lobe about X A
covered; notch rays covered by skin entirely; free portions
of upper and lower lobes appear separate and unjoined by
fin membrane. Radials 2 (2+0+0), fenestrae between radials
absent. Scapula with a strong shaft. Coracoid with a very
long thin helve.
Body not deep, greatest depth at anterior dorsal-fin ray,
about 95% HL. Not hump-backed, dorsal profile gradually
rounded; ventral contour straight. Preanal distance short,
31.6% SL. Haemal spine of last abdominal vertebra (12)
equals about half next (13). Vertebrae 35 and 36 damaged,
healed. First dorsal rays not rudimentary, gradually
elongated; first ray-bearing interneural between neural
spines 6 and 7; two free anterior interneurals present
between 4th, 5th, and 6th neural spines. Costal ridges weakly
developed. Epineural and epipleural ribs slender, thin, short,
on vertebrae 3-19, 2 vertebrae or less in length. Nephro¬
haemal canal present: parapophyses of vertebrae 3-11
joined to form very short haemal spines, directed anteriorly
at vertebrae 5-7. C 4/4. Vertical fins overlap caudal about
Stein et al .: Australian liparids 367
Figure 22. Paraliparis atrolabiatus n.sp. A, holotype, CSIRO H550-11, mature ?, 127 mm TL, 114 mm SL. B,
ventral view of mouth and upper jaw tooth plate. C, P 788, cleared and stained right pectoral girdle.
half. Anus below Vs of postorbital space, aAf short. Skin
unprickled, thin, semitransparent. Subcutaneous gelatinous
tissue well developed. Pyloric caeca of similar lengths.
Specimen an adult male with ripe testes and prominent cone¬
like urogenital papilla below middle of postorbital space. Males
mature at small length, about 114 mm SL.
Colour. Skin light brown with broad markings; head much
lighter than body, yellowish. Pectoral fin, margin of
opercular flap and urogenital papilla blackish. Inner surface
of subrostral fold and lips distinctly darker than head, dark
blackish-brown. Mouth dark grey, darker than head, tongue
black-dotted. Tooth plates pale, gill arches light grey-dotted,
branchial cavity dark grey, almost black. Peritoneum black,
stomach and pyloric caeca pale.
Distribution. West coast of Tasmania, 1120-1220 m.
Etymology. The name derives from Latin atro —dark and
labium —lip.
Comparative notes. Paraliparis atrolabiatus is in group
IIIc; it is distinguished by its broad head, peculiarly skin-
bound pectoral fins with short lower lobes, short gill
opening, short distance between anus and anal fin, light-
brown body and yellowish head, two pectoral radials, and
the nephrohaemal canal. Externally it is most similar to P.
auriculatus, which has a similar pectoral fin and chin pores,
but it differs from the latter in presence of a nephrohaemal
canal (v. absence), two radials (v. 3), interradial fenestrae
absent (v. two small present), snout normal (v. strongly
angled, almost absent), light brown head (v. dark brown),
lips blackish-brown, darker than head (v. grey, lighter than
head), mouth dark grey and darker than head (v. grey, lighter
than head), anterior dorsal rays gradually elongated (v. 4
anterior rays distinctly shorter than following), and anus below
anterior third of postorbital space (v. below posterior third).
Paraliparis auriculatus n.sp.
Fig. 23
Material examined. Holotype CSIRO H749-06, $, 145
mmTL, 131 mm SL. FR VSoela, stn. So 3/86/32,41°45.8’S
144°24.8'E, W coast of Tasmania, W of Granville Harbour,
1000-992 m, 16 May 1986; radiograph 680 F4; pectoral
girdle 740.
Diagnosis. Vert. 68, D 62 (4+58), anterior 4 rays short and
thin, C 8, radials 3, notched. Mouth sub terminal, teeth tiny,
tooth plates smooth. Snout not protruding, angled
posterodorsally from immediately above upper lip. Eye
25.4%, io 48% HL. Chin pore interspace equal to their
diameter, with a common shallow skin fold anteriorly. Gill
opening small, 0.6 eye. Pectoral fin 16+2+3, with short
lobes, posterior half only of upper and lower lobes
protruding from skin. Anus slightly anterior to gill opening.
HL 18.6%, preA distance short, 29%, aAf 16.7% SL. Skin
dark brown, peritoneum black, mouth grey, tongue black-
dotted.
Further description. Counts: D c. 62 (4+58), A c. 55, P
21, C 8 (4/4), Vert. 68 (12+56), radials 3 (3+0), gr 9, pc 4,
pores 2-6-7-1. Ratios: HL 18.6, its width 11.1 (60), and
depth 15.3 (82), bd 16.8 (90), bdA 16.8 (90), preD 20.6,
preA 29.0, ma 13.7, aAf 16.7, UPL 11.7 (63), LPL9.9 (53%
HL, 84% UPL), NL 1.6 (13.6% UPL), sn 6.5 (34.8), E 4.7
(25.4), gs 3.1 (16.4), po 9.1 (49), io 9.0 (48.3), so 2.4 (13.1),
uj 8.0 (43), lj 7.3 (39), pc 5.
Head short, 5.4 in SL, not much compressed, its width
60% HL, its depth 1.4 its width. Dorsal contour of head
deep at occiput, widely rounded in anterior profile. Snout
deep, strongly angled, almost absent in lateral view, its
highest point horizontal with upper margin of eye, tip just
above upper lip; length from maxillary symphysis to anterior
368 Records of the Australian Museum (2001) Vol. 53
B
Figure 23. Paraliparis auriculatus n.sp. A, holotype, CSIRO H749-06, 9, 145 mm TL, 131 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 740, cleared and stained right pectoral girdle.
margin of eye 1.4 eye diameter. Subrostral fold present,
deep but not completely covering upper lip. Eye quite large,
3.9 in HL, its contour not touching dorsal profile of head,
suborbital short, half of eye. Pupil c. % eye diameter.
Interorbital space rounded, broad, 1.8 eye diameter. Nostril
small, pore-like, horizontal with lower margin of pupil.
Mouth subterminal, its cleft reaching to below anterior
margin of pupil. Lower jaw subterminal, almost included,
deep below posterior of oral cleft. Teeth tiny, tooth plates
appear smooth, in 23-25 rows of up to 8-9 teeth anteriorly.
Diastemae absent. Circumoral pores small, chin pores
interspaced at their diameter, small, longitudinally oval,
interspace pigmented, anterior shallow skin fold present.
Gill opening short, 0.6 eye diameter, its upper end slightly
above eye level, lower end on horizontal through eye centre;
dorsal end anterior to the lower. Opercular flap small, ear¬
shaped, its tip above level of upper margin of eye.
Dorsalmost pectoral ray on horizontal through centre of
eye, base of ventralmost on vertical through middle of
postorbital space. P 16+2+3, notch rays short (14% UPL),
rudimentary rays absent. Pectoral lobes both short, not
reaching anal-fin origin. Skin closely attached to body
covering anterior half of upper lobe, leaving only posterior
half of fin free and movable; lower fin lobe similar; notch rays
covered by skin entirely; free parts of upper and lower lobes
appear to be separate, not joined by fin membrane. Radials 3
(3+0), two small fenestrae between radials present. Scapula
with a long shaft, on which a dorsal lateral rib is present.
Coracoid with a small slit-like opening and a strong helve.
Body not deep, greatest depth at anterior ray of dorsal
fin, about 90% HL. Not hump-backed, dorsal profile
gradually rounded; ventral contour almost straight.
Horizontal midline of body passes through upper margin
of pupil. Preanal distance short, 29% SL. Haemal spine of
last abdominal vertebra (12) about half the length of the
next. Lirst dorsal ray rudimentary, its interneural between
neural spines 5 and 6; free anterior interneurals present
between 3rd, 4th, and 5th neural spines. Next three dorsal
rays short and thin. Epineural ribs on vertebrae 2-14,
epipleural ribs on 3-20, well developed but thin, about 2-
2.5 vertebrae in length. Costal ridges weakly developed. C
4/4. Vertical fins overlap caudal by about half. Anus slightly
anterior to gill opening. Skin quite thin, semitransparent.
Subcutaneous gelatinous tissue well developed. Pyloric
caeca of similar lengths. Specimen an adult female, one
large ripe egg 3 mm in diameter present near oviduct
opening, ovarian eggs much smaller and unripe.
Colour. Head and body dark, blackish-brown with irregular
broad darker areas. Inner surface of subrostral fold, lips
and mouth distinctly lighter, grey; tongue densely black-
dotted. Tooth plates pale, gill arches light grey-dotted,
branchial cavity dark brown. Peritoneum black, stomach
and pyloric caeca pale.
Distribution. West coast of Tasmania, 1000-992 m.
Etymology. The specific epithet auriculatus from the Latin
for small ear, auricula refers to the shape of the opercular flap.
Comparative notes. Paraliparis auriculatus is in group
IIIc; it is distinguished by its peculiar skin-bound pectoral
fin with short upper and lower lobes, small gill opening,
ear-shaped opercular flap, snout not protruding, compara¬
tively large eye and broad interorbital, more posterior
position of the anus, the shortened anterior four dorsal rays,
and tiny teeth. Among species of this group which have no
nephrohaemal canal, it is the only one with 3+0 radials (v.
4). It is most similar to P. retrodorsalis in the posterior
position of the dorsal origin, but differs from it in its short
preanal distance of 29% SL (v. 37%), anus-anal-fin distance
Stein et al .: Australian liparids 369
Figure 24. Paraliparis australiensis n.sp. A, holotype, NMV A21497, 2,176 mm TL, 164 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 789, cleared and stained right pectoral girdle.
of about 17% SL (v. 26%), insertion of the first dorsal-fin
interneural between neural spines 5 and 6 (v. between 7
and 8), two interneurals (v. 3), radials 3 (3+0) v. 4 (3+1),
two small pectoral girdle fenestrae (v. none), diastemae
absent (v. distinctly developed), tiny teeth (v. normal), dark
brown colour (v. black), pale tooth plates (v. dark), and other
characters.
Paraliparis australiensis n.sp.
Fig. 24
Material examined. Holotype NMV A21497, 2 ,176 mm
TL, 164 mm SL. FRV Soela, stn. So 1/88/09, 37°01.09’S
137°25.44'E, 100 km S of Kangaroo Island, South Australia,
1090-1160 m, 24 Jan. 1988; radiograph NMV 5873E;
pectoral girdle 789.
Diagnosis. Vert. 65, D 60, C 8, radials 3, round. Eye nearly
touching dorsal contour. Snout blunt, large, 37.5% HL.
Mouth terminal, horizontal, teeth very small. Chin pore pair
in a common pit, interspace equals pore diameter. Gill
opening ventral end horizontal with lower third of eye. P
16+0+4, upper lobe short, 63% HL, notch rays absent. Body
deep, 125% HL. HL 19.5% SL, preA 34.5% SL, aAf long,
30% SL. Colour very dark blackish-brown, peritoneum
black, mouth and tongue grey.
Further description. Counts: D 60, A 54, P 20, C 8 (4/4),
Vert. 65 (11+54), radials 3 (3+0), round, gr 9, pc 5, pores 2-
6-7-1. Ratios: HL 19.5, its width 12.8 (66) and depth 16.3
(84), bd 24.4 (125), bdA 20.4 (105), preD 24.4, preA 34.5,
ma 12.5, aAf 30.0, UPL 12.2 (63), LPL 10.5 (59% HL,
95% UPL), notch rays absent, E 4.6 (23.8), gs 3.8 (19.3),
sn 7.3 (37.5), po (51.6), io 7.5 (38.4), so 2.7 (14.0), uj 8.5
(44), lj 7.9 (40.6), pc 9.8.
Head small, 5.1 in SL, quite deep at occiput, dorsal
contour slopes steeply anteriorly and much more gently
posteriorly. Head not much compressed, depth 1.3 its width.
Snout large, deep, and blunt, its highest point horizontal
with upper margin of pupil; in lateral view, snout length
about equal to eye; length from symphysis of upper jaw to
anterior margin of eye is 1.6 eye diameter. Subrostral fold
deep, covering upper lip almost entirely. Eye almost
touching upper contour of head, suborbital about 0.6 eye
diameter, pupil about half eye diameter. Interorbital space
1.6 larger than eye. Nostril level with eye centre, with raised
rim, twice diameter of snout pores. Mouth terminal,
horizontal, its cleft not quite reaching to below anterior
margin of pupil. Lower jaw subterminal, almost included,
chin widely rounded, quite gelatinous, shallow below
posterior end of oral cleft. Teeth very small, slightly
prominent, in 25-26 quite regular rows, curving anteriorly
on tooth plates, up to 10-12 teeth per row anteriorly.
Diastema of upper jaw narrow, absent in lower jaw.
Circumoral pores small, with slightly raised rims; chin pores
in a shallow small oval pit, interspace pigmented, equal to
pore diameter. Chin pore diameter half that of pm 2 . Gill
opening 0.9 eye diameter, its upper end slightly above level
of upper margin of eye, lower end horizontal with lower
third of eye. Opercular flap ear-shaped, upper margin
notched, tip level with upper margin of pupil.
Upper pectoral ray horizontal with lower margin of eye,
lowermost ray about below posterior margin of eye. Pectoral
fin 16+0+4, deeply notched, lowest upper lobe ray slightly
farther from remainder of rays; upper and lower lobes
connected by fin membrane, normal notch rays absent.
Upper lobe short, not reaching anal-fin origin. Skin of
proximal surface of upper lobe attached to body at ray bases,
in notch at about Vi ray length, and in lower lobe at about Vs
ray length. Pectoral girdle with 3+0 round radials, fenestrae
370 Records of the Australian Museum (2001) Vol. 53
in cartilaginous basal lamina absent. Coracoid with a very
long thin helve.
Body deep, elliptic, 125% HL, greatest depth at dorsal-
fin origin, depth at A origin about equal to eye (105%).
Horizontal midline touches lower margin of pupil. Anterior
dorsal rays embedded in gelatinous tissue; first dorsal ray
short, its interneural between vertebral spines 4 and 5;
anterior free interneurals absent. Parapophyses at least of
abdominal vertebrae 10 and 11 joined, length of haemal
spine of vertebra 11 almost equal to next, not reaching first
interhaemal. Costal ridges weakly developed. Epineural ribs
present on vertebra 2-17, epipleural ribs on vertebrae 4-
17, both thin, not stout, lengths up to 1.5 vertebrae. Anus
just behind a vertical through eye; distance between anus
and anal fin noticeably greater than HL. Skin quite thin,
prickles absent. Vertical fins overlapping anterior third of
caudal. Pyloric caeca similar, quite thick at their bases and
long. Ripe eggs in oviduct c. 2.7 mm in diameter.
Colour. Head and body very dark, uniformly blackish-
brown, chin and lips dark grey, inner side of subrostral fold
black. Mouth and tongue uniformly light grey, tooth plates
dark, branchial cavity black, gill rakers grey. Pores as dark
as head, not whitish. Peritoneum black, stomach and pyloric
caeca pale.
Distribution. Off South Australia at 1090-1160 m.
Etymology. The specific epithet— australiensis —refers to
the country of origin of this new species.
Comparative notes. Paraliparis australiensis belongs to
group Illb, but is distinguished by its large blunt snout, short
upper pectoral lobe, great distance between anus and anal
fin, deep body, small teeth, absence of notch rays in the
pectoral fin, 3 round radials, by a deep subrostral fold, and
by lip and head of similar colour. In this group, only P.
csiroi lacks normal rays in the pectoral-fin notch, but P
australiensis clearly differs from it in having the
parapophyses of only abdominal vertebrae 10 and 11 joined
(v. vertebrae 5-11 joined in very short spines creating a
nephrohaemal canal), eye touching the dorsal contour of
the head (v. not touching), a well-developed subrostral fold
almost covering the upper lip (v. absent), wider head 66 (v.
54)% HL, chin pores in a co mm on depression on lower
surface of chin (v. in a common depression on the anterior
surface of chin), diastema of upper jaw narrow, of lower
jaw absent (v. diastema of lower jaw slightly wider than
that of upper jaw), pyloric caeca lengths 9.8 (v. 4.6)% SL,
darker blackish-brown colour (v. dark brown), fenestrae in
pectoral girdle absent (v. one present), and lip colour similar
to that of head (v. lighter).
Paraliparis avellaneus n.sp.
Fig. 25
Material examined. Holotype NMV A5873, 8, 149 mm
TL, 132 mm SL. LRV Soela, stn. So 1/88/09, 37°01.09’S
137°25.44'E, 100 km S of Kangaroo Island, South Australia,
1090-1160 m, 24 Jan. 1988; radiograph NMV 5873E;
pectoral girdle 735.
Diagnosis. Vert. 65, D 58,7 anterior rays short; C 8, radials
3, round. Head and body low, mouth oblique. Teeth tiny.
Chin pores interspaced by their diameter, not in a pit. Gill
opening short, lower end slightly below level of eye.
Pectoral fin 21, upper lobe short, about 60% HL;
rudimentary rays absent. Head 19% SL, preA 35%; bd 90%
HL. Colour nut-brown, skin thin. Peritoneum black.
Further description. Counts: D 58, A 54, P 21, C 8 (4/4),
Vert. 65 (10+55), radials 3 (3+0), pc 5, gr 8, pores 2-6-7-1.
Ratios: HL 18.8, its width 10.6 (56) and depth 12.9 (69), bd
16.7 (89), bdA 14.5 (72), preD 25.8, preA 35, ma 13.6, aAf
22.7, UPL 11.4 (60.5), LPL 10.6 (93% UPL), NLc. 3.4 (30%
UPL), E 4.5 (24.2), gs 3.9 (20.6), sn 6.1 (32.3), po 9.1 (48), io
6.4 (34.3), uj 8.4 (45.2), lj 8.2 (43.5), so 2.9 (15.3), pc 4.5.
Head small, 5.3 in SL, quite compressed, its width 56%
its length, and low, its depth 69% HL or 1.2 its width, dorsal
contour almost horizontal, slightly declining anteriorly.
Snout deep, blunt, 1.3 eye, not projecting anterior to upper
jaw, not very gelatinous; its highest point level with upper
margin of eye. Subrostral fold deep, entirely covering upper
lip. Eye 4.1 in HL, almost entering upper profile of head;
pupil about half eye diameter, suborbital distance 0.6 eye.
Interorbital space narrow, a little more than Vs HL, 1.4 eye.
Nostril small, on horizontal through upper margin of pupil.
Mouth terminal, oblique; symphysis of upper jaw almost
horizontal with lower margin of eye. Oral cleft reaching to
vertical through anterior margin of eye. Lower jaw included
by upper; moving upper lip allows margins of upper tooth
plates to be seen. Lower jaw tapered anteriorly; chin
rounded, in lateral view slanted, not gelatinous, not deep
below end of oral cleft. Teeth extremely small, barely
projecting above gums; surface of tooth plates appears
smooth. Teeth in about 24-26 oblique rows of 7-8 teeth
each anteriorly. Diastemae absent from both jaws.
Circumoral pores small, not in pits; chin pores separated
by a distance equal to their diameter, anterior skin fold
absent, clearly not in a pit. Gill opening short, its length 0.9
eye diameter, its upper end level with upper half of pupil, its
lower end slightly below horizontal through lower margin of
eye. Opercular flap small, triangular with rounded tip, covering
lower Vs of gill opening, tip level with lower margin of pupil.
Uppermost pectoral ray on horizontal through middle of
suborbital space. Upper lobe short, not nearly reaching anal-
fin origin. Lin deeply notched, 21 (16+2+3), rudimentary
rays absent. Lower lobe comparatively long, of 3 rays,
uppermost longest (93% UPL); lowermost inserted just behind
the vertical through posterior margin of eye. Membrane of
proximal surface of upper lobe attached at bases of rays, of
notch at about Vs ray length, of lower lobe at about Vs lobe
length. Radials 3+0, round. Lenestrae absent. Scapular helve
short, with small upper rib; coracoid helve long, thin.
Body relatively shallow, its dorsal contour almost
straight, greatest depth (at dorsal-fin origin) less than head
length (ca 90% HL). Horizontal through midbody touches
lower margin of eye. Preanal length 35% SL. Vertebral
column almost straight anteriorly. Abdominal vertebrae 10.
Parapophyses of 10th vertebra short, not forming a haemal
spine. Lirst dorsal-fin ray rudimentary, between 5th and 6th
neural spines; next 6 dorsal rays thin, shortened. Lree
interneurals absent. Costal ridges absent. Epineural ribs on
3rd to 6th vertebrae, thin, their length not exceeding the
length of 2.5 vertebral bodies. Epipleural ribs on 4th-llth
vertebrae, short and thin, lengths about 1.5 vertebrae. Caudal
fin 4/4, procurrent rays absent. Anus on vertical just behind
eye. Skin thin, translucent. Gelatinous tissue poorly
Stein et al .: Australian liparids 371
Figure 25. Paraliparis avellaneus n.sp. A, holotype, NMV A5873, 6 ,149 mmTL, 132 mm SL. B, ventral view of mouth
and upper jaw tooth plate. C, P 735, cleared and stained right pectoral girdle, fin rays reconstructed from left side.
developed. Vertical fins overlapping caudal about Vi of its
length. Pyloric caeca 5, similar in length, 14.5% SL.
Colour. Head and body pale nut-brown, slightly darker
posteriorly. Mouth grey, tongue pale, densely black-dotted.
Inner surface of subrostral fold light nut-brown. Gill arches
black-dotted. Branchial cavity dark brown. Peritoneum
black. Stomach and pyloric caeca pale.
Distribution. Off South Australia at 1090-1160 m.
Etymology. The specific epithet derives from the Latin
avellaneus , nut-brown.
Comparative notes. Paraliparis avellaneus is a member
of Group I, and is similar to P. eastmani and P. brunneo-
caudatus in having an oblique mouth, but differs from the
former in the number of vertebrae 65 (v. 69), dorsal-fin rays
58 (v. 64), pectoral radials 3+0 (v. 2+0+0), and in the absence
of procurrent caudal-fin rays (v. 1+3/3+1), tiny teeth (v.
large, stout), lower jaw included (v. protruding), eye almost
entering dorsal profile of head (v. distinctly below it), body
colour light brown (v. black), absence of diastemae (v. present),
and light brown inner surface of the subrostral fold (v. black-
dotted). See description of P brunneocaudatus, below, for
differences from it.
Paraliparis badius n.sp.
Fig. 26
Material examined. Holotype CSIRO T1981-01,
juvenile, 90 mm TL, 82 mm SL. Coordinates of capture
location unknown, off Tasmania, depth unknown, 20 Oct.
1984; radiograph 687B; pectoral girdle 781.
Diagnosis. Vert. 65, D 63, A 55, C 8, radials 3, round. Mouth
inferior, lower jaw included. Subrostral fold absent. Teeth
simple, not large, but prominent. Chin pores closely set,
with common thin skin fold anteriorly. Gill opening short,
half of eye. Lower end of gill opening and uppermost
pectoral fin on horizontal with upper margin of pupil. P
20-21 (14-15+2+4), rudimentary notch rays absent. Head
20% SL, eye large, 29% HL. Preanal length c. 33%, aAf
short, 15.9% SL. Vertical fins overlap half of caudal. Head
brown, mouth black, tongue black-brown-dotted.
Further description. Counts: D 63, A 55, P 20-21, C 8 (4/
4), Vert. 65 (9+56), radials 3 (3+0), gr 5, pc 5, pores 2-6-7-
1. Ratios: HL 20.2, its width 11.5 (56.6), and depth 14.6
(72.3), bd 14.0 (69.3), bdA 13.4 (66), preD 24.4, preA 32.7,
ma 13.4, aAf 15.9, UPL 13.4 (66), LPL 12.8 (95% UPL), E
5.9 (28.9), gs 3.0 (15.1), sn 6.7 (33.1), po 9.5 (47), io 7.4
(36.7), so 3.7 (18), uj 9.0 (44.6), lj 8.3 (41.0).
Head moderately large, about 4.9 in SL, not deep at
occiput but greatly sloping anteriorly; quite compressed
(width 57% HL), depth 1.3 its width. Ventral surface of
head flat. Eye large, almost 3.5 in HL (probably slightly
smaller in adults), almost touching dorsal profile of head.
Pupil about half eye diameter. Interorbital width 1.3 eye,
suborbital distance 0.6 eye. Snout wide, gelatinous, its
length 1.1 eye; profile slanted, clearly projecting beyond
upper jaw, highest point on horizontal with upper margin
of pupil. Subrostral skin fold absent, upper lip entirely
visible. Nostril on level with upper half of pupil. Mouth
inferior, its cleft reaching to below anterior margin of eye,
lower jaw included. Teeth simple, not small, prominent,
about 15 rows of up to 7 teeth each anteriorly in jaws.
Diastema absent in both jaws. Circumoral pores small, not
contoured, rims not thickened. Chin pores almost touching
each other, interspace pigmented, a t hin anterior skin fold
present. In ventral view, upper tooth plates entirely visible.
Postorbital length short. Gill opening small, about half of
eye diameter, lower end on horizontal with upper margin
of pupil. Opercular flap small, rounded, its tip slightly above
level of upper margin of eye.
372 Records of the Australian Museum (2001) Vol. 53
Figure 26. Paraliparis badius n.sp. A, holotype, CSIRO T1981-01, juvenile, 90 mm TL, 82 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 781, cleared and stained right pectoral girdle, partially reconstructed
using detail from left side. Incompletely ossified.
Uppermost pectoral ray level with upper margin of pupil.
Upper pectoral lobe long, reaching to above anal-fin origin. P
(L) 21 (15+2+4), (R) 20 (14+2+4). Rudimentary rays absent.
Origin of lower lobe rays below first quarter of postorbital
space. Pectoral skin missing. Cartilaginous basal lamina with
3 (3+0) round radials, fenestrae absent. Scapula, coracoid and
radials barely ossified, consisting mainly of cartilage.
Body low, maximum depth about 70% HL, in adults
probably slightly deeper. Dorsal contour gently curves
ventrally anteriorly and posteriorly from above anal-fin
origin, ventral profile straight. Preanal distance short.
Horizontal through midline anteriorly touching upper
margin of eye. Free dorsal interneurals absent; first dorsal
ray between vertebrae 4 and 5. Anterior dorsal rays not
rudimentary, embedded in gelatinous tissue. Vertical fins
overlapping caudal slightly more than one-half its length.
Anus below midst of postorbital space, aAf short. Skin
lacking prickles, quite thin, semitransparent. Pyloric caeca
similar.
Colour. Head and body uniformly brown, blackish around
gill opening and on chin. Mouth black, tongue black-brown
dotted, tooth plates pale. Subrostral fold inside and lips
similar in colour to head. Circumoral pores whitish pale,
distinctly contrasting with head colour. Branchial cavity
dark grey, gill arches dotted. Peritoneum dark brown, pyloric
caeca and stomach light.
Distribution. Off Tasmania, depth unknown.
Etymology. Badius from the Latin meaning “dark brown”,
a reference to the overall body colour.
Comparative notes. Paraliparis badius is in group II. It
differs in its large eye, short gill opening equal to half of
eye, absent subrostral fold, chin pores almost touching,
common skin fold anteriorly, aAf short, vertical fins
overlapping half of caudal, and by its brown colour. It is
most similar to P. plagiostomus, but differs from it in its
brown colour (v. brownish-black), shorter mouth cleft
reaching to below anterior margin of eye (v. its centre), larger
eye about 29 (v. 23)% HL, and shorter distance from anus
to anal-fin origin 16% SL (v. 19). In addition, the subrostral
skin fold is absent, so that the upper lip is entirely visible
(v. a wide subrostral skin fold entirely covering the upper
lip), pale tooth plates (v. dark grey), normal sized teeth (v.
tiny, tooth plates smooth), and circumoral pores not
contoured (v. distinctly contoured).
Paraliparis brunneocaudatus n.sp.
Fig. 27
Material examined. Holotype CSIRO T1980-01, S ,142
mm TL, 127 mm SL. FV Petuna Endeavour, collection
coordinates unknown, W coast of Tasmania, collection depth
unknown, Apr. 1984; radiograph 681 A; pectoral girdle 780.
Diagnosis. Vert. 66, D 62, C 8, radials 2. Eye low on side
of head, suborbital space equal to half of eye. Mouth
oblique, lower jaw included. Teeth strong, conical. Chin
pores not in a pit. Dorsal end of gill opening on horizontal
through eye centre. Pectoral fin 20-21 (15-16+2+3), with
long lobes, rudimentary notch rays absent. HL 19% SL,
preA 37%. Anus on vertical just behind eye, aAf long.
Colour bright reddish-brown, end of tail darker, blackish-
brown. Skin thin, semitransparent. Mouth and tongue
blackish-brown, brightly dotted.
Further description. Counts: D 62 (5+57), A 53, P 20-21, C
8 (4/4), Vert. 66 (11+55), radials 2 (2+0+0), pc 6, gr 7, pores
2-6-7-1. Ratios: HL 19.0, its width 10.6 (56.0), and depth 15.0
(79.0), bd 15.7 (83), bdA 18.1 (95), preD 21.6, preA 37.0, ma
15.0, aAf 24.9, UPL 14.2 (75), LPL 13.4 (94% UPL), NL c.
5.5 (39% UPL), E 4.7 (24.9), rim of dense whitish skin around
eye 6.1 (32.4), so 2.8 (14.5), gs 4.3 (22.8), sn 6.3 (33.2), io 7.1
(37.3), uj 8.8 (46.5), lj 7.9 (41.5), pc 3.9.
Head small, 5.2 in SL, depth shallow at occiput (c. 80%
HL), its dorsal contour only slightly sloping anteriorly. Head
very compressed, its width 56% HL, depth 1.4 its width.
Stein et al .: Australian liparids 373
Figure 27. Paraliparis brunneocaudatus n.sp. A, holotype, CSIRO T1980-01, <S, 142 mm TL, 127 mm SL. B,
ventral view of mouth and upper jaw tooth plate. C, P 780, cleared and stained right pectoral girdle.
Snout deep, its length 1.4 eye, slightly projecting above
upper jaw, its highest point above eye level. Subrostral fold
well developed, entirely covering upper lip. Eye large, 4 in
HL, upper margin far below dorsal profile of head,
suborbital distance very short, about half eye diameter. Eye
surrounded by thick, whitish, dense skin, its diameter
slightly larger than eye diameter, pupil about 3 A eye diameter.
Interorbital space 1.5 eye. Nostril pore-like, with raised rim,
on horizontal with upper half of pupil. Mouth oblique,
symphysis of upper jaw horizontal with lower margin of
eye. Mouth cleft reaching to below anterior margin of eye.
Lower jaw included, upper tooth plates entirely visible in
ventral view. Lower jaw tapering anteriorly, tip of chin
rounded, not gelatinous, quite deep below oral cleft. Teeth
stout, quite sharp, conical, in 21-22 rows of 8-9 teeth
anteriorly. Diastema of upper jaw wide, about half of tooth
plate in width, that in lower jaw narrower. Circumoral pores
small, not in pits or protruding. Chin pore pair separated by
about their diameter, interspace pigmented. Gill opening
small, 0.9 eye diameter, entirely above pectoral base; upper
end on horizontal through eye centre, lower end below level
of eye. Opercular flap small, ear-like, tip horizontal with
lower margin of pupil, gill opening comparatively long.
Upper pectoral-fin ray slightly below level of lower
margin of eye. P (L) 20 (15+2+3), (R) 21 (16+2+3), clearly
notched, rudimentary rays absent. Upper lobe long, but not
reaching anal-fin origin. Lower lobe also long, length c.
94% UPL. Pectoral skin attached almost at bases of upper
lobe rays, elsewhere unknown. Basal cartilaginous lamina
of pectoral girdle with two small radials, both located
dorsally just below scapula; uppermost round, lower (R2)
notched on both sides. Two fenestrae present at upper and
lower margins of R2. Helve of scapula stout, with upper lateral
rib. Coracoid has an elongated helve with upper lateral rib.
Body low, its greatest depth above anal-fin origin, 95%
HL, dorsal contour only slightly curved ventrally. Horizontal
midline of body anteriorly touching lower margin of eye.
Preanal 37% SL. Anteriormost 5 dorsal rays very short.
Interneural of 1st dorsal ray between 3rd and 4th neural
spines, free interneurals absent. Parapophyses of the last
abdominal vertebrae not joined together. Haemal spine of
12th (first caudal) vertebra slightly shorter than the next.
Costal ridges absent. Epineural ribs present on 3rd to 18th
vertebrae, anteriormost thin and short, others not longer
than three body vertebrae. Epipleural ribs on 3rd-16th
vertebrae thin, anteriormost not longer than 2.5 body
vertebrae. Vertical fins overlapping caudal about half. Anus
on vertical just behind eye. Pyloric caeca 6, sizes similar, c.
4.7% SL. Skin thin, unprickled, semitransparent. Gelatinous
subcutaneous layer weakly developed.
Colour. Head and trunk very pale, bright reddish-brown,
darker posteriorly, end of tail dark, blackish-brown. Lips
and chin darker than head, blackish; inner surface of
subrostral fold black-dotted; mouth and tongue blackish-
brown, brightly dotted. Pectoral fin dark brown. Peritoneum
blackish-brown, stomach and pyloric caeca pale. Branchial
cavity blackish, gill arches dusky.
Distribution. West coast of Tasmania, depth unknown.
Etymology. Prom Latin brunneo and caudatus, brown¬
tailed, referring to the more darkly pigmented tail.
Comparative notes. Paraliparis brunneocaudatus belongs
to group I, but is distinguished by its reddish-brown skin
with tail darker than head, narrow suborbital space (c. half
of eye diameter), whitish, dense skin surrounding the eye,
comparatively long distance between mandible and anus,
long pectoral-fin lobes, blackish lips and chin, and black-
dotted inner surface of the subrostral fold. It is most similar
to P. avellaneus but differs in having 2 (v. 3) radials and 2
(v. no) fenestrae in the pectoral girdle, in having normally
developed epineurals on vertebrae 3-18 and epipleurals on
vertebrae 3-16 (v. poorly developed ribs on vertebrae 3-6
374 Records of the Australian Museum (2001) Vol. 53
and 4-11); 1st dorsal interneural between neural spines 3
and 4 (v. between 5 and 6); colour reddish with darker tail
(v. uniformly nut-brown); eye surrounded by a thick whitish
rim (v. not), eye not nearly touching upper contour of head
(v. almost touching), teeth stout, conical (v. tiny), and
diastemae present and wide (v. absent).
Paraliparis brunneus n.sp.
Fig. 28
Material examined. Holotype CSIRO H749-05, 9, 169
mm TL, 151 mm SL. FR V Soela, stn. So 3/86/32,41°45.8’S
144°24.8'E, W coast of Tasmania, W of Granville Harbour,
1000-992 m, 16 May 1986; radiograph 682 A; pectoral
girdle 741.
Diagnosis. Vert. 65, D 58, 2 anterior rays shortened, C 8,
radials 4, two with rudimentary notches. Mouth subterminal,
lower jaw subterminal. Teeth small. Chin pores one pore
diameter apart, not in a pit but with anterior skin fold. Gill
opening ventral end slightly below level of lower margin
of eye. Pectoral fin 23 (18+1-1-4), rudimentary rays absent.
HL 19.9% SL, preA 35%. Body dark brown, peritoneum
black, mouth black, tongue densely black-dotted.
Further description. Counts: D 58, A 53, P 23, C 8, Vert.
65 (11+54), radials 4 (3+1) with 3 rudimentary fenestrae in
pectoral girdle; pc 6, gr 7, pores 2-6-7-1. Ratios: HL 19.9,
its width 11.7 (59), and depth 14.2 (72), bd 17.9 (90), bdA
16.2 (82), preD 24.5, preA 35.0, ma 13.9, aAf 22.6, UPL
13.2 (67), LPL 11.3 (57% HL, 85% UPL), NL 3.3 (25%
UPL), E 4.8 (24.0), gs 4.0 (20), sn 7.3 (36.7), po 9.4 (47.3),
io 8.5 (40), so 3.4 (17), uj 8.7 (44), lj 7.6 (38.3), pc 5.0.
Head small, 5.0 in SL, quite low and compressed, its
depth 1.2 its width. Dorsal contour gradually sloping
anteriorly. Snout large, deep, rounded, its length 1.5 eye,
most dorsal point horizontal with eye centre; not gelatinous,
slightly projecting anterior to upper jaw. Subrostral fold
present, deep but not covering upper lip entirely. Nostril
small, pore-like, horizontal with eye centre. Eye quite large,
its upper contour almost touching dorsal margin of head;
suborbital space about 0.7 eye, pupil about half eye
diameter. Interorbital 1.7 eye diameter. Mouth horizontal,
subterminal, cleft reaching to below anterior margin of eye.
Lower jaw subterminal, slightly shorter than upper, not
included. Teeth simple, small, only slightly prominent, in
24-25 rows of up to 8-9 teeth anteriorly. In ventral view
when upper lip is moved, margins of upper tooth plates
visible. Diastemae absent. Lower jaw tapering anteriorly,
chin rounded, deep. Lower jaw below oral cleft deep.
Circumoral pores small, contoured; chin pores longi-
tudinally-oval, interspace unpigmented, equal to their
diameter, not in a pit but with thin skin fold anteriorly.
Distance between nasal pores Vs eye diameter. Gill opening
short, 0.8 eye diameter, its dorsal end horizontal with upper
margin of eye, ventral end slightly below level of lower
margin. Gill opening not vertical, its dorsal end in front of
its ventral end. Opercular flap rounded, covering 3 A of gill
opening, its tip level with upper margin of pupil.
Uppermost pectoral ray horizontal with lower margin of
eye, upper lobe not reaching anal-fin origin. P 18+1+4,
rudimentary rays absent. Lowermost ray inserted behind
vertical through posterior margin of eye. Skin of proximal
side of upper lobe attached at about % distance from fin
base to ray tips, in notch almost to tips, in lower lobe about
Vi. Radials 3+1, R2 largest; R2 and R3 with tiny rudimentary
notches. Three rudimentary fenestrae present. Scapula with
rudimentary notch; helve short, with upper lateral rib. Helve
of coracoid long, with two ribs.
Body elliptic, shallow, maximum depth at anterior of
dorsal fin. Upper and lower body profiles are similarly
curved. Tail quite deep anteriorly, posterior half rather thin.
Horizontal midline of body passes below eye. Anterior
Figure 28. Paraliparis brunneus n.sp. A, holotype, CSIRO H749-05, $, 169 mm TL, 151 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 741, cleared and stained right pectoral girdle.
Stein et al .: Australian liparids 375
halves of dorsal and anal fins entirely embedded in
gelatinous tissue, entirely covered by fin membrane, tips
of rays not projecting. Anterior two dorsal rays short.
Interneural of first dorsal ray between neural spines 5 and
6, two free interneurals present anteriorly between neural
spines 3, 4, 5. Vertebral column almost completely straight
anteriorly. Haemal spine of vertebra 11 short, about half as
long as next. Epineural ribs on vertebrae 2-15, thin, as long
as 3 vertebrae. Epipleural ribs on vertebrae 3-18, shorter
than epineurals, as long as two vertebrae. Slight keel-like
ridge above anterior half of pectoral fin present on side of
body. Vertical fins overlapping caudal almost to its midpoint.
Anus about below middle of postorbital space. Skin quite
thick, prickles absent. Subcutaneous gelatinous layer
moderately developed. Pyloric caeca similar. Eggs unripe.
Colour. Head and body uniformly dark, blackish-brown,
lips slightly paler than head. Subrostral fold black, densely
dotted inside. Mouth black, tongue densely black-dotted,
tooth plates pale, gill arches densely dotted, almost black,
branchial cavity black. Pores at least on lower jaw whitish,
contrasting with dark skin. Peritoneum black, stomach and
pyloric caeca pale.
Distribution. West coast of Tasmania, 1000-992 m.
Etymology. From the Latin brunneus, brown, in reference
to the dense brown colour of the body.
Comparative notes. Paraliparis brunneus belongs to group
IIIc and is most similar to P. auriculatus and P. atrolabiatus
in having uniform black-brown colour, 3+1 radials and 3
rudimentary fenestrae in pectoral girdle, whitish contoured
pores on the lower jaw, and the pectoral-fin upper lobe inner
side free for 3 A of its length. Paraliparis brunneus differs
from P. auriculatus in having a longer snout (v. very slanted,
almost absent in lateral view), radials 3+1 (v. 3+0), 3
rudimentary fenestrae (v. 2) in pectoral girdle, and anus
located below mid-postorbital space (v. below gill opening).
It differs from P. atrolabiatus in its dark, blackish-brown
colour (v. lighter, brown with yellowish head), radials 3+1
(v. 2+0+0) fenestrae 3 (v. absent), P 18+1+4 (v. 16+2+3),
and nephrohaemal canal absent (v. present).
Paraliparis coracinus n.sp.
Fig. 29
Material examined. Holotype CSIRO H1935-02, 9,
unknown TL (tail missing), 170+ mm SL. FRV Soela, stn.
So 1/89/56, 37°34.53'S 138°57.00’E, off South Australia,
W of Cape Martin, 1205-1175 m, 1 Feb. 1989; radiograph
682 E2; pectoral girdle 795.
Diagnosis. Vert. 62+ (11+51+), D 54+, C unknown, radials
4, the lowest half-moon shaped, forming part of posterior
margin of basal cartilaginous lamina. Mouth subterminal,
lower jaw included. Teeth not large, simple. Chin pores
touching each other, anterior skin fold present. Subrostral
fold absent. Gill opening reaching ventrally in front of 4th
pectoral ray. Ventral end of gill opening and uppermost
pectoral ray on horizontal with upper margin of pupil.
Pectoral fin 16-17+2r+3, two notch rays rudimentary. Head
less than 20% SL, eye large, 27.0% HL, snout short, deeply
rounded. Body deep, about 1.3 HL, slight costal ridge
present. Head black, peritoneum black.
Figure 29. Paraliparis coracinus n.sp. A, holotype, CSIRO H1935-02, 9, unknown TL, 170+ mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 795, cleared and stained right pectoral girdle.
376 Records of the Australian Museum (2001) Vol. 53
Further description. Counts: D 54+, A 50+, P 21-22, C
unknown, Vert. 62+(ll+51+), radials 4 (3+1), one fenestra
present; pc 5, gr 8, pores 2-6-7-1. Ratios: HL less than 20%
SL; head width 54% HL, its depth 88, bd c. 132, bdA 100,
preD 129, preA 176, ma 65, aAf 117, UPL 66, LPL damaged;
sn 29.4, E 27.0 (measured between the edges of the retina), po
53, io 39.7, so 17, gs 20.6, uj 47.6, lj 42.3, pc 33.
Head slightly less than 20% SL, deep, compressed, depth
1.6 its width. Dorsal contour greatly rounded from deep
occiput to snout tip, ventral profile straight. Eye very large,
3.7 in head, almost entering upper contour of head.
Suborbital space about % eye diameter. Pupil about half
eye diameter; interorbital 1.4 eye. Snout comparatively
short, deeply rounded, its length from symphysis of upper
jaw to anterior margin of eye equal to the latter. Snout most
prominent on level with lower margin of eye. Subrostral
fold absent, upper lip entirely visible; lower lip fold covered
by upper lip. Nostril pore-like, level with lower margin of
pupil. Mouth subterminal, oral cleft reaching to below eye
centre, posterior of upper jaw extending to below posterior
margin of pupil. Lower jaw shorter than upper, included.
Upper tooth plates entirely visible in ventral view. Teeth
simple, not large, in 27-32 rows of up to 9-10 teeth
anteriorly. Teeth at distal ends of rows (anterior edge of
tooth plates) tiny; interior teeth larger, especially in upper jaw.
Diastema of upper jaw wide, in lower jaw narrower. Lower
jaw below mouth cleft deep; chin skinned, honeycomb tissue
clearly visible, well developed. Circumoral pores not large,
chin pores touching, interspace unpigmented, a thin anterior
skin fold present. Gill opening short, 0.7 eye diameter, its dorsal
end above eye level, ventral end on level with ventral end
on level with pupil, reaching ventrally to 4th pectoral ray.
Opercular flap triangular, sharp-angled, its upper side deeply
notched, covering % of gill opening.
Uppermost pectoral-fin ray level with upper margin of
pupil. Upper lobe not reaching to anal-fin origin. P 16+2+3
(L), 17+2+3 (R); both notch rays rudimentary. Base of lower
lobe rays quite far posterior, below last quarter of postorbital
space. Ends of lower lobe rays missing. Pectoral girdle very
unusual, radials 3+1, two upper large, R2 with rudimentary
ventral notch, R3 small, round. R4 unusual, in shape a half¬
moon (half-round, hoof-like), its straight side even with and
forming posterior margin of cartilaginous basal lamina
exactly between rudimentary notch rays, its rounded side
notched. Interradial fenestrae absent, but one unusual
fenestra present at anterior surface of notched R4.
Body deep (132% HL), maximum depth at beginning of
dorsal fin; upper contour of body abruptly rounded
anteriorly. Ventral contour of body almost straight, dorsal
very curved. Anterior dorsal and anal rays entirely
embedded in gelatinous tissue. Parapophyses of vertebra
11 form a short haemal spine, absent on others. Interneural
of first (rudimentary) dorsal ray between neural spines 5
and 6, 1 free anterior interneural present. Epineural ribs
present on vertebrae 2-13, epipleural ribs on 3-25, length
of anterior ribs of both series not longer than 3 vertebrae,
but stout; a slight costal ridge present. Anus below posterior
third of postorbital space. Ovarian eggs at different stages
of maturity, largest 2.8 mm. Body partially skinned. Skin
on head without prickles, dense, opaque. Subcutaneous
gelatinous tissue moderately developed. Pyloric caeca of
similar lengths.
Colour. Body partially skinned, reddish-brown shreds of
dermis remain on muscles. Remnants of skin on head ink-
black, body probably the same colour. Lips lighter, dark
grey, mouth dark grey, tongue grey-dotted, tooth plates dark.
Branchial cavity ink-black, gill arches grey. Peritoneum
black, pyloric caeca and stomach pale.
Distribution. Off South Australia in the Great Australian
Bight at about 1200 m.
Etymology. Coracinus, Latin, meaning jet-black.
Comparative notes. Paraliparis coracinus belongs to
group IIIc, and is so distinctive we have no reluctance in
describing it as new. It is particularly noteworthy for its
pectoral girdle, with two rudimentary rays and a hoof-like
R4 between them, forming the posterior margin of the girdle
at that point. This is similar to the arrangement in P. hureaui
Matallanas, 1999, and P charcoti Duhamel, 1992, which
have pectoral girdles with both R3 and R4 hoof-shaped,
both forming part of the posterior margin of the pectoral
girdle. Other characters distinguishing this species from all
others include the gill opening reaching to 4th pectoral ray
(v. in all others no farther than the second ray), absence of
subrostral fold, large eye (27% HL), chin pores touching,
and honeycomb tissue on the chin.
Paraliparis costatus n.sp.
Lig. 30
Material examined. Holotype CSIRO H561-02, 3 , 224
mm TL, 204 mm SL. LRV Soela, stn. So 3/86/33,41°51.4’S,
144°23.8'E, W coast of Tasmania, W of Granville Harbour,
1366-1370 m, 16 May 1986; radiograph 684A; pectoral
girdle 733. Paratype CSIRO H1378-01, $, 261 mmTL,
235 mm SL. LV Petuna Endeavour , Stn 101/18, 42°12'S,
144°38'E, W coast of Tasmania, W of Cape Sorell, 1042-1080
m, 21 Apr. 1988; radiograph 684B; pectoral girdle 731.
Diagnosis. Vert. 70-71, D 62-66, P 22-24, C 8, radials 4,
round. Snout high, blunt, and large; nasal pores widely
spaced, chin pores not in a pit and without skin fold
anteriorly. Mouth horizontal, terminal. Anterior 7-8
epineural and epipleural ribs straight, elongated, thickened.
Keel-like lateral ridge protruding along and above anterior
part of abdominal cavity. Gill opening lower end below eye
level. Head 19.3-19.8% SL, preA length about 40, aAf 22-
26% SL. Body black; orobranchial cavity black-dotted to
black; peritoneum black.
Further description. Counts: D 66 [62], A 57 [56], P 22
[24], C 8 in both (1+3/3+1), Vert. 71 [70] (11+59-60),
radials 4 (3+1), pc 5 [3], gr 10 [8]. Ratios: head 19.8 [19.3],
its width 12.9 [12.7], and depth 13.8 [18.7], bd 16.2 [23.4],
bdA 16.3 [19.6], preD 25.1 [28.5], preA 39.1 [40.3], ma
14.0 [12.8], aAf 21.7 [26.4], UPL 13.2 [12.7], LPL 11.8
[10.8], E 4.7 [4.7], sn 6.7 [7.4], gs 3.6 [4.5], io 9.8 [10.6], po
9.8 [10.6], so 2.9 [—], uj 9.6 [9.2], lj 9.1 [9.2], pc 5.3 [5.5]; as
% HL: UPL 66.8 [65.6], E 23.8 [22.0], gs 19.5 [21.0], sn 36.6
[35.0], so 15.6 [—], io 54 [50], uj 48.3 [47.8], lj 44.6 [47.6].
Head small, not as deep as body, moderately compressed;
its width 0.7 [0.6] HL and depth 1.1 [1.4] its width. Dorsal
contour of head only slightly sloping to nearly vertical, deep,
bluntly rounded snout, slightly protruding anterior to upper
Stein et al .: Australian liparids 377
Figure 30. Paraliparis costatus n.sp. A, holotype, CSIRO H561-02, 6, 224 mm TL, 204 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 733, cleared and stained right pectoral girdle.
lip. Subrostral fold deep, partially covering upper lip.
Interorbital space flat or a little convex, 1.7 [1.7] eye
diameter. Mouth terminal, horizontal; lower jaw sub¬
terminal. Mouth cleft extending to below anterior of orbit,
maxilla reaching to below mid-eye; teeth simple, small,
subconical, closely set in about 30 oblique rows of up to 8-
10 teeth each anteriorly, forming moderately wide bands in
both jaws. Diastemae present in both jaws, that in lower
wider than in upper; lower jaw long, about Vi HL. Eye
moderately large, 4.2 [4.1] eye diameter, well below dorsal
outline of head. Suborbital space 2 A eye. Nostril with low
raised rim, level with upper margin of pupil. Pupil half eye
diameter. Circumoral pores small, round; chin pores well
spaced, interspace between them (pm,-pm,) equalling two
pore diameters, pigmented. Nasal pores unusually widely
spaced, the lower just above upper lip, the upper at the top
of snout; distance between them equals eye diameter. Gill
opening completely above pectoral-fin base; its upper end level
with upper margin of pupil, lower end level with middle of
suborbital space; opercular lobe small, roundly pointed, on
horizontal with lower margin of eye.
Uppermost pectoral ray level with middle of suborbital
space, lowermost ray inserted below posterior margin of
eye. In the holotype P 22 (15+4+3), in the paratype 24
(16+4+4). Upper pectoral lobe rather short, not reaching
anal-fin origin. Pectoral-fin notch moderately deep, dividing
fin into two distinct lobes. Notch rays 3-4, shortened
gradually ventrally but not becoming rudimentary; more
widely spaced in notch but more closely spaced dorsally,
difficult to distinguish from upper lobe rays. Lower lobe
short, of 3-4 rays, slightly shorter than upper lobe. Skin on
proximal surface attached at bases of all rays. Pectoral girdle
morphology identical in both specimens: radials 4 (3+1),
round, rather large, rudimentary fenestrae absent. Scapula
and coracoid with well-developed shafts; additional side
ribs absent. Coracoid without opening.
Body elliptical, relatively slender, maximum depth at
anal-fin origin, 6.6 in SL, less than HL (0.8). Anterior dorsal-
fin rays rudimentary. Interneural of first dorsal ray between
neural spines 5 and 6, one free interneural present anteriorly,
between 4th and 5th neural spines. Epineural ribs present
on 2nd-16th, epipleural ribs on 2nd-15th vertebrae, anterior
ones short, but on vertebrae 4-7 longer, straight, thickened
in the middle (up to 3-4 vertebrae long); becoming
gradually shorter caudally, discernible up to vertebrae 13-
16. A keel-like costal ridge present above and along anterior
part of abdominal cavity. Horizontal midline passes through
lower margin of eye. Caudal fin of 6 principal rays, a single
procurrent ray present above and below (1+3/3+1),
overlapped by dorsal and anal fin rays to one third of its
length. Anus below middle of postorbital space, preA long.
Skin thin, semitransparent; subcutaneous gelatinous layer
moderately developed. Pyloric caeca wide, similar. Urogenital
papilla in male short, conical, below % postorbital space.
Lemale (paratype) with three generations of oocytes, largest
of 3.2 mm diameter (its belly enlarged because of eggs so
body depth much larger than in male holotype).
Colour. Skin black, head and belly darker than body of
which pale muscles are visible through skin. Subrostral fold
of the same colour as head, lips dark grey. Mouth and tongue
grey, tooth plates pale. Pores whitish, distinctly contrasting
with head colour. Branchial cavity dark-dotted to black,
gill arches dark-dotted, peritoneum black, stomach pale,
pyloric caeca and posterior part of intestine grey.
Distribution. West coast of Tasmania between 1042-1370 m.
Etymology. The specific epithet is formed from the Latin
word costa, rib, to emphasize the unusual development of
epineural and epipleural ribs in this species.
Comparative notes. Paraliparis costatus is in group Ilia.
It has well-developed epipleural and epineural ribs forming
a keel-like lateral ridge above the pectoral fins, a large, blunt,
378 Records of the Australian Museum (2001) Vol. 53
deep snout with widely spaced nasal pores, 4 round radials,
maximum body depth at anal-fin origin, and pale circumoral
pores. The paratype (a ripe female) differs from the holotype
(unripe adult male) in having a deeper head and body [hd
86, bd 110% HL] owing to having ripe ovarian eggs. The
ratio of head depth to width is 1.5 v. 1.0. The teeth also
appear stronger and stouter, the diastemae are wider, and
the remnants of the skin on the body have a reddish tint.
Paraliparis costatus differs from nearly all Paraliparis (with
the exceptions of P. dewitti, P. lasti, and Paraliparis sp. 2)
in the unusually well-developed ribs forming a costal ridge.
See description of P. dewitti for comparison. Paraliparis
costatus differs from P. lasti, which has similar chin pores,
by its black (v. reddish-brown) colour, more ventral gill
opening (lower end below eye v. level with eye centre),
and radials 3+1 (v. 3+0).
Paraliparis csiroi n.sp.
Fig. 31
Material examined. Holotype NMV A5874, ripe $, 182
mm TL, 163 mm SL. FR \ Soela, stn. 01/88/86, 38°37.58’S
141°01.12'E, 60 km S of the Victoria-South Australia border,
1080-1110 m, 8 Feb. 1988; radiograph NMV A5874;
pectoral girdle 739.
Diagnosis. Vert. 66, D c. 60, C 8, radials 3, uppermost
ventrally notched. Parapophyses of vertebrae 5-11 fused
to form a nephrohaemal canal. Pectoral fin 20 (15+1+4),
one rudimentary ray each present in notch and in lower
lobe. Mouth terminal, short. Chin pores separated by one
pore diameter, in a common oval, pigmented depression on
anterior surface of chin. Gill opening ventral end horizontal
with lower margin of pupil. Head compressed, HL 19.6%
SL, preanal 39% SL. Colour dark brown, lips light grey,
peritoneum black, mouth and tongue grey.
Further description. Counts: D c. 60, A c. 55, P 20
(15+lr+4=3+lr), C 8 (4/4), Vert. 66 (11+55), radials 3 (3+0,
see below), pc 5, gr 8, pores 2-6-7-1. Ratios: HL 19.6, its
width 10.7 (54) and depth 17.4 (88), bd 24.6 (125), bdA
21.5 (109), preD 25.1, preA 38.7, ma 13.9, aAf 24.6, UPL
13.9 (70), LPL 9.8 (85% HL, 71% UPL), E 4.6 (23.4), gs
3.3 (17.1), sn 6.4 (32.8), po 10.4 (53), so 2.6 (13.4), uj 9.2
(46.8), lj 6.7 (34.4), io 6.1 (31.3), pc 4.6.
Head small, 5.1 in SL, strongly compressed and deep,
depth 1.6 its width. Dorsal contour gradually sloping
anteriorly to highest point of snout; snout deep and blunt,
not projecting anterior to upper jaw. Distance from upper
jaw symphysis to eye 1.4 eye. Subrostral fold absent, upper
lip not covered. Eye 4.3 in HL, its upper contour far below
dorsal contour of head, distance from lower margin of eye
to mouth very short, about equal to 0.6 eye diameter. Pupil
slightly larger than half eye diameter. Interorbital narrow,
1.3 eye. Nostril with raised rim, horizontal with upper half
of pupil. Mouth horizontal, terminal, oral cleft reaching to
below anterior margin of eye. Lower jaw included, in ventral
view broadly rounded anteriorly, but in lateral view,
symphysis in reverse angle so that it slopes anteroventrally;
depth shallow below posterior of oral cleft. Teeth simple,
small, only slightly projecting, with blunt tips, in 23-24
regular rows, curved on the anterior surface of tooth plates;
c. 11 teeth in each anterior row. Diastema of upper jaw
slightly narrower than that of lower jaw. Circumoral pores
small; chin pore interspace equal to pore diameter,
pigmented, in a common oval depression located on anterior
surface of gelatinous chin. Gill opening vertical, short, 0.7
eye diameter, entirely above base of upper P ray; opening
vertical; its upper end slightly above horizontal through
upper margin of eye, ventral end level with lower margin
of pupil. Opercular flap small, ear-shaped, dorsally notched;
its tip about level with upper margin of eye.
Base of upper pectoral ray below end of opercular flap,
level with lower margin of pupil; lowermost ray inserted
Figure 31. Paraliparis csiroi n.sp. A, holotype, NMV A5874, ripe $, 182 mm TL, 163 mm SL. B, ventral view of
mouth and upper jaw tooth plate. C, P 739, cleared and stained right pectoral girdle.
Stein et al .: Australian liparids 379
behind vertical through posterior margin of eye. P 15+1
(rudimentary) + 3 (and 1 rudimentary), deeply notched,
rudimentary notch ray clearly seen only with clearing and
staining; upper and lower lobes united by pectoral-fin
membrane; lowermost lower lobe ray also rudimentary.
Upper lobe rays not nearly reaching anal-fin origin, lower
lobe long, length c. 70% UPL. Pectoral-fin skin missing.
Pectoral radials 3 (3+0). The 4th (lowest) radial is probably
undeveloped, a distinctly contoured round opening present
at normal radial location. Upper radial with ventral notch,
others unnotched. One fenestra (f2) present below upper
radial, two small rudimentary slit-like openings present
below scapula and third radial. Helve of scapula with two
lateral ribs, coracoid with long thin unribbed shaft.
Anteroventral part of basal cartilaginous lamina above
coracoid slightly angled, not joined to cleithrum (the latter
not visible in Fig. 31b).
Body deep, leaf-like, greatest depth above posterior half
of abdomen, 4.1 in SL; depth above A exceeds head length
(109%). Dorsal contour gradually rounded anteriorly and
posteriorly. Horizontal midline passes through eye centre.
Anterior dorsal rays gradually lengthening posteriorly,
embedded in gelatinous tissue. First dorsal ray rudimentary,
its interneural between neural spines 4 and 5, a free
interneural present between spines 3 and 4. First anal ray
on vertebra 11, below dorsal ray 7. Parapophyses of
vertebrae 5-11 joined to form short spines, forming a
nephrohaemal canal; spines 5-7 directed anteroventrally;
haemal spine of posteriormost abdominal vertebra (11)
about Vs length of haemal spine of first caudal vertebra.
Epineural ribs on vertebrae 3-14, epipleural ribs on
vertebrae 3-18, all thin, not long, lengths of anterior 7-8
not longer than 2-2.5 vertebrae. Anus below midst of
postorbital space. Caudal fin overlapped by vertical fins
about Vs of its length. Skin thin, slightly transparent; prickles
absent; subcutaneous gelatinous tissue moderately
developed. Pyloric caeca with wide bases and sharp tips, of
similar size. Almost ripe eggs c. 4.1 mm in diameter.
Colour. Head and body dark blackish-brown, body lighter
than head and caudal; lips and chin margin light grey. Chin
pore depression pigmented, pores as dark as head. Anal
region black. Mouth grey, tongue dark grey, branchial cavity
greyish-brown. Peritoneum black, stomach and pyloric
caeca pale.
Distribution. Off South Australia at 1080-1110 m.
Etymology. Named after the Commonwealth Scientific and
Industrial Research Organization (CSIRO), the supporting
agency for Australian fisheries research.
Comparative notes. Paraliparis csiroi belongs to group
Illb, and is distinguished by its nephrohaemal canal, low
eye (suborbital space about half eye diameter), narrow
interorbital, rudimentary single notch ray, deep body (125%
HL) with maximum depth in posterior of abdomen, blunt,
short, snout, short mouth, subrostral fold absent, chin pore
depression on anterior surface of gelatinous chin (a unique
character), and lips paler than head. In group Illb, it is most
similar to P. australiensis, which also has no normally
developed rays in the pectoral-fin notch, but differs from it
in the absence of foramina in the pectoral girdle (v. present),
lower eye (v. touching upper contour of head), chin-pore
pit on the anterior surface of the chin (v. on lower surface),
anus below middle of postorbital head (v. just below rear of
eye), nephrohaemal canal (v. absent), absence of the
subrostral fold (v. deep, almost entirely covering upper lip),
and body paler than head and caudal (v. the same). It is also
similar to P atrolabiatus, but in addition to the above
characters, differs in having 3+0 radials (v. 2+0+0), one
fenestra in the pectoral girdle (v. none), and head darker
than body, blackish - brown (v. head lighter than body,
yellowish-brown).
Paraliparis delphis n.sp.
Fig. 32
Material examined. Holotype CSIRO H749-03, S , 140 mm
TL, 127 mm SL. FRV Soela, So 3/86/32,41°45.8’S 144°24.8'E,
W coast of Tasmania, W of Granville Harbour, 1000-992 m,
16 May 1986; radiograph 680 F 3; pectoral girdle 791.
Diagnosis. Vert. 67, D 61, C 8, radials 2, round. Mouth
subterminal, lower jaw included, teeth simple, small. Chin
pores almost touching, not in a pit but with thin skin fold
anteriorly, interspace unpigmented. Lower end of gill
opening and uppermost pectoral-fin ray horizontal with
pupil. Opercular flap triangular with rounded end. P 15+2+4,
rudimentary rays absent. Nephrohaemal canal present on
vertebrae 5-11. HL 18% SL, preA 31%. Colour very dark,
uniformly brownish-black, mouth grey, tongue grey-dotted,
peritoneum black.
Further description. Counts: D 61, A53, P21 (15+2+4), C 8
(4/4), Vert. 67 (11+56), radials 2+0+0, gr 8, pc 6, pores 2-6-7-
1. Ratios: HL 18.1, its width 11.0 (61) and depth 15.0 (83), bd
16.1 (89), bdA 15.0 (83), preD 21.2, preA 31, ma 12.0, aAf
20.4, UPL 13.0 (71), LPL 11.0 (85% UPL), NL2.2 (17% UPL),
E 4.4 (24.3), gs 3.2 (17.8), sn 6.0 (33.0), po 8.8 (49), io 7.6
(41.7), so 3.1 (17.4), uj 8.3 (45.7), lj 7.7 (43.5), pc 5.5-7.0.
Head small, 5.5 in SL, not very compressed, depth 1.4
its width. Dorsal profile deep at occiput, slanting anteriorly
to rounded snout. Eye not touching dorsal profile of head,
suborbital distance about 0.7 eye. Pupil equals about half
eye diameter. Interorbital width 1.6 eye. Snout deep,
rounded, clearly projecting; gelatinous, 1.4 eye, its dorsal
edge level with lower margin of pupil. Nostril pore-like,
horizontal with lower margin of pupil. Subrostral fold
deeper anteriorly than laterally, where upper lip clearly
visible. Mouth horizontal, subterminal (almost inferior, but
lower surface of head slanted, not horizontal), cleft almost
reaching to below anterior margin of pupil. Lower jaw
included; in ventral view, upper tooth plates entirely visible
anteriorly when upper lip is turned out; nasal and two
anterior infraorbital pores also visible. Teeth simple, small,
only slightly prominent, in about 21-24 regular oblique rows
of up to 11 and 9 teeth each anteriorly in both jaws. In upper
jaw, rows extend as very small teeth onto anterior surface
of tooth plate. Diastemae not clearly developed. Lower jaw
below end of mouth cleft deep. Circumoral pores small,
with raised rims. Chin pores closely set, interspace less than
their diameter, unpigmented, even with chin surface but
with a thin, pigmented, skin fold anteriorly. Gill opening short,
dorsal end slightly above eye, ventral end about level with eye
centre. Gill opening not vertical, dorsal end slightly anterior to
ventral end. Opercular flap triangular, rounded ventrally,
380 Records of the Australian Museum (2001) Vol. 53
Figure 32. Paraliparis delphis n.sp. A, holotype, CSIRO H749-03, 8 , 140 mm TL, 127 mm SL. B, ventral view of
mouth and upper jaw tooth plate. C, P 791, cleared and stained right pectoral girdle.
dorsally unnotched, tip level with upper margin of eye.
Uppermost pectoral ray level with lower margin of pupil,
base of ventralmost ray on vertical just behind posterior
margin of eye. Skin closely attached to body, covering
anterior l A of upper lobe, only posterior of fin free and
movable; lower fin lobe similar, about Vi attached; notch rays
about 80% attached, free parts of upper and lower lobes appear
to be separate, unjoined by fin membrane. Upper pectoral lobe
not reaching to anal-fin origin. P 15+2+4, lowermost notch
ray short, 17% UPL, but not rudimentary. Radials 2+0+0,
round, fenestrae absent. Coracoid with a long thin helve.
Body not deep, distinctly humpbacked, greatest depth
89% HL, at first dorsal-fin ray. Ventral contour of body
almost straight, dorsal contour much curved at dorsal-fin
origin. Horizontal midline of body passes through centre
of eye. Preanal distance short, 31% SL. Interneural of first
dorsal ray between neural spines 4 and 5; one free anterior
interneural present. Anterior of dorsal fin covered by
gelatinous tissue. Parapophyses of vertebrae 5-11 joined,
forming nephrohaemal canal. Costal ridges weakly developed.
Epineural ribs on vertebrae 3-16, short, thin, not as long as
two vertebrae; epipleural ribs on vertebrae 5-11, also slim.
Vertical fins overlapping anterior half of caudal. Anus below
middle of postorbital space. Gelatinous tissue moderately
developed. Skin quite dense, opaque, prickles absent. Pyloric
caeca elongated. Small, cone-like, urogenital papilla present.
Colour. Head, lips, chin, and body uniformly dark,
brownish-black. Mouth dark grey, slightly lighter than head,
tongue grey-dotted. Inner surface of subrostral fold as dark
as head, densely black-dotted. Branchial cavity and gill
arches dark grey. Pores paler inside but not strongly
contrasting with head colour. Peritoneum black, stomach
and pyloric caeca pale. Urogenital papilla black.
Distribution. West coast of Tasmania, 1000-992 m.
Etymology. The name is derived from the Greek delphis ,
dolphin, to which the new species is similar in body shape.
Comparative notes. Paraliparis delphis belongs to
group IIIc. It is distinguished by its hump-backed body,
very dark uniform brownish-black colour, round radials
2+0+0, nephrohaemal canal on vertebrae 5-11, and chin
pores touching with a very narrow unpigmented
interspace. It is most similar to P. atrolabiatus and P.
ater, but differs from the former in its very dark, uniform,
brownish-black colour (v. non-uniform, brown, light-
yellowish head and blackish pectoral fin, opercular flap
and genital papilla), lips similar to head colour (v. darker
than head, blackish), mouth dark grey but lighter than
head (v. dark grey and darker than head), body more
hump-backed and slightly shallower (bd 89 v. 95% HL,
bdA 83 v. 91%), anus below middle of suborbital space
(v. below first third), first dorsal interneural between
neural spines 4 and 5 (v. 6 and 7), one free interneural (v.
2), nephrohaemal canal on vertebrae 5-11 (v. 3-11),
lower jaw included (v. subterminal), chin pores almost
touching (v. spaced by 2 diameters), triangular opercular
flap with rounded tip and unnotched upper edge (v.
rounded), and a longer lower pectoral lobe of 85 (v. 61%
UPL). It differs from P. ater in having 2+0 radials (v.
3+0), shape of opercular flap, preanal length 31 (v. 37),
lower jaw included (v. subterminal), subrostral fold deep
anteriorly (v. almost absent), and other characters.
Stein et al .: Australian liparids 381
Figure 33. Paraliparis dewitti n.sp. A, holotype, CSIRO T889-02, !, 213 mm TL, 192 mm SL. B, ventral view of
mouth and upper jaw tooth plate. C, P 742, cleared and stained right pectoral girdle.
Paraliparis dewitti n.sp.
Fig. 33
Material examined. Holotype CSIRO T889-02, $, 213 mm
TL, 192 mm SL. FV Margaret Philippa, 34°26.5'S 132°04'E,
South Australia, Great Australian Bight, 1175-1118 m, 14Nov.
1984; radiograph 684E; pectoral girdle 742.
Diagnosis. Vert. 65, C 8, radials 4, round; one rudimentary
fenestra present. About 7 anterior epineural and epipleural
ribs long, stout. Mouth horizontal, subterminal, teeth tiny.
Chin pores in common wide-oval depression, interspace equals
twice their diameter. P 22-23 (17+2+3-4), rudimentary notch
rays absent. Prominent costal ridge present. Head 21% SL,
wide (67% HL). Eye diameter half interorbital width. BdA
90% HL, preanal length 34% SL. Colour light nut-brown,
skin semitransparent, peritoneum black.
Further description. Counts: D 59, A 52, P 22-23, C 8 (4/
4), Vert. 65 (12+53), radials 4, pc 6, gr 11, pores 2-6-7-1.
Ratios: HL 21.4, its width 14.3 (67) and depth 18.3 (86), bd
20.1 (93), bdA 19.3 (90), preD 24.5, preA 33.9, ma 14.1,
aAf 19.5, UPL 13.5 (63), LPL 12.0 (56% HL, 88% UPL),
NL 4.2 (31% UPL), E 4.1 (20.2), gs 4.1 (19.0), sn 7.8 (36.5),
postocular 10.6 (49.8), io 8.9 (41.5), so 3.3 (15.6), uj 10.4
(48.8), lj 10.1 (47.6), pc 3.0.
Head moderately large, 4.7 in SL and wide, its depth 1.3
its width. Dorsal contour high at occiput, sloping
anteroventrally from above eye. Snout large, 1.8 eye,
comparatively deep, rounded in lateral profile, significantly
projecting anterior to upper jaw. Subrostral fold shallow,
not covering upper lip. Eye small, 4.9 in HL, upper margin
not touching dorsal profile of head, suborbital space 0.8
eye. Pupil about half of eye diameter. Interorbital width
twice eye diameter. Nostril small, rim low, on horizontal
with lower half of pupil. Mouth horizontal, subterminal,
oral cleft reaching to below anterior margin of pupil. Lower
jaw included. Chin below posterior of oral cleft deep. In
ventral view, when upper lip pushed aside, margins of upper
tooth plates visible; chin wide and blunt. Upper lip wide,
lower lip fold distinct. Teeth simple, sharp, tiny, tooth plates
appear smooth, in about 29-30 oblique rows of 10-13 teeth
each anteriorly. Diastema of lower jaw narrower than in
upper. Circumoral pores small, without thickened rims. Chin
pores small, two pore diameters distant, interspace
pigmented, in a shallow, wide, oval pit placed on the lower
surface of the gelatinous chin. Gill opening 0.9 eye diameter,
entirely above P fin base, slanted anteriorly, its upper end
slightly above horizontal of upper margin of eye, lower end
level with lower margin of pupil. Opercular flap small,
triangular, tip level with upper margin of pupil. Gill rakers
knob-like, prickles absent.
Base of uppermost pectoral-fin ray level with lower
margin of eye, just below lower end of gill opening. Upper
P lobe short, almost reaching anal-fin origin. Pectoral fin
deeply notched, rays quite stout, length about 30% UPL. P
17+2+3-4. Lower lobe rays 3 or 4, lowermost ray inserted
below posterior margin of eye. Pectoral radials rounded, 4
(3+1). One rudimentary fenestra present below scapula.
Scapula and coracoid each with a long, strong shaft, with
an upper lateral rib.
Body humpbacked, depth comparatively shallow (93%
HL), deepest at D origin, depth at A origin similar.
Horizontal midline of body passes through mid-eye. Skin
thin, semitransparent. Subcutaneous gelatinous tissue
moderately developed, anterior dorsal rays embedded. First
dorsal ray interneural between neural spines 4 and 5, free
interneurals absent. Parapophyses of two last abdominal
382 Records of the Australian Museum (2001) Vol. 53
vertebra (11, 12) forming rather short haemal spines.
Epineural ribs on vertebra 3-11, epipleural ribs on vertebra
3-18; anterior 7-8 epipleural and epineural ribs long and
stout, about as long as 4 vertebrae. Ribs producing a
prominent keel-like lateral ridge on each side of body above
first half of pectoral fin. Vertical fins overlap caudal for
about one fourth of its length. Anus below posterior third
of postorbital space. Pyloric caeca short and quite thick,
with rounded ends, of similar size. Eggs small, unripe.
Colour. Body uniformly light nut-brown (without reddish
tint), skin semitransparent, showing pale musculature.
Interspace between chin pores pigmented. Mouth and
tongue light grey, inner surface of subrostral fold and pores
same colour as head; lips and chin slightly lighter. Tooth
plates pale. Peritoneum black, stomach pale, pyloric caeca
grey. Branchial cavity dark greyish-brown, gill arches light
brownish-grey.
Distribution. Off South Australia, Great Australian Bight,
1175-1118 m.
Etymology. Named in honour of Hugh H. Dewitt (1933 -
1995), prominent student of the Antarctic fish fauna.
Comparative notes. Paraliparis dewitti is a member of
group Illb. It is distinguished by having well-developed
epineural and epipleural ribs, a costal ridge, 4 radials, wide
head, large snout, tiny teeth, a large chin pore pit, and short
upper pectoral-fin lobe and pyloric caeca. It is similar to P.
costatus, but differs in lower counts: Vert. 65 (v. 70-71), D
59 (v. 62-66), A 52 (v. 56-57), C without procurrent rays,
4/4 (v. 1+3/3+1), one rudimentary fenestra in pectoral girdle
(v. absent), coracoid with long helve (v. thin, small), chin
pores in a wide oval depression (v. not), nut-brown colour
(v. black), and other characters.
Paraliparis eastmani n.sp.
Fig. 34
Material examined. Holotype AMS 128900-003, S, 203
mm TL, 183 mm SL. FV Kapala, fid. no. K 88-17-06,
33°29.5'S, 152°12.5'E, E of Broken Bay, New South Wales,
1035-1070 m, 1 Sep. 1988; radiograph 28900; pectoral
girdle 743.
Diagnosis. Vert. 69, D 64, P 20 (15+3+2), C 8, with 2
procurrent rays, radials 2, round. Mouth oblique. Teeth stout,
subconical. Chin pores at surface, interspace equal to
diameter. Gill opening small, 0.8 eye diameter, with dorsal
end level with eye centre. HL 19.0% SL, preA 36%. Body
depth equal to head length, bdA 90% HL. Skin thick, not
transparent. Colour solid black, inner surface of subrostral
fold and tooth plates black-dotted.
Further description. Counts: D 64, A 57, P 20, Vert. 69
(11+58), C 8 (1+3/3+1), radials 2 (2+0+0), pc 4. Ratios:
HL 19.0, its width 9.3 (49.0), bd 18.5 (100), bdA 16.9 (89),
preD 16.1, preA 36.0, ma 11.9, aAf 24.6, UPL 12.6 (66),
LPL 12.1 (64.0% HL, 107% UPL), NL c. 3.2, io 8.2 (42.9), so
2.7 (14.4), E 4.9 (25.9), uj 9.7 (51.3), lj 9.2 (48.4), gs 3.9 (20.7).
Head small, greatly compressed, its width nearly half its
length. Dorsal contour of head very slowly sloping to high,
blunt, snout, not protruding anteriorly. Snout 1.2 eye
diameter, its highest point level with upper margin of eye.
Subrostral fold deep. Interorbital space gently curved, broad,
1.7 eye diameter. Eye large, 3.9 in head, very large pupil
almost equal to eye diameter, dorsal margin of eye well
below dorsal profile of head, suborbital distance 0.6 eye.
Nostril with slightly raised rim, level with eye centre. Mouth
oblique, terminal, large, lower jaw projecting. Posterior of
mouth cleft nearly reaching vertical through anterior edge
Figure 34. Paraliparis eastmani n.sp. A, holotype, AMS 128900-003, 6 , 203 mmTL, 183 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 743, cleared and stained right pectoral girdle.
Stein et al .: Australian liparids 383
of eye. Teeth simple, stout, subcorneal, with blunted tips,
closely set in 21 and 28 oblique rows in both jaws, 7-8
teeth in each row anteriorly. A wide diastema present at
symphysis of upper jaw, that in lower jaw narrower; chin
slanted. Chin below posterior of mouth cleft deep.
Circumoral pores small, round, hardly discernible in black
skin. Chin pores very closely set, interspace equals their
diameter, not in a pit or skin depression. Opercular flap
small, triangular, its tip level with lower half of eye, covering
about Vs of gill opening. Gill opening very small, 0.8 eye
diameter, vertical, completely above pectoral-fin base; its
dorsal end level with eye centre, ventral end with middle of
suborbital space.
Uppermost pectoral-fin ray origin about level with
posterior end of oblique upper jaw. P 20 (15+3+2), deeply
notched, its upper lobe of 15 rays, short, not reaching anal-
fin origin. Notch rays three, moderately short, the shortest
Vs the length of upper lobe rays. Lower pectoral lobe distinct,
consisting of two elongate rays only; origin about at posterior
margin of eye. Pectoral membrane of proximal fin surface
attached to body at base of upper lobe, at about 70% of notch
ray length, and at about 33% of lower lobe length. Basal
cartilaginous lamina with two round radials, both located
dorsally just below scapula; foramina absent. Scapula helve
of unusual form, thin and comparatively long with additional
lateral ribs. Coracoid without foramen, helve long.
Body moderately deep, elongated, its greatest depth at
middle of pectoral upper lobe, equal to head length, bdA c.
90% HL. Dorsal contour of body straighter than ventral
contour. Predorsal length short, interneural of first dorsal-
fin ray between 4th and 5th neural spines; one free (rayless)
interneural present anteriorly. Epineural ribs on 2nd-17th
vertebrae, epipleural on 3rd-15th, thin and short, lengths
equal to 1.5-2.5 vertebrae. Parapophyses of lOth-llth
vertebrae joined, forming short haemal spines. Vertebral
column without a curve anteriorly. Caudal fin of 6 principal
rays; a single procurrent ray present above and below (1+3/
3+1). Caudal fin about Vs overlapped by dorsal- and anal-fin
rays. Anus below Vi of postorbital space. Skin thick, opaque,
not loose. Pyloric caeca of similar size, short, about 4.6% SL.
Gelatinous tissue not well developed.
Colour. Body uniformly solid black, palate black; tongue, inner
surface of subrostral fold, branchial cavity, mouth between
lower jaws, and tooth plates black-dotted. Stomach, pyloric
caeca and intestine unpigmented. Peritoneum solid black.
Distribution. Tasman Sea a little north of Sydney, 1035-
1070 m.
Etymology. The new species is named after Joseph T.
Eastman in honour of his valuable studies on the natural
history, physiology, and origins of the Antarctic fish fauna.
Comparative notes. Paraliparis eastmani belongs to group
I. It is distinguished by its solid black colour, thick opaque
skin, and short gill opening (equal to 0.8 eye diameter). It
is most similar to P. brunneocaudatus, but differs in its black
(v. reddish-brown) colour, absence of pectoral fenestrae (v.
2), projecting lower jaw (v. included), dark tooth plates (v.
pale), and other characters. Paraliparis eastmani is similar
to P. trunovi Andriashev from the Meteor Seamount and
the slope of SE Africa (Andriashev, 1986) in having two
dorsally located radials, a dark body and orobranchial cavity,
and especially in having two elongated lower pectoral-fin
rays that are slightly longer than the longest upper pectoral-
fin lobe rays. However, it differs distinctly in having a
shorter head (19.0% SL v. 22.7-23.0), darker colour
(uniformly solid black v. dark brown, unpigmented muscles
visible through the skin), and fully developed notch rays
(v. rudimentary). In addition, Vert. 69 (v. 63-66), D 64 (v.
57-59), A 57 (v. 51-53), and C 1+3/3+1 (v. 4/4).
Paraliparis gomoni n.sp.
Fig. 35
Material examined. Holotype NMV A7124, ripe S, 121
mm TL, 110 mm SL. FRV Soela, stn. So 3/88/8, 39°00.92'S
148°43.71'E, Tasmania, 100 km NE of Flinders Island,
1140-1160 m, 13 May 1988; radiograph NMV A; pectoral
girdle 801.
Diagnosis. Vert. 69, D c. 62, C 8, radials 3, round. Mouth
horizontal, terminal, teeth small, simple. Chin pores in a
pit, interspace equal to one pore diameter. Gill opening 0.8
eye, ventral end on horizontal with lower margin of eye. P
17+2+4, rudimentary rays absent, upper lobe long, 76 HL.
Head 19.3 SL, preanal 33.5, body low, bd 90.4 HL. Colour
brown with paler areas. Skin thin, peritoneum black.
Further description. Counts: D c. 62, Ac. 56, P 23, C 8 (4/
4), Vert. 69 (11+58), radials 3 (3+0), pc 5, gr 8, pores 2-6-
7-1. Ratios: HL 19.3, its width 10.6 (55) and depth 16.5
(86), bd 17.7 (90.4), bdA 15.9 (82), preD 22.0, preA 33.5,
ma 12.9, aAf 22.0, UPL 14.7 (76), LPL 12.8 (67), UPL 87.5,
NL —, E 4.7 (24.3), gs 3.8 (19.5), sn 6.9 (35.7), po 9.9 (51.4),
io 7.2 (34.3), uj 8.7 (45), lj 8.3 (43), so 3.8 (19.5), pc 3.8-5.5.
Head small, 5.2 in SL, compressed, depth 1.6 its width.
Dorsal contour straight, slanted anteriorly to vertical snout.
Snout highest point level with upper half of eye, deep, blunt,
not projecting, 1.5 eye. Subrostral fold deep, but not
covering upper lip entirely. Eye 4.1 in HL, almost touching
dorsal contour. Suborbital distance 0.8 eye. Pupil c. Vi eye.
Interorbital flat, 1.5 eye. Nostril small, level with upper
margin of pupil, with raised rim. Mouth terminal, horizontal,
cleft to below anterior margin of eye. Lower jaw included,
chin in ventral view rounded, in lateral view right-angled;
lower jaw shallow below oral cleft, honeycomb tissue
absent. Lips not wide. Teeth small, slightly prominent.
Upper jaw including about 20 rows of up to 8 teeth each,
diastema absent; in lower jaw about 24 rows of up to 8
teeth, narrow diastema present. Circumoral pores large,
whitish, contrasting with head colour, not in pits, n r n 2
widely spaced by a distance about equal to eye, chin pores
pm 7 close to eye at about first third of postorbital distance.
Chin pores in a distinct pit, oval, interspace about equal to
their diameter. Gill opening 0.8 eye, upper end above eye
level, lower about on level with lower margin of eye.
Opercular flap triangular with rounded ventral margin,
covering Vs length of gill opening, tip above level of eye
centre. Gill opening extending ventrally to base of upper
pectoral ray.
Upper pectoral lobe about level with mid-eye, not
reaching anal fin. Lower pectoral lobe at vertical through
anterior third of orbit. Both lobes long; P 17+2+4,
rudimentary rays absent. Radials 3 (3+0), round, scapula
384 Records of the Australian Museum (2001) Vol. 53
Figure 35. Paraliparis gomoni n.sp. A, holotype, NMV A7124, ripe 6, 121 mm TL, 110 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 801, cleared and stained right pectoral girdle.
with short helve, coracoid helve long. Pectoral membrane
on proximal side of upper lobe attached at about Vs length,
about Vs of notch and about Vs of lower lobe.
Body cigar-like, elongated, dorsal contour slightly
straighter than ventral contour; shallow, 5.7 in SL, not
humpbacked, maximum depth at anal-fin origin. Horizontal
midline through lower margin of eye. Vertebral column
straight anteriorly, last abdominal vertebra with a short
parapophysis without haemal spine. First caudal vertebra
with long haemal spine. First dorsal ray shortened, inserted
between vertebra 5 and 6, 2 free interneurals present
between vertebrae 3-4,4-5. Costal ridges absent. Epineural
ribs only on vertebrae 2-9, epipleural ribs on 2-5, thin, not
stout, about 2.5 vertebrae long. Anus at about second third
of postorbital length. Urogenital papilla small. Vertical fins
overlapping caudal about half. Gelatinous tissue poorly
developed. Skin thin, semitransparent, prickles absent.
Pyloric caeca unequal, gradually increasing in length.
Colour. Head and body brown with paler areas, belly
blackish, subrostral fold, lips, mouth, and tongue blackish
or dark grey; pectoral fin darker than body. Tooth plates
pale. Peritoneum black, stomach, pyloric caeca pale.
Branchial cavity dark grey, arches grey.
Distribution. Off NE Tasmania, at about 1150 m depth.
Etymology. Named after Martin F. Gomon, Senior Curator,
Ichthyology, Museum Victoria, Melbourne, Australia, for
his contributions to Australian ichthyology.
Comparative notes. A member of group Illb, P gomoni is
distinguished by radials 3+0, fenestrae absent, long pectoral-
fin lobes, pyloric caeca not only unequal, but progressively
increasing in length; colour brown, shallow body lacking
humpback, epineural ribs on vertebrae 2-9, epipleural ribs
on 2-5, and haemal spine absent on vertebra 11. Paraliparis
gomoni is most similar to P. infeliciter and Paraliparis sp.
1; it differs from the former in the absence of pectoral girdle
foramina (v. 1), shallower body with a terminal mouth (v.
sub terminal), lower jaw included (v. subterminal), chin
right-angled (v. rounded), Vert. 69 (v. 66), and other
characters. It differs from the latter in fenestra (0 v. 3), head
depth 86 (v. 77.5), body depth 90.4 (v. 116), pectoral rays
23 (v. 19-21) and other characters.
Paraliparis hobarti n.sp.
Fig. 36
Material examined. Holotype CSIRO H3170-01, $, 140
mm TL, 124 mm SL. Continental slope of Tasmania, no
other data; radiograph 681 C; pectoral girdle 792.
Diagnosis. Vert. 66, D 60, C 8, radials 2, round. Mouth
almost terminal, teeth simple, small. Subrostral fold absent.
Chin pores even with chin surface, slit-like. Eye large, 26%
HL, snout short. Ventral end of gill opening and uppermost
pectoral ray horizontal with lower margin of pupil. Last
preoperculo-mandibular pore very close to eye. Pectoral
fin long, 83% HL; P 20 (14+2+4), rudimentary notch rays
absent. Body deepest at anal-fin origin, bdA 102% HL. Head
18.5% SL, ma long, 14.5%, preanal length 35.5%. Colour
light brown, skin semitransparent, peritoneum black.
Further description. Counts: D 60, A 54, P 20, C 8 (4/4),
Vert. 66 (11+55), radials 2+0+0, gr 8, pc 6, pores 2-6-7-1.
Ratios: HL 18.5, its width 10.8 (58), and depth 16.1 (87),
bd 19.0 (102), bdA 19.0 (102), preD 21.7, preA 35.5, ma
14.5, aAf 21.8, UPL 15.3 (83), LPL 12.1 (65% HL, 79%
UPL), NL 3.2 (21% UPL), E 4.8 (26.0), gs 4.0 (21.7), sn
5.7 (30.8), po 8.4 (45), io 6.9 (37.0), so 2.5 (13.9), uj 8.9
(48), lj 8.1 (43), pc 2.4-4.8.
Head small, 5.4 in SL, deep at occiput, sloping anteriorly
to rounded snout. Head depth 1.5 its width. Eye large, not
close to dorsal contour of head, suborbital short, Vi eye.
Stein et al .: Australian liparids 385
Figure 36. Paraliparis hobarti n.sp. A, holotype, CSIRO H3170-01, $, 140 mm TL, 124 mm SL. B, ventral view
of mouth and upper jaw tooth plate. C, P 792, cleared and stained right pectoral girdle.
Pupil about half of eye diameter. Interorbital width 1.3 eye,
postorbital length short. Snout deeply rounded, short, 1.1
eye, not gelatinous, slightly projecting above upper jaw,
subrostral fold absent. Upper lip not wide. Nostril pore¬
like, level with lower margin of pupil. Mouth almost
terminal, horizontal, cleft reaching to below anterior margin
of pupil. Chin margin deep, almost right angled but with
rounded tip, gelatinous. Lower jaw subterminal, almost
equal to upper. Teeth simple, very small, not prominent, in
21-22 rows of up to 9 teeth anteriorly. Diastemae narrow,
almost absent. Lower jaw tapering anteriorly, chin edge
rounded. Circumoral pores with slightly thickened rims.
Chin pores slit-like, interspace (pigmented) less than their
diameter, level with chin surface; no skin fold anteriorly.
Preoperculo-mandibular pore series unusually close to
infraorbital pore series on cheek: pm 7 close to eye, distant
from it by Vi of postorbital space (usual state in Paraliparis
is Vi). Gill opening small, 0.8 eye, dorsal end level with
upper margin of eye, ventral end level with lower margin
of pupil. Gill opening not vertical, dorsal end anterior to
ventral end. Opercular flap triangular, tip rounded, covering
% of gill opening, level with upper margin of eye.
Dorsal pectoral ray on horizontal with lower margin of
pupil, ventralmost ray on vertical just behind posterior
margin of eye. P 20 (14+2+4). Skin on proximal surface of
pectoral fin attached to body at bases of upper lobe rays,
about % of notch ray length, and at about Vi of lower lobe
length. Pectoral upper lobe not quite reaching anal-fin origin,
comparatively long. Rudimentary rays absent. Radials 2+0+0,
round; fenestrae absent. Coracoid with long thin helve.
Body elliptic, not deep, 5.3 in SL, deepest at anal-fin
origin, dorsal contour broadly rounded anteriorly and
posteriorly from this point. Dorsal and ventral contours
similar. Horizontal midline touches lower margin of pupil.
First dorsal ray interneural between neural spines 5 and 6,
1 free interneural between neural spines 4 and 5. Anterior
dorsal rays embedded in gelatinous tissue. Posteriormost
two or more abdominal parapophyses joined to form short
haemal spines. Costal ridges weakly developed. Epineural
ribs on vertebrae 2-14, epipleural ribs on 10-11, not long
or stout, not more than 2.5 and 2 vertebrae long respectively.
Anus below about one-third of postorbital space, distance
from mandible to anus long. Vertical fins overlapping caudal
fin about half. Skin quite thin, semitransparent. Gelatinous
tissue moderately developed. Very small urinary papilla
present. Pyloric caeca thick, ends rounded. Ovarian eggs
up to 2.5 mm.
Colour. Head and body brown, not uniform but with whitish
areas from musculature barely visible through semitrans¬
parent skin; ventral surface of belly, lips, chin and pectoral
fins darker, uniformly blackish-brown. Mouth grey, tongue
slightly lighter, also grey. Branchial cavity and gill arches
dark grey. Pores, especially on lower jaw, contrasting
whitish. Peritoneum black, pyloric caeca and stomach pale.
Distribution. Continental slope of Tasmania.
Etymology. Named after the capital of Tasmania—Hobart.
Comparative notes. In group Ilia, distinguished by having
only two radials (unnotched), brown colour with whitish
areas, absence of a subrostral fold, the preoperculo-
mandibular sensory pore series unusually close to the
infraorbital series on the cheek, short snout and postorbital
head length, long upper pectoral-fin lobe, and other
characters.
386 Records of the Australian Museum (2001) Vol. 53
Figure 37. Paraliparis impariporus n.sp. A, holotype, CSIRO H3168-01, $, 173 mm TL, 162 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 734, cleared and stained right pectoral girdle. D, seventh vertebra.
Paraliparis impariporus n.sp.
Fig. 37
Material examined. Holotype CSIRO H3168-01, $, 173
mm TL, 162 mm SL. FRV Soela, stn. So 2/89/70,41°51.97'S
144°27.16'E, W coast of Tasmania, W of Trial Harbour,
1040-1050 m, 14 Mar. 1989; radiograph 682F; pectoral
girdle 734.
Diagnosis. Vert. 70, D 64, P 20-21 (15-16+2+3), C 9,
radials 3 (3+1?, see below), round. Mouth subterminal.
Teeth large, blunt. Chin pore single, unpaired. Gill opening
small, entirely above P base, reaching ventrally to level of
eye centre. HL 19.7% SL, about equal to its depth, preanal
long, 40%. Body deep, 132% HL, bdA 111%. Colour solid
black. Orobranchial cavity and peritoneum black, tongue
densely dotted.
Further description. Counts: D 64, A 56, P 20-21, C 9 (5/
4), Vert. 70 (11+59), radials 3 (3+1?, see below), pc 4, pores
2-6-7-1. Ratios: HL 19.7, its width c. 10.5 (53.5), and depth
15.5 (103), bdc. 20 (132), bdAc. 17 (111), preD 26.8, preA
c. 40, ma 14.8, aAf c. 23.4, UPL 14.0 (66), LPL 9.6 (69%
UPL), NL 1.7 (13% UPL), E 4.1 (25.8), gs 4.3 (25.6), sn
5.3 (35), io 5.7 (37), po 7.9 (53), uj 7.1 (47), pc 6-7.
Head small, about Vs SL, deep and compressed, depth
and length about equal, width more than half HL, 1.9 width.
Dorsal contour of head rounded; snout high, slightly
protruding. Interorbital space gently curved, broad, 1.4 eye
diameter. Eye moderately large, not touching the upper
contour of head; pupil very large, about % eye diameter.
Horizontal midline passing through eye centre. Nostril level
with lower margin of eye, its rim slightly raised. Mouth
subterminal, lower jaw almost equal to upper jaw. Maxilla
reaching slightly behind vertical through mid-eye, oral cleft
almost reaching centre of eye. Teeth quite large, stout, blunt
canines, closely set in about 30-35 oblique rows of 6-8
teeth each anteriorly, forming moderately wide band in each
jaw; narrow diastema present between teeth of each side in
upper and lower jaws. Subrostral fold deep, almost entirely
covering upper lip anteriorly. Lower jaw partly skinned,
honeycomb tissue present on chin; jaw below end of mouth
cleft deep. Circumoral pores not large; round, contoured.
Chin pore single, unpaired, round, similar in size to other
mandibular pores. Free neuromasts on head and body
apparently absent. Gill opening vertical, small, about equal
to eye, above P fin base, relatively high on side of head,
reaching ventrally to level of mid-pupil and in front of first
pectoral ray. Opercular lobe small but distinct, triangular,
pointed; its tip level with upper margin of eye. Gill rakers 7.
Uppermost pectoral-fin ray about level with middle of
pupil. Upper pectoral lobe not nearly reaching anal-fin
origin. P 20-21: (16+2+3) (L), (15+2+3) (R). Pectoral-fin
notch distinct, moderately deep, of 2-3 more widely spaced
rays; rudimentary rays absent. Lower lobe short, of 3 rays,
the longest about % upper lobe length. Pectoral radials
round, 3 (3+0), but probably 4 (3+1); at normal location of
lowest radial only an empty pit present. Two rudimentary
fenestrae present at upper and lower margins of uppermost
radial. Helve of scapula well developed; coracoid with
elongated helve with two lateral ribs.
Body tapering gradually towards tail, very deep, greatest
depth at about the level of posterior part of pectoral fin,
much greater than HL. Trunk depth similar, predorsal long.
Anterior dorsal rays short, embedded in gelatinous tissue.
Interneurals of first and second dorsal rays between 5th and
Stein et al.: Australian liparids 387
6th spines; no ray between 6 and 7th neural spines. One
free rayless interneural discernible before anteriormost ray.
Parapophyses of last (11th) abdominal vertebra grown
together, forming rather short haemal spine. Epineural ribs
at 2nd-10th vertebrae, epipleural ribs at 3rd-17th; all thin,
2.5-3 vertebrae long. Caudal fin of 9 (5/4) rays, procurrent
rays absent. Degree of overlap with vertical fins unknown
owing to damage. Anus below 2 A of postorbital space,
mandible to anus long. Skin thick, prickles absent;
subcutaneous gelatinous layer moderately developed.
Pyloric caeca digitate, of similar length.
Colour. Body colour solid black, oral cavity densely black,
tongue densely dotted, tooth plates pale, skin fold behind
tooth plates inside mouth black. Inner surface of subrostral
fold densely dotted, almost black. Lower lip dark grey. Gill
arches light grey. Branchial cavity and peritoneum black.
Stomach and pyloric caeca pale.
Distribution and biology. West coast of Tasmania,
Australia, at a depth of 1040-1050 m. Collected by a
demersal trawl, P. impariporus is probably benthic or
benthopelagic. The holotype is a female, 173 mm TL, that
appears to have spawned shortly before capture (early
March): rather than being filled with eggs to form a solid
mass, the cavity of the ovary is hollow. The largest eggs are
1.7 mm in diameter, and unripe.
Etymology. Impariporus from the Latin impar — unequal
and porus —pore, to note the single (rather than paired)
terminal mandibular pore.
Comparative notes. Paraliparis impariporus belongs to
group Ilia. It differs in having a single chin pore, a very
deep and compressed head and body with long predorsal
and preanal lengths, gill opening high on the side and
reaching ventrally to the eye centre, the subrostral fold inside
densely dotted, and other characters. It is similar to P.
monoporus Andriashev & Neelov (1979) from the Antarctic
in having a single, unpaired symphyseal mandibular pore,
but distinctly differs in being entirely black including the
orobranchial cavity (v. a uniformly pale body and
orobranchial cavity, orange-red in life), longer gill opening
(4.3% SLv. 2.1-2.2%), andradials 2+1 (v. 3+1). Differences
in counts are not great (Vert. 70 v. 72-74, A 56 v. 58-62, P
20-21 v. 22-25).
Paraliparis infeliciter n.sp.
Pig. 38
Material examined. Holotype CSIRO H3169-01, ripe $,
170 mm TL, 153 mm SL. PRV Soela, stn. So 1/88/48,
37°37.9'S 139°00.60'E, South Australia, W of Cape Banks,
1070-1090 m, 30 Jan. 1988; radiograph 682 D; pectoral
girdle 798. Specimen from Hoplostethus atlanticus stomach.
Diagnosis. Vert. 66, D 60, C 8, radials 3+0, uppermost
notched. Interneural of first dorsal ray between neural spines
6 and 7. Mouth subterminal, lower jaw subterminal. Teeth
simple, small. Chin pores in a pit on lower surface of chin,
space between unpigmented, equalling their diameter.
Ventral end of gill opening and dorsal pectoral ray level
with lower margin of pupil. P 17+2+3, rudimentary notch
rays absent. Head 19.1% SL, compressed (width 55% HL),
and depressed (depth 72% HL); preanal distance 34%,
mandible to anus short, 10.5% SL. Colour blackish- brown,
paler laterally; peritoneum black.
Further description. Counts: D 60, A 55, P 22, C 8 (4/4),
Vert. 66 (11+55), radials 3+0, gr 8, pc 6, pores 2-6-7-1.
Ratios: HL 19.1, its width 10.5 (55), and depth 13.7 (72),
A
B
Figure 38. Paraliparis infeliciter n.sp. A, holotype, CSIRO H3169-01, ripe $, 170 mm TL, 153 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 798, cleared and stained right pectoral girdle.
388 Records of the Australian Museum (2001) Vol. 53
bd 20.9 (109), bdA 15.4 (80), preD 23.2, preA 34.0, ma
10.5, aAf 21.6, UPL 11.8 (62), LPL 8.5 (44.5% HL, 72%
UPL), NL 2.9 (25% UPL), E 4.7 (24.7), gs 3.3 (17.5), sn
6.5 (34.2), po 8.6 (45.2), io 7.4 (38.7), so 2.9 (15.1), uj 8.5
(45.4), lj 8.0 (42), pc 4.3.
Head small, 5.2 in SL, compressed, low, depth 1.3 its
width. Eye not entering dorsal contour of head, suborbital
space and pupil 2 A eye. Interorbital space rounded, 1.6 eye.
Snout deep, rounded, 1.4 eye, slightly projecting anterior
to upper jaw, highest point level with upper margin of pupil.
Subrostral fold well developed, deep, not covering upper
lip. Nostril level with lower half of eye. Mouth subterminal,
horizontal, cleft reaching to below anterior third of eye.
Lower jaw subterminal, almost included. Teeth simple,
small, in upper jaw in 22-24 rows of up to 9-10 teeth
anteriorly. Diastemae of both jaws narrow. Chin in ventral
view rounded; skinned, thin honeycomb tissue visible; lower
jaw shallow below posterior of oral cleft. Circumoral pores
small, chin pores in a small unpigmented pit, rims thickened,
interspace equals their diameter; on lower surface of chin.
Postorbital head length short. Gill opening small, vertical,
ventral end on horizontal with lower half of pupil, dorsal
slightly above eye. Opercular flap triangular, covering lower
2 A of gill opening, lower margin rounded, tip level with
upper margin of eye.
Uppermost pectoral ray level with lower margin of pupil.
P 17+2+3, rudimentary rays absent. Upper lobe short, not
reaching anal-fin origin; lower lobe short, lowest ray below
posterior margin of eye. Fin membrane missing. Radials
3+0, lowest small. R1 notched with a small ventral fenestra
in notch; R2 with a tiny rudimentary ventral notch. Coracoid
with unusually strong, long helve.
Body deepest at dorsal-fin origin; body shape unclear
owing to damage. Dorsal contour broadly rounded
anteriorly, evenly tapering posteriorly. Horizontal midline
passes through eye centre. Anterior of dorsal fin possibly
damaged; interneural of first dorsal ray apparently between
neural spines 6 and 7; 3 free interneurals present anteriorly
between neural spines 3-6. Anterior dorsal rays embedded
in gelatinous tissue. Parapophyses of abdominal vertebrae
distinctly separate. Costal ridges absent. Epineural ribs on
vertebrae 2-8, anteriorly not long, not stout, length less than
3 vertebrae; epipleurals not clearly visible on radiograph. Anus
below anterior third of postorbital space, mandible-anus length
short. Genital papilla absent. Skin thin, translucent, prickles
absent. Gelatinous tissue moderately developed. Ovarian
eggs to 2.9 mm. Pyloric caeca similar, pointed.
Colour. Light blackish-brown, skin on sides of trunk paler,
finely dotted; anal region blackish. Subrostral fold brightly
black-dotted. Lips slightly paler than head, grey. Oral cavity
black-dotted: tongue and palatal skin fold behind upper
tooth plates grey, soft palate itself paler. Branchial cavity
brown, gill arches grey. Peritoneum black, pyloric caeca
and stomach pale.
Distribution. Off eastern South Australia, 1070-1090 m.
Etymology. Infeliciter from the Latin “bad luck”; thus
named because the specimen was eaten by an orange roughy
(Hoplostethus atlanticus).
Comparative notes. A member of group Illb, P infeliciter
is distinguished by its comparatively light brown colour,
thick layer of honeycomb tissue on the chin, chin pores
with an unpigmented interspace, radials 3+0, notched R1
and R2 with one fenestra, coracoid with an unusually stout
and long helve, nephrohaemal canal absent, triangular
opercular flap, and rounded snout with a short oral cleft. It
is most similar to P. gomoni and Paraliparis sp. 1, but differs
from both in having a subterminal lower jaw (v. included).
See descriptions of both species for specific differences.
Paraliparis labiatus n.sp.
Fig. 39
Material examined. Holotype CSIRO H749-02, ?, 166
mm TL, 150 mm SL. FRV Soela, stn. So 3/86/32,44°45.8’S
144°24.8'E, W coast of Tasmania W of Granville Harbour,
1000-992 m, 16 May 1986; radiograph 682 B; pectoral
girdle 790.
Diagnosis. Vert. 66, D 59, C 8, radials round. Mouth
terminal, teeth small. Lateral profile of chin right-angled;
chin with thick gelatinous layer, pores protected by anterior
skin fold. Upper lip wide posteriorly, entirely covering lower
lip fold. Eye comparatively small, 21% HL, interorbital
broad, 44%. Lower end of gill opening and uppermost
pectoral ray level with lower margin of eye. P 16+2+3,
rudimentary rays absent. HL 19.3% SL, preA 35%. Vertical
fins overlapping caudal fin slightly more than half. Females
with prominent genital papilla. Colour dark blackish-brown,
peritoneum black, tongue grey-dotted.
Further description. Counts: D 59, A 53, P 21, Vert. 66
(11+55), C 8 (4/4), radials 4 (3+1), pc 5, gr 7, pores 2-6-7-
1. Ratios: HL 19.3, its width 12.7 (66), and depth 15.3 (79),
bd 20.7 (107), bdA 20 (103), preD 24.7, preA 34.7, ma
13.7, aAf 22.0, UPL 13.3 (69), LPL 10.7 (55% HL, 80%
LPL), NL c. 2.0 (15% UPL), E 4.1 (21.0), sn 6.9 (35.9), gs
4.3 (22.4), io 8.5 (44.1), po 10.0 (52), so 3.2 (16.6), uj 8.3
(43), lj 7.7 (40), pc 6.6.
Head small, 5.2 in SL, its dorsal contour rounded
anteriorly; wide, its depth 1.2 its width. Eye small, 4.8 in
HL, upper contour clearly below dorsal profile of head,
suborbital space about % eye. Pupil half eye diameter.
Interorbital space gently curved, broad, 2.1 eye. Snout deep,
bluntly rounded, anterior profile vertical, highest point level
with lower margin of pupil; not projecting beyond upper
lip; its length 1.7 eye. Subrostral fold present, deep
anteriorly and covering upper lip almost entirely; shallow
posteriorly where upper lip is wide and prominent. Nostril
short, tubular, level with lower half of pupil. Mouth
horizontal, terminal, cleft reaching to below anterior margin
of eye. Lower jaw terminal, almost equal to upper. Upper
lip wider laterally than anteriorly, entirely covering lower
lip fold. Teeth very small, not prominent, in about 25 oblique
rows of up to 9 teeth each anteriorly; lower jaw tooth plates
damaged. Diastema of upper jaw narrow, wider than that
of lower jaw. Circumoral pores small; chin pores closely
set, interspace equal to their diameter; anterior skin fold
vertical, high; in lateral view, chin appears right -angled. In
ventral view, chin blunt; skin partially missing, a peculiar
honeycomb structure of subcutaneous tissue clearly visible.
Lower jaw shallow below posterior of mouth cleft. Gill
Stein et al .: Australian liparids 389
Figure 39. Paraliparis labiatus n.sp. A, holotype, CSIRO H749-02, $, 166 mm TL, 150 mm SL. B, ventral view
of mouth, lower jaw, and genital papilla. C, P 790, cleared and stained right pectoral girdle.
opening small, about equal to eye, vertical, completely
above pectoral base; lower end level with lower margin of
eye. Opercular flap ear-shaped, tip on horizontal through
upper margin of eye.
Uppermost pectoral ray level with lower margin of eye.
Pectoral fin 16+2+3, rudimentary notch rays absent. Upper
lobe not reaching anal-fin origin, lower lobe short. Lowest
ray below posterior margin of eye. Pectoral-fin membrane
missing. Pectoral girdle with 4 (3+1) radials; all round, of
similar sizes, distance between R3 and R4 short. Fenestrae
absent. Scapula and coracoid each with a long helve.
Body elliptic, deep, 4.8 in SL; greatest depth close to
anal-fin origin. Upper and lower body profiles similar.
Horizontal midline touching lower margin of eye. Anterior
half of dorsal-fin rays embedded in gelatinous tissue.
Interneural of the first (rudimentary) dorsal ray between
neural spines 5 and 6; one free interneural present anteriorly.
Caudal-fin rays 4/4. Short haemal spines present on
vertebrae 9-11. Epineural ribs present on vertebrae 2-18,
thin and short, length not exceeding 2 vertebrae. Epipleural
ribs not visible in radiograph; if present, very slim. Vertical
fins overlapping caudal fin slightly more than half. Anus
below posterior third of postorbital space. Anal-genital area
unusual: prominently swollen, a short, thick cone-shaped
genital papilla present behind oviduct opening, its length
about equal to half of eye. Skin on body entirely missing;
skin on head thin, translucent. Subcutaneous gelatinous
tissue well developed. Pyloric caeca similar. Ovaries full
of eggs at different stages of maturity, not ripe, largest about
1 mm in diameter.
Colour. Remains of skin on head dark blackish-brown.
Inner surface of subrostral fold bright black-dotted, similar
to head colour; lips and chin margin lighter, grey. Mouth
dark grey, tongue lighter, grey-dotted, tooth plates pale.
Branchial cavity dark brown, gill arches grey, black-dotted.
Peritoneum black, anal area and genital papilla black.
Pyloric caeca pale.
Distribution. West coast of Tasmania at 1000-992 m.
Etymology. Labiatus from the Latin, having large lips.
Comparative notes. Paraliparis labiatus is in group IIIc.
It is distinguished by its chin shape and structure (right-
angled in lateral profile with honeycomb tissue), upper lip
covered only anteriorly by the posteriorly wider subrostral
fold, broad head, small eye, broad interorbital, unusual
female genital papilla, pectoral girdle structure with 4 radials
of similar size (distance between radials 3 and 4 short) and
absence of fenestrae. It is most similar to P. brunneus but
differs (in addition to the above characters) in the absence
of fenestra in the pectoral girdle (v. 3 present), the shape of
the opercular flap, and smaller eye (21% SL v. 24%). It is
also similar to P. anthracinus, but differs by the gill opening
reaching ventrally to the uppermost pectoral-fin ray (v. to
the second), subrostral fold (v. absent), and blackish-brown
colour (v. coal black).
Paraliparis lasti n.sp.
Fig. 40
Material examined. Holotype CSIRO T982-02, 3 , 205
mm TL, 185 mm SL. FV Petuna Endeavour , stn. PE 4/10,
33°45.5'S 129°37.5'E, South Australia, Great Australian
Bight, 1152-1000 m, 5 June 1983; radiograph 684 C;
pectoral girdle 779.
Diagnosis. Vert. 70, D 63, C 8, P 22 (15 + 3+4),
rudimentary rays absent. Radials 3, round. Mouth
390 Records of the Australian Museum (2001) Vol. 53
Figure 40. Paraliparis lasti n.sp. A, holotype, CSIRO T982-02, 6, 205 mm TL, 185 mm SL. B, ventral view of
mouth and upper jaw tooth plate. C, P 779, cleared and stained right pectoral girdle.
horizontal, subterminal, lower jaw included; teeth large,
stout; chin pores paired, small, not in pits. Uppermost
pectoral ray horizontal with eye centre. HL 19.7% SL,
preA 37% SL. Mandible to anus short, 11.4% SL. Skin
semitransparent, reddish-brown, peritoneum brownish-
black.
Further description. Counts: D 63 (1 rudimentary+62), A
57, P 22, C 8 (4/4), Vert. 70 (12+58), radials 3 (3+0), pc 7,
gr 9, pores 2-6-7-1. Ratios: HL 19.7, its width 11.4 (57.5),
and depth 14.9 (75), bd 21.1 (107), bdA 17.8 (90), preD
26.0, preA 36.8, ma 11.4, aAf 23.1, UPL 14.9 (75), LPL
13.5 (90% UPL), NL 4.1 (27% UPL), sn 6.7 (33.1), E 4.6
(23.6), gs 4.3 (22.0), io 7.3 (37.0), po 12.2 (49.3), so 2.8
(14.2), uj 10.3 (52), lj 8.6 (43.8), pc 5.4.
Head not large, 5 in SL, deep at occiput; dorsal contour
significantly sloping anteroventrally. Head quite com¬
pressed, its width more than half its length, its depth 1.3
its width. Snout quite large, bluntly rounded, 1.4 eye,
projecting above upper jaw, its highest point horizontal
with eye centre. Subrostral skin fold developed, covering
upper half of upper lip. Eye 4.2 in HL, not entering dorsal
contour of head. Pupil 3 A eye diameter. Interorbital space
1.6 eye. Suborbital distance almost % eye. Nostril large,
pore-like, level with lower half of pupil. Mouth
horizontal, subterminal, its cleft reaching to below centre
of eye. Lower jaw included; in ventral view, if upper lip
pushed aside upper tooth plates visible. Upper jaw long,
chin tapering in ventral view, symphysis slanted but not
right-angled in lateral view, lips not wide. Lower jaw
below angle of mouth cleft quite deep. Teeth simple,
large, stout, slightly blunt, in 23-25 quite regular oblique
rows, 8-10 teeth per row anteriorly. Diastema of upper
jaw wider than at lower jaw. Circumoral pores small, not
in pits. Chin pores spaced at 1.5 times their diameter,
interspace pigmented, not in a pit and without skin fold
anteriorly. Gill opening length 0.9 eye diameter, entirely
above pectoral base, vertical, dorsal end clearly above
level of eye upper margin, lower end horizontal with eye
centre. Opercular flap small, triangular, with notched
upper rim, covering lower half of gill opening, tip level
with upper margin of eye.
Uppermost pectoral-fin ray level with eye centre and
below opercular tip, lowermost ray inserted below
posterior margin of eye. Upper and lower pectoral lobes
long; upper lobe not reaching anal-fin origin, 22
(15+3+4). Notch ray length about l A UPL, lower lobe
length not quite equal to UPL. Skin on proximal surface
of upper lobe attached at ray bases, notch and lower lobe
unknown. Basal cartilaginous lamina of pectoral girdle
with 3 (3+0) round radials in upper half of basal lamina,
R3 smallest. Lenestrae absent. Helve of scapula stout.
Coracoid with long helve with ventral lateral rib.
Body deep, humpbacked, greatest depth at dorsal-fin
origin, depth at A origin slightly less. Predorsal length
long. Dorsal contour much more curved than ventral
contour. Horizontal midline of body goes through eye
centre. Lirst dorsal-fin ray rudimentary, its interneural
between neural spines 5 and 6; one free interneural
between neural spines 4 and 5. Haemal spine of the last
(12th) abdominal vertebra short. Epineural ribs on
vertebrae 2-15, epipleurals on 3-15, thin and not long,
lengths not more than 2-2.5 vertebrae. Ribs producing
lateral keel-like body ridges above first half of pectoral
fins, but not prominent. Anus below the middle of
postorbital space, mandible-anus distance short. Vertical
fins overlapping caudal at l A of its length. Skin thin,
semitransparent. Subcutaneous gelatinous tissue slightly
developed. Pyloric caeca similar in size.
Colour. Uniform bright reddish-brown, anteriorly slightly
lighter. Snout and chin brown, lips paler, greyish-brown.
Stein et al .: Australian liparids 391
Figure 41. Paraliparis obtusirostris n.sp. A, holotype, NMV A7123, ripe $, TL unknown, 137 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 800, cleared and stained right pectoral girdle.
Mouth light grey, tongue grey-dotted. Branchial cavity
blackish, gill arches pale. Anal region blackish. Peritoneum
brownish-black, stomach and pyloric caeca pale.
Distribution. Great Australian Bight, depth 1152-1000 m.
Etymology. The new species is named in honour of Peter
R. Last of the CSIRO Division of Marine Research,
Hobart, Tasmania, whose studies of Australian fishes
have added fundamentally to knowledge of the deep-sea
fauna of that region.
Comparative notes. Paraliparis lasti is in group Ilia,
and is distinguished by its thin, semitransparent, reddish-
brown skin, dorsally-located gill opening (lower end level
with eye centre), long predorsal length, short mandible-
anus distance, and long pectoral-fin lobes. It is most
similar to P. piceus, see that species for comparison. It is
also similar to P. dewitti n.sp. but differs in having 3
(3+0) (v. 4 (3+1) radials), teeth large and stout (v. small),
symphyseal mandibular pores not embedded (v. in an oval
depression), chin angled (v. bluntly rounded), skin bright,
reddish-brown (v. nut-brown), pyloric caeca pale (v.
grey), snout 1.4 eye (v. 1.8), interorbital width 1.6 eye
(v. 2), and vertebrae 70 (v. 65).
Paraliparis obtusirostris n.sp.
Fig. 41
Material examined. Holotype NMV A7123, ripe $, TL
unknown, 137 mm SL. FRV Soela, stn. So 3/88/10,
38°57.09'S 148°41.95'E, off Tasmania, 100 km NE of
Flinders I., 1270-1290 m, 13 May 1988; radiograph NMV
B; pectoral girdle 800.
Diagnosis. Vert. 66, D 62, C 9, radials 2, lowest notched.
First dorsal ray interneural between neural spines 4-5.
Mouth terminal, lower jaw subterminal. Teeth simple, small.
Chin pores in small unpigmented pit, almost touching.
Lower end of gill opening and uppermost pectoral ray level
with lower half of pupil. P 14+3+4 to 14+2+3, rudimentary
rays absent. Head 18.6% SL, preanal 33%. Body light brown
with variable pattern, peritoneum black.
Further description. Counts: D 62, A 55, P19-21, C9 (1+4/
4), Vert. 66 (11+55), radials 2+0+0, fenestrae 2, pc
unknown, gr 8, pores 2-6-7-1. Ratios: HL 18.6, its width
11.3 (61), and depth 16.4 (88), bd 20.4 (110), bdA 17.2
(92), preD 22.8, preA 33.3, ma 12.3, aAf 21.5, UPL 12.8
(69), LPL 9.5 (50.1% HL, 74% UPL), NL 1.8 (14.2% UPL),
E 4.7 (25.5), gs 4.2 (22.7), sn 7.3 (39.2), po 9.9 (53), io 6.6
(35.6), so 3.6 (19.6), uj 8.2 (44), lj 7.9 (42.4).
Head small, 5.3 in SL, deep at occiput, dorsal contour
sloping straight anteriorly to blunt snout. Head not much
compressed, its width 0.6 its length, depth 1.5 width. Eye
large, 3.9 in HL, almost entering dorsal contour of head,
suborbital almost equal to eye. Pupil about Vi eye.
Interorbital 1.4 eye, suborbital distance about 0.8 eye. Snout
deep, blunt, 1.5 eye, not projecting anterior to upper jaw,
its highest point horizontal with upper margin of pupil.
Subrostral fold not deep, not entirely covering upper lip.
Upper lip not wide. Nostril almost on level with eye centre.
Mouth terminal, horizontal, its cleft reaching to below
anterior half of pupil. Lower jaw subterminal, almost equal
to upper. Teeth simple, small, barely projecting; in upper
jaw in 21-22 rows of 9 teeth each anteriorly. Diastemae
absent. Chin rounded in ventral view, tip right-angled in
lateral view, shallow below posterior of mouth cleft.
Circumoral pores not in pits, contoured. Nasal pores widely
spaced at about % eye diameter. Chin pores in a shallow
unpigmented depression almost touching. Gill opening not
392 Records of the Australian Museum (2001) Vol. 53
short, 0.9 eye, lower end level with lower half of pupil.
Opercular flap triangular, covering Vs of gill opening, its
tip rounded, level with upper margin of eye.
Uppermost pectoral ray on horizontal with lower half of
pupil. Base of lowermost ray below first third of postorbital
space. Pectoral rays not reaching anal-fin origin. Pectoral
fin 14+2+3 (L), 14+3+4 (R), rudimentary rays absent. Skin
of pectoral-fin proximal side attached at about Vs upper lobe
length, 3 A notch length, Vs lower lobe length. Radials 2+0+0,
R1 round, R2 notched above and below, two fenestrae
present. Coracoid with long thin helve.
Body humpbacked, deepest at dorsal-fin origin, 4.9 in
SL. Horizontal midline passes through eye centre. Preanal
length 33% SL. First dorsal ray interneural between neural
spines 4 and 5, first and second dorsal rays shortened, one
free interneural present anteriorly between neural spines
3-4. Anterior dorsal rays embedded in gelatinous tissue.
Parapophyses of abdominal vertebrae unjoined. Costal
ridges absent. Epineural ribs on vertebrae 2-16, epipleural
ribs on 3-18, anterior ribs not long, not stout, shorter than
2.5 vertebrae. Anus below posterior Vs of postorbital space.
Caudal fin with one dorsal procurrent ray. Skin quite thin,
slightly transparent, prickles absent. Gelatinous tissue
moderately developed. Genital papilla absent. Eggs in
ovaries of different size, the largest 2.4 mm. Pyloric caeca
not studied.
Colour. Uniformly light brown, with variable markings,
pectoral fins and anal area blackish. Subrostral fold and
upper lip blackish, slightly darker than head. Mouth grey,
tongue pale, black-dotted, tooth plates pale. Branchial cavity
black, gill arches grey. Peritoneum black. Pyloric caeca and
stomach pale.
Distribution. Coast of NE Tasmania, 1270-1290 m.
Etymology. Obtusirostris from the Latin, meaning “with
blunt snout.”
Comparative notes. Paraliparis obtusirostris is in group
Illb, and is distinguished by its brown colour, blunt deep
snout, presence of a caudal procurrent ray, two shortened
anterior dorsal-fin rays, and unjoined parapophyses of all
abdominal vertebrae. Most similar to Paraliparis sp. 2, it
differs in having 2 pectoral girdle fenestrae (v. none), a well-
developed subrostral fold (v. absent), absence of costal keels
(v. present), and light brown colour (v. black).
Paraliparis piceus n.sp.
Fig. 42
Material examined. Holotype CSIRO H805-04, ripe S ,
167 mm TL, 149 mm SL. FRV Soela, stn. So 3/86/41,
41°51.25'S 144°23.FE, W coast of Tasmania, W of Trial
Harbour, 1384-1416 m, 18 May 1986; radiograph 682 C;
pectoral girdle 796.
Diagnosis. Vert. 66, D 59, C 8, 3 radials, upper dorsally
notched. Mouth terminal, teeth simple, small; chin pores
not in a pit or other depression, interspace equals their
diameter. Gill opening ventral end and dorsal pectoral ray
on horizontal through eye centre. Opercular flap dorsally
rounded, tip of operculum dorsally directed. P 16+3+3,
rudimentary rays absent. HL 18% SL, preanal length 34%.
Body and head dark blackish-brown, mouth and tongue
grey, peritoneum black.
Further description. Counts: D 59, A 55, P 22, C 8 (4/4),
Vert. 66 (11+55), radials 3+0, one fenestra, gr 7, pc 5, pores
2-6-7-1. Ratios: HL 18.1, its width 11.8 (65), and depth
16.8 (93), bd 21.5 (119), bdA 19.7 (108), preD 21.1, preA
34.2, ma 12.6, aAf 21.8, UPL 14.1 (78), LPL 12.1 (67%
HL, 86% UPL), NL 3.4 (24% UPL), E 4.3 (23.7), gs 3.4
(18.5), sn 6.0 (33), io 6.7 (37), po 9.5 (52.6), so 3.4 (18.5),
u j 8.1 (44.4), lj 8.1 (44.4), pc 6.7.
Head small, 5.5 in HL, slightly compressed, depth 1.4
its width. Dorsal contour slopes significantly from deep
occiput to vertical snout. Eye large, 4.2 in head, close to
dorsal contour of head; suborbital space about 0.8 eye. Pupil
about half eye. Interorbital space slightly rounded, 1.6 eye.
Snout 1.4 times eye, deep, bluntly rounded, not projecting
anteriorly beyond jaw, its highest point horizontal with eye
centre. Subrostral fold present, much deeper anteriorly than
laterally, not covering upper lip posteriorly. Lips not wide.
Nostril pore-like, level with lower margin of pupil. Mouth
terminal, horizontal, cleft reaching to below anterior margin
of eye. Lower jaw subterminal, slightly shorter than upper.
Teeth simple, small, slightly prominent, blunt, in 23-24 rows
of up to 7-8 teeth anteriorly. Diastema of upper jaw narrow,
but wider than that in lower jaw. Chin right-angled in lateral
view, symphyseal knob large, vertical, clearly seen on
radiograph. Lower jaw below angle of mouth cleft shallow.
Circumoral pores small. Chin pores closely set, interspace
equal to their diameter, not in a pit or other depression;
interspace pigmented. Gill opening short, 0.8 eye diameter,
its ventral end level with eye centre. Opercular flap rounded
dorsally; not vertical (dorsal end in front of vertical of
ventral end), tip of operculum directed dorsally, about level
with upper margin of eye.
Uppermost pectoral ray on horizontal through eye centre.
Upper pectoral lobe long. Lowermost ray insertion below
first quarter of postorbital space. P 16+3+3, rudimentary
rays absent. Skin on proximal surface of upper pectoral-fin
lobe attached at base, in notch attached almost at ray tips,
in lower lobe at about l A lobe length. Radials 3+0, R1 with
a small dorsal notch, small fenestra present. Coracoid with
long thin helve.
Body leaf-like, deep and compressed, maximum depth
at anal-fin origin, 4.7 in SL. Dorsal contour very rounded,
ventral contour almost straight. Horizontal midline through
lower half of eye. First dorsal ray interneural between neural
spines 5 and 6; one free interneural present anteriorly.
Parapophyses of anterior abdominal vertebrae separate, last
three joined. Nephrohaemal canal present on vertebrae 9-
11, short. Epineural ribs on vertebrae 3-13, thin and short,
lengths about equal to two vertebrae; epipleural ribs not
clearly visible on radiograph. Costal ridge absent. Anus
below anterior third of postorbital space, small urogenital
papilla present. Vertical fins overlap anterior half of caudal
fin. Skin without prickles, opaque. Gelatinous tissue
moderately developed. Pyloric caeca of similar lengths.
Colour. Body, head, lips, inner surface of subrostral fold,
and chin uniformly dark blackish-brown. Mouth dark grey,
tongue grey, tooth plates pale. Pores pale, clearly
contrasting. Branchial cavity dark grey, gill arches lighter,
also grey. Urogenital papilla black. Peritoneum black,
stomach and pyloric caeca pale.
Stein et al .: Australian liparids 393
Figure 42. Paraliparis piceus n.sp. A, holotype, CSIRO H805-04, ripe S, 167 mm TL, 149 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 796, cleared and stained right pectoral girdle.
Distribution. West coast of Tasmania, 1384-1416 m.
Etymology. Piceus from the Latin, blackish-brown, tar-
coloured.
Comparative notes. Paraliparis piceus is in group Ilia. It
is distinguished by its leaf-like, deep and compressed body
and head, dorsally rounded opercular flap, blunt vertical
snout, long upper pectoral lobe, radials 3+0 with R1 dorsally
notched, uniform dark tar-colour, absence of rudimentary
rays and costal keel, and short nephrohaemal canal. It is
most similar to P. lasti but differs in tooth size (small v.
large), terminal mouth (v. sub terminal), Vert. 66 (v. 70), D
59 (v. 63), oral cleft reaching anterior margin of eye (v.
centre of eye), having one fenestra (v. absent), deeper body
(maximum 119 v. 107 HL, bdA 108 v. 91), and colour (dark
blackish v. light brown with distinct reddish tint).
Paraliparis plagiostomus n.sp.
Fig. 43
Material examined. Holotype CSIRO T488-02, 3 ,148 mm
TL, 137 mm SL. F \ Petuna Endeavour, stn. PE 6/8,42°19.5’S
144°42'E, W coast of Tasmania, W of Cape Sorell, 993-987
m, 9 July 1983; radiograph 680 B; pectoral girdle 782.
Diagnosis. Vert. 65, D 59, C 8. Radials 3, round. Jaws long,
mouth inferior, teeth tiny, tooth plates appear smooth. Pores
with distinctly contoured rims. Chin pores one diameter
apart. Snout slanted, protruding, subrostral fold deep and
entirely covering upper lip. Interorbital narrow. P 20
(15+2+3), upper ray horizontal with upper part of eye.
Upper pectoral lobe short, 63% HL, lower lobe short. Gill
opening equal to eye, dorsolaterally located, completely
above horizontal through upper margin of eye. HL 18.7%
SL, mandible-anus short, 10.9% SL. Uniformly brownish-
black, mouth and tooth plates dark grey, tongue black-
dotted, gill arches dark grey, peritoneum black.
Further description. Counts: D 59, A 54, P 20, C 8 (4/4),
Vert. 65 (11+54), radials 3 (3+0), pc 5, gr 8, pores 2-6-7-1.
Ratios: HL 18.7, its width 10.9 (57.8), and depth 16.4 (88),
bd 19.2 (103), bdA 17.2 (91), preD 21.0, preA 30.1, ma
10.9, aAf 19.1, UPL 11.7 (63), LPL 8.8 (75% UPL), NL c.
22% UPL, E 4.2 (22.7), gs 3.9 (20.7), sn 6.6 (35.5), io 7.7
(30.1), po 9.9 (53.1), so 2.9 (15.6), uj 10.2 (54.7), lj 8.9 (47.7).
Head small, 5.4 in SL, deep at occiput, dorsal contour
sloping rapidly to snout tip; very compressed, its width
slightly more than its length. Snout deep, rounded, large,
1.6 eye; greatly protruding beyond upper jaw, gelatinous.
Nostril short, tube-like, with stout rim; horizontal with lower
half of eye. Subrostral fold deep, entirely covering upper
lip. Eye 4.4 in head, upper contour not touching dorsal
profile. Interorbital narrow, 1.8 eye; suborbital 0.7 eye.
Mouth inferior, its cleft reaching to below anterior margin
of pupil. Upper and lower jaws large. Lower jaw subterminal
(shorter than upper jaw, but not included); in ventral view,
lower surface of snout and pores of upper jaw visible. Chin
widely rounded, gelatinous. Teeth simple, tiny; tooth plates
look smooth, in 25-27 regular rows, anteriorly of 7-9 teeth
each. Diastema absent in upper jaw, narrow in lower jaw.
Circumoral pores distinctly contoured, not in pits but on
surface; chin pore interspace equalling their diameter, in a
very shallow depression but not in a pit, anterior skin fold
absent. Gill opening 0.9 eye diameter, vertical, dorsally
located; its lower end horizontal with upper margin of eye.
Opercular flap small, ear-shaped, with notched upper rim,
its tip above a horizontal through upper margin of eye by
distance equal to eye diameter.
Uppermost pectoral ray about on horizontal through
upper margin of eye. Pectoral fin deeply notched, 20
(15+2+3), rudimentary rays absent. Upper lobe quite short,
394 Records of the Australian Museum (2001) Vol. 53
Figure 43. Paraliparisplagiostomus n.sp. A, holotype, CSIRO T488-02, 6, 148 mm TL, 137 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 782, cleared and stained right pectoral girdle.
not reaching anal-fin origin; of 3 thick rays. Skin of proximal
surface of upper lobe attached at ray bases, about half way
to tips of notch rays, and Vs of lower lobe ray lengths.
Cartilaginous basal lamina of pectoral girdle with 3 (3+0)
round radials, all in upper half of lamina; the lowermost
small. Fenestrae in cartilaginous basal lamina absent. Helve
of coracoid long and thin.
Body not deep but humpbacked, greatest depth at first
dorsal rays, about equal to HL; dorsal contour sloping
anteriorly and more gently posteriorly from this point,
ventral profile straight. Horizontal midline anteriorly
touching upper margin of eye. Predorsal length short.
Parapophyses of vertebra 11 short, haemal spine absent.
Interneural of first dorsal ray between neural spines 6
and 7, two free interneurals present anteriorly. Epineural
ribs on vertebrae 2-10, anterior 7 quite stout and short,
length about equal to 2.5 vertebrae. Epipleural ribs well
developed on vertebrae 3-16, and as small but distinct
commas on vertebrae 17-23; anterior 6-8 ribs quite stout,
length up to 3 vertebrae. Lateral keel-like ridges at sides
of body above pectoral fin present, not well developed.
Vertical fins overlap caudal Vs. Anus below first third of
postorbital space; mandible-anus distance short. Skin
quite dense, opaque, prickles absent. Subcutaneous
gelatinous tissue well developed, especially on lower
surface of head and pectoral fins. Pyloric caeca similar
to one another.
Colour. Head and body uniformly dark, brownish-black; lips,
inner surface of subrostral fold, and chin black, anal area and
urogenital papilla black. Mouth black, and tooth plates dark
grey, tongue black-dotted, branchial cavity black and gill arches
dark grey. Peritoneum black, pyloric caeca and stomach pale.
Distribution. West coast of Tasmania, around 990 m.
Etymology. The specific epithet, plagiostomus is from the
Greek plagios “placed on lower surface of head” and stoma
mouth.
Comparative notes. The new species is in group II. It differs
from all other described Paraliparis, with the exception of
P. badius, in having an inferior mouth and the gill opening
entirely above eye level. See description of P. badius for
comparison.
Paraliparis retrodorsalis n.sp.
Fig. 44
Material examined. Holotype CSIRO H1935-01, 9 , 159
mm TL, 145 mm SL. FRV Soela, stn. So 1/89/56,37°34.53’S
138°57.00'E, South Australia, W of Cape Martin, 1205-1175
m, 1 Feb. 1989; radiograph 682 E-l; pectoral girdle 736.
Diagnosis. Vert. 68, D 62, C 8, radials 4 (3+1? see below),
round. Body not hump-backed. Mouth horizontal,
subterminal, lower jaw included; chin pore interspace equal
to pore diameter, not in a pit, without skin fold anteriorly. P
21 (15+2+4), rudimentary notch rays absent. Interneural of
first dorsal ray between neural spines 7 and 8. Body and
peritoneum black, mouth black, tongue black-dotted, tooth
plates dark.
Further description. Counts: D 62, A 56, P 21, C 8 (4/4),
Vert. 68 (12+56), radials 4 (3+1?), pc 5, gr 7, pores 2-6-7-
1. Ratios: HL 18.1, its width 18.3 (57), and depth 16.7 (92),
bd c. 19.9 (110), bdA 14.8 (82), preD 28.3, preA 36.6, ma
13.1, aAf 25.9, UPL 12.4 (69), LPL 11.0 (42% HL, 89%
UPL), NL 4.1 (33% UPL), E 4.2 (23.3), gs 3.4 (19.1), sn
6.3 (34.4), po 9.0 (50), io 6.9 (38.2), so 2.8 (15.3), uj 9.0
(49.6), lj 8.3 (45.8), pc 4.5.
Head small, 5.5 in SL, dorsal contour gradually sloping
anteriorly to low snout; compressed and deep, depth 1.6 its
width. Snout projecting anteriorly, acutely angled, length
1.5 eye. Subrostral fold present, not entirely covering upper
lip. Upper margin of eye almost touching dorsal profile of
head, suborbital about Vs eye; pupil very large. Interorbital
width 1.6 eye. Nostril small, with low rim, on horizontal
Stein et al .: Australian liparids 395
Figure 44. Paraliparis retrodorsalis n.sp. A, holotype, CSIRO H1935-01, $, 159 mmTL, 145 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 736, cleared and stained right pectoral girdle, rays reconstructed
from left side.
with lower half of pupil. Mouth horizontal, sub terminal;
lower jaw included. In ventral view, upper tooth plates
entirely visible below lip; lower jaw tapering anteriorly, chin
rounded. Oral cleft reaching to below eye centre, lower jaw
deep below end of cleft. Honeycomb tissue thick, well
developed on chin. Teeth simple, quite large on upper jaw,
smaller on lower jaw, about 24-25 regular rows of up to 8-
10 teeth each anteriorly. Diastema of upper jaw distinctly
wider than in lower jaw. Circumoral pores small, chin pores
closely set, interspace equal to pore diameter, not in any
depression, anterior skin fold absent, interspace
pigmented. Gill opening small, 0.8 eye diameter, entirely
above pectoral base; dorsal end slightly above horizontal
through upper eye margin, lower end level with lower
margin of pupil. Opercular flap small, triangular, with
notched upper side, covering 3 A of gill opening, tip level
with upper margin of pupil.
Uppermost pectoral ray horizontal with lower margin of
eye. P 15+2+4, bilobed, upper lobe not reaching anal-fin
origin; notch rays not rudimentary, lower lobe short, its
middle rays thin, elongated. Basal cartilaginous lamina
short, wide in middle. Radials 3+1, round, third radial
smaller, located at posterior margin of lamina; only part of
the fourth radial present in cleared and stained preparation.
Fenestrae absent. Scapula small, helve short with two thin
lateral ribs, coracoid helve thin, not long.
Body elliptic, shallow, 5 in SL, deeper than HL, dorsal
contour evenly sloping from deepest point (above dorsal
origin) anteriorly and posteriorly. Horizontal midline close
to oral cleft. Predorsal length long. Anterior dorsal rays
reduced; intemeural of first dorsal ray between neural spines
7 and 8; three free interneurals present anteriorly between
spines 4 and 7. Length of haemal spine of last abdominal
vertebra (12) about half that of next one. Costal ridges weak,
but distinct. Epineural ribs present on vertebrae 3-18,
epipleural ribs on vertebrae 3-15, thin, length of anterior
not more than two vertebrae. Anus-anal-fin long. Vertical
fins overlap caudal fin for about one-third. Skin quite thick,
opaque, prickles absent. Subcutaneous gelatinous layer
weakly developed. Pyloric caeca thick at their bases, of
similar length. Diameter of ripe eggs c. 2.5 mm.
Colour. Head and body very dark, uniformly ink-black,
subrostral fold as black as head, lips paler, dark grey. Mouth
black, tongue black-dotted, tooth plates dark. Pores inside
pale but not contrasted with head. Branchial cavity black,
gill arches dark grey, peritoneum black, stomach and pyloric
caeca pale.
Distribution. Off South Australia at 1205-1175 m.
Etymology. Retrodorsalis from the Latin retro —rear, and
dorsalis —dorsal, referring to the relatively posterior position
of the first dorsal-fin ray between vertebrae 7 and 8.
Comparative notes. Paraliparis retrodorsalis belongs to
group Ilia and is distinguished by its very dark, ink-black,
uniform colour, dark tooth plates, black mouth, more
posterior dorsal-fin origin (first dorsal ray between neural
spines 7 and 8), included lower jaw, upper jaw teeth
somewhat larger than in lower jaw, diastema of upper jaw
wider than in lower. It is most similar to P. costatus but
differs in having weak ribs and lacking a costal keel (v.
epineural and epipleural ribs strongly developed, keel
distinct), D 62 (v. 66), first dorsal intemeural between neural
spines 7 and 8 (v. 5 and 6), 3 free interneurals present (v.
one), C 4/4 (v. 1+3/3+1),and other characters.
396 Records of the Australian Museum (2001) Vol. 53
Figure 45. Paraliparis tasmaniensis n.sp. A, holotype, CSIRO H2679-02, $, 213 mm TL, 201 mm SL. B, ventral
view of mouth and upper jaw tooth plate. C, P 732, cleared and stained right pectoral girdle.
Paraliparis tasmaniensis n.sp.
Fig. 45
Material examined. Holotype CSIRO H2679-02, $, 213
mm TL, 201 mm SL. FRV Soela, stn. So 2/89/32,40°26.64’S
143°18.36'E, NW Tasmania, SW of King Island, 1000-1100
m, 6 Mar. 1989; radiograph 684 F; pectoral girdle 732.
Diagnosis. Vert. 67, D 59, C 8, radials 3, upper two notched.
Pectoral-fin upper lobe of 17 rays, widely separated from
lower lobe of 4 rays by a wide notch with 1 rudimentary
ray. Mouth horizontal, subterminal, teeth very small. Chin
pores two, pit and anterior skin fold absent. Body deep, 125%
HL, depth bdA 113. Eye small, 18.8% HL; gill opening equal
to eye, reaching ventrally to horizontal through eye centre.
Body deep, hump-backed, bdA 113% SL. HL 20% SL, preA
37% SL. Colour black, peritoneum black, mouth and tongue
dark grey, tooth plates pale.
Further description. Counts: D 59, A 55, P 17+lr+4, C 8
(4/4), Vert. 67 (11+56), radials 3 (3+0), pc 7, gr 8, pores 2-
6-7-1. Ratios: HL 19.9, its width 10.9 (55) and depth 17.4
(88), bd 24.9 (125), bdA 22.4 (113), preD 24.9, preA 36.8,
ma 13.9, aAf c. 22.9, UPL 13.2 (66), LPL unknown, E 3.7
(18.8), gs 4.0 (20), sn 7.0 (35), po 10.6 (53.5), io 8.2 (41.3),
so 3.6 (18.0), uj 8.5 (42.5), lj 8.4 (42), pc 7.5.
Head not large, 5 in SL, deep at occiput, depressed above
eye; not wide, depth 1.6 its width. Snout large, 1.9 eye,
rounded, slightly protruding above upper jaw, most dorsal
point horizontal with upper third of eye. Subrostral fold
deep, covering upper lip almost entirely. Nostril not large,
rim raised, on level with lower half of pupil. Eye small, 5.3
in HL, not entering dorsal profile of head. Suborbital
distance about equal to eye, postorbital length large. Mouth
horizontal, subterminal, reaching to below anterior margin
of pupil. Lower jaw subterminal, slightly shorter than upper;
jaw deep below end of mouth cleft; honeycomb tissue thick.
In ventral view, lower jaw tapers anteriorly, chin rounded;
when upper lip displaced, upper tooth plate edges visible.
Teeth very small, not prominent, tips blunt, arranged in 23-
26 regular rows of up to 9-12 teeth each anteriorly. Diastema
of upper jaw wide, of lower jaw narrower. Circumoral pores
small, chin pores not in a pit or depression, anterior skin fold
absent, interspace equals 1.5 pore diameters; interspace
pigmented. Gill opening vertical, small, 1.1 eye; dorsal end
above eye level, ventral end level with eye centre. Opercular
flap covering about 2 A of gill opening, small, triangular, upper
margin notched; tip horizontal with upper margin of eye.
Uppermost pectoral ray level with eye centre. P
17+(l)+4, with upper lobe rays widely separated from lower
lobe rays (lowest shortest) by a wide notch including a single
ray, that one rudimentary, visible only by clearing and
staining. Upper lobe not reaching anal-fin origin, lower lobe
length unknown (broken). Skin on proximal surface of upper
lobe attached at about % lobe length, unknown in notch.
Lowermost pectoral ray below posterior margin of eye.
Radials 3+0, upper two notched; R3 small, round. Two
fenestrae present below scapula and first radial; a slit-like
rudimentary fenestra present below 2nd radial. Scapula
notched ventrally, helve short, with two stout lateral ribs.
Coracoid notched ventrally, with very long, thin, shaft.
Body hump-backed, elliptic, deep, 4 in SL, deepest at
dorsal-fin origin; depth at A origin also greater than HL.
Dorsal profile significantly rounded ventrally, both
anteriorly and posteriorly from point of greatest body depth.
Horizontal midline of body touching lower margin of eye.
Stein et al .: Australian liparids 397
Interneural of first dorsal ray between vertebral spines 6
and 7; anteriorly, 3 free interneurals present between spines
3-6. Haemal spine of last abdominal vertebra about half as
long as next. Epineural ribs on vertebra 2-15, epipleural
ribs on 3-17; thin, not long, anteriormost not longer than
2.5 vertebrae. Costal keel thin, slightly prominent. Anus
below middle of postocular space. Degree of caudal overlap
by vertical fins unknown. Skin quite thick, prickles absent.
Subcutaneous gelatinous tissue moderately developed.
Pyloric caeca similar. Egg diameter (ripening) 1.7-2.0 mm.
Colour. Head, body, snout, chin, and subrostral fold
uniformly black; upper lip dark grey. Mouth and tongue
dark grey, tooth plates pale. Pores paler than head.
Peritoneum black, stomach and pyloric caeca black;
posterior of rectum near anus blackish. Branchial cavity
black, gill arches dark grey.
Distribution. Off NW Tasmania at 1000-1100 m.
Etymology. The specific epithet is a reference to
Tasmania—the place of capture.
Comparative notes. Paraliparis tasmaniensis is in group
Ilia. It is distinguished by having a rudimentary notch ray
(v. notch rays not rudimentary) and 2 pectoral girdle
fenestrae (v. absent), a deep body, small eye, and short upper
jaw. In addition, P. tasmaniensis differs fromP. retrodorsalis
in being hump-backed (v. not hump-backed), radials 3+0,
upper two notched (v. 3+1, unnotched), two fenestrae in
pectoral girdle present (v. absent). It differs from P.
impariporus in its entirely black colour, in having two chin
pores (v. one pore), eye 19 (v. 26% HL); pectoral radials
3+0 (v. 4), two large fenestrae in cartilaginous basal lamina
(v. two rudimentary), and a wide pectoral notch with one
rudimentary ray (v. 2-3 normal notch rays).
Paraliparis sp. 1
Fig. 46
Material examined. CSIRO H549-05, ripe $, length c.
134+ mm SL (a few posteriormost vertebrae missing). FRV
Soela, stn. So 3/86/58, 41°50.4’S 144°23.45'E, W coast of
Tasmania, W of Trial Harbour, 1328-1288 m, 25 May 1986;
radiograph 680E; pectoral girdle 799.
Diagnosis. Vert. 56+, D —, C —, radials 3, all notched.
Mouth subterminal, lower jaw included. Teeth simple, small.
Subrostral fold absent. Chin pores in a small pit, interspace
unpigmented, equal to their diameter. Ventral end of gill
opening and dorsal pectoral-fin ray level with lower margin
of pupil. Body straight-backed, ventral contour much more
curved. Costal ridges slightly developed. P 19-21,
rudimentary rays absent. Head less than 20% SL, predorsal
120% HL, preanal 193%. Body black, peritoneum black.
Further description. Counts: Vert. 10+46+, D —, A—, P
19-21, C —, radials 3+0, all notched; fenestrae three, pc 6,
gr 8, pores 2-6-7-1. Ratios: HL less than 20% SL; in% HL:
head width 58, and depth 77.5, bd 116, bdA 89, preD 120,
preA 193, ma79, aAf 114,UPL69,LPL46.5HL(67%UPL),
NL 12 (18% UPL), E 24.8, gs 19.4, sn 31.0, po 50.4, io 35.7,
so 17.0, uj 42.6, lj 39.5, pc 31 (i.e., less than 6.3 SL).
Head less than Vs SL, compressed, depth 1.3 width.
Dorsal contour straight, slanting anteroventrally. Snout
rounded, projecting anterior to upper jaw, its highest point
level with upper pupil margin, the most prominent point
level with eye centre. Subrostral fold absent. Nostril level
with lower margin of pupil. Eye almost entering dorsal
contour of head. Suborbital distance short, less than 3 A eye.
Pupil 2 A eye. Mouth subterminal, cleft reaching to anterior
Figure 46. Paraliparis sp. 1. A, CSIRO H549-05, ripe 9, c. 134+ mm SL (a few posteriormost vertebrae missing).
B, ventral view of mouth and upper jaw tooth plate. C, P 799, cleared and stained right pectoral girdle.
398 Records of the Australian Museum (2001) Vol. 53
margin of pupil, jaws short. Lower jaw included. Lips not
wide. Chin deep below end of mouth cleft; in lateral view
tip slanted, honeycomb tissue well developed. Teeth simple,
very small, slightly prominent, in c. 26 rows on upper jaw,
19 on lower, about 13 and 10 teeth per row anteriorly. Tooth
plate on upper jaw wider than on lower. Diastema of upper
jaw narrow, in lower jaw absent. Circumoral pores small;
chin pores in a small pit, interspace unpigmented, equal to
one pore diameter. Gill opening shorter than eye, ventral
end level with lower margin of pupil. Opercular flap
triangular, dorsally notched, sharp tip level with upper
margin of pupil.
Upper pectoral ray level with lower margin of pupil, not
reaching anal-fin origin; lower lobe short. Lowermost ray
insertion below first third of postorbital space. Skin on fin
missing. Radials 3+0, all notched. Fenestrae three, below
scapula, below R1 and R2. Coracoid helve long, thin. Helve
of scapula unusually strong. P 14+2+3 (L), 15+2+4 (R),
rudimentary rays absent.
Body straight-backed, ventral contour much more curved
than dorsal. Horizontal midline through lower margin of
eye. Greatest depth at dorsal-fin origin. Abdominal and
caudal vertebrae clearly distinguishable; last abdominal
vertebra lacks haemal spine, that of first caudal vertebra
very long. Parapophyses of abdominal vertebrae obviously
not joined. First dorsal-fin ray short, its interneural between
neural spines 4 and 5, 1 free interneural present anteriorly
between neural spines 3 and 4. Costal ridges weakly developed.
Epineural ribs on vertebrae 2-8, as long as 3 vertebrae;
epipleural ribs on 2-14, thin, hardly visible. Anus almost below
gill opening. Pyloric caeca slender. Ovarian eggs to 2 mm.
Colour. Body skinned. Snout and lower jaw ink-black. Upper
lip as dark as head, lower lip and chin slightly paler. Pores not
contrasting. Mouth and tongue dark grey. Branchial cavity
black, gill arches grey. Peritoneum black, pyloric caeca and
stomach pale. Anal area and end of rectum black.
Distribution. West coast of Tasmania at 1328-1288 m.
Comparative notes. A member of group Illb, distinguished
by its body shape, a straight back and greatly curved ventral
contour, chin pore interspace unpigmented, lower jaw
included, subrostral fold absent, anus almost below gill
opening, and lower jaw below mouth cleft deep. Most
similar to P. infeliciter, it differs in mouth cleft reaching to
below pupil (v. to below anterior margin of eye), P 14/
15+2+3/4 (v. P 17+2+3), fenestrae three (v. one), helve of
coracoid thin (v. stout), helve of scapula not strong (v.
strong), lower jaw included (v. subterminal), and anus almost
below gill opening (v. first third of postorbital space).
Paraliparis sp. 2
Fig. 47
Material examined. CSIRO H555-04, unripe S , 144 mm TL,
131 mm SL. FRV Soela, stn. So 3/86/38, 42°20.6'S
144°37.25’E, W coast of Tasmania, W of Cape Sorell, 1376-
1404 m, 17 May 1986; radiograph 679-1; pectoral girdle 797.
Diagnosis. Vert. 67, D 61, C 8, radials 2, round. Mouth
sub terminal, short, lower jaw sub terminal. Teeth simple,
small. Subrostral fold absent. Chin pores in a small pit,
interspace pigmented equal to their diameter. Ventral end
of gill opening and dorsal pectoral ray level with lower
margin of orbit, gill opening reaching 2nd pectoral ray.
Anterior 3 dorsal rays short. Body low, shape elliptic, not
humpbacked. Costal ridges present. P 16+2+4, rudimentary
rays absent. Head small, 17.7% SL, predorsal length short,
18.5% SL, preanal 34%. Head black, peritoneum black with
pale background.
Figure 47. Paraliparis sp. 2. A, CSIRO H555-04, unripe 6 , 144mmTL, 131 mmSL. Eye missing. B, ventral view
of mouth and upper jaw tooth plate. C, P 797, cleared and stained right pectoral girdle.
Stein et al .: Australian liparids 399
Further description. Counts: D 61, A 54, P 22, C 8 (1+4/4),
Vert. 67 (11+56), radials 2+0+0, round; fenestrae absent, pc 4,
gr 7, pores 2-6-7-1. Ratios: HL 17.7, its width 9.4 (53), and
depth 12.8 (72.4), bd 15.1 (85), bdA 13.6 (77), preD 18.5,
preA 34.0, mac. 12.2, aAf c. 22.1, UPL 13.0 (73.3), LPL 10.7
(60.3% HL, 82% UPL), NL 3.1 (23% UPL), E —, orbit 4.7
(26.7), gs c. 3.7 (20.7), sn* 5.3 (30.2), po* 8.5 (48.3), io* 6.3
(35.3), uj 7.9 (44.4), lj 7.5 (42.2), pc 5.7 (* measured from
margin of orbit, eyeball missing).
Head very small, 5.6 in SL, compressed, depth 1.4 its
width. Eyeball missing, orbit close to upper contour of
head but suborbital distance very short, about half of
orbit. Snout deep, rounded, slightly projecting anterior
to upper jaw; its highest point close to level of upper
margin of orbit, most prominent point slightly below.
Subrostral fold obviously absent. Nostril slightly below
level of eye centre. Mouth barely subterminal, its cleft
short, reaching only to below anterior margin of orbit.
Lower jaw subterminal, almost included. Teeth very
small, simple, slightly prominent, in 16-17 rows, about
8 teeth anteriorly. Narrow diastema in upper jaw. Chin
pores in a common small pit, interspace pigmented,
equalling their diameter. Chin tip slanted in lateral view
(not right-angled), deep below end of mouth cleft. Gill
opening reaching 2nd pectoral ray, its ventral end level
with lower margin of orbit. Opercular flap triangular, tip
directed posteriorly, level with eye centre.
Upper pectoral ray level with lower margin of orbit. P
16+2+4, notch rays about V* UPL. Fin membrane missing.
Base of lowest ray below posterior margin of orbit. Upper
lobe not reaching anal-fin origin. Radials 2+0+0, round;
fenestrae absent. Coracoid helve thin, long.
Body not humpbacked, low, of elliptic shape, dorsal
and ventral contours similar. Maximum depth 6.6 in SL,
deepest at anal-fin origin, contours sloping evenly
anteriorly and posteriorly. Horizontal midline through
centre of orbit. Predorsal length short. Two free anterior
dorsal interneurals, the first between vertebrae 2 and 3;
anterior 3 dorsal rays short. First dorsal ray above tip of
opercular flap. Costal ridges not large, but distinct.
Vertical fins overlapping about half of caudal. Anus
below first third of postorbital space. Pyloric caeca of
similar lengths, with sharp tips.
Colour. Body and head skinned, snout, lips, and chin black
with pale background, skin thin and semitransparent. Tooth
plates pale, mouth grey, tongue grey, dotted. Branchial
cavity dark grey, gill arches grey. Peritoneum black with
distinct pale background. Pyloric caeca and stomach pale.
Distribution. West coast of Tasmania, 1376-1404 m.
Comparative notes. This specimen is quite different
from the other species of its group. It seems to be a new
species, but because of its poor condition (skinned body,
missing eye balls) we prefer not to name it but designate
it as “sp. 2”. It should be described when better specimens
become available. It is a member of group IHb,
distinguished by its low, elliptically-shaped shallow body,
short predorsal length, peritoneum not ink-black but with
a pale background, anterior three dorsal rays short, mouth
cleft reaching only to below anterior of orbit, costal
ridges present, gill opening reaching to 2nd pectoral ray,
and radials 2+0+0, round.
Incertae sedis
Paraliparis sp. (cf. copei group)
Fig. 48
These specimens are in such poor condition that we can
provide only a partial description of them. They are clearly
distinct from the other specimens examined for this paper,
but so many characters are unknown that description awaits
better material. Because of their condition and their
similarities, we treat them together although we recognize
that they are probably not the same species.
Material examined. AMS 122809-036, ripe S, c. 178 mm
SL. FRV Soela, 18°40’S 116°42'E, Indian Ocean off Western
Australia, 250 km NW of Port Hedland, 584-592 m, 4 Apr.
1982; radiograph ZISP 2054; pectoral girdle 738. AMS
122813-018, ripe S, 154+ mm TL. FRV Soela, 18°32'S
116°50'E, Indian Ocean off Western Australia, 250 km NW
of Port Hedland, 658-660 m, 6 Apr. 1982; radiograph ZISP
20253; pectoral girdle 737.
Description. The specimens are similar in having the head
short, profile rounded; mouth subterminal, lower jaw
included, tooth plates entirely visible in ventral view. Teeth
small, stout canines, uniserial for posterior % of jaws,
biserial for anterior third, similar in both jaws. Opercle
horizontal, broadly curved, with tip pointing dorsally. Gill
opening damaged. Pectoral fin damaged, upper lobe of at
least 14 rays, lower lobe of at least 3 rays. Radials 3 or 4.
Prominent costal keels present. Scapula stout, helve short,
triangular; coracoid helve very long, with ventral
strengthening blade. Body relatively deep but not hump¬
backed, elliptical. Pyloric caeca 3?, stout, digitate. The
specimens differ significantly:
In I 22813-018, Vert. 63+ (=11+52+), epineural ribs on
vertebrae 2-13, epipleural ribs on 2-15; anteriormost 8 of
both series strong, long, sabre-like, up to 4 vertebrae in
length. D insertion between vertebrae 4/5. Right girdle:
2+0+1, R1 ventrally notched, R2 dorsally and ventrally
notched. Fenestrae between R1 and 2, one rudimentary
fenestra below R2. Coracoid helve long, thin. Left girdle
with 3+1 radials.
Figure 48. Paraliparis sp. (cf. copei group), AMS 122809-036,
6 , c. 178 mm SL. P 738, cleared and stained right pectoral girdle.
400 Records of the Australian Museum (2001) Vol. 53
In I 22809-036, Vert. 42+ (=11+31+), epineural ribs on
at least vertebrae 2-10, epipleural ribs on at least vertebrae
2-9. Radials 3+1, R1 and 2 unnotched, R3 notched dorsally.
One foramen above R3. Rays partially missing. Two
rudimentary rays in notch.
Colour. Body colour unknown, lips brownish. Orobranchial
cavity and gill arches pale, brown-dotted. Peritoneum very
dark brown or black, stomach and caeca pale.
Distribution. Indian Ocean off Western Australia at 584-
660 m.
Comparative notes. These two specimens differ
significantly in radial number and presence of notches.
There is no question that in their dentition, the two differ
greatly from all the other Australian species we have
examined, which have teeth forming wide bands. In
dentition these two are similar to the U P. copei-P. rosaceus
group”, that includes the Pacific P. attenuatus Garman,
1899, P. rosaceus Gilbert, 1890, P. grandiceps Garman,
1899, P. nassarum Stein & Fitch, 1984, and P. paucidens
Stein, 1978b; the Atlantic P. copei Goode & Bean, 1896
and P. copei wilsoni Richards, 1966; and the Antarctic P
copei gibbericeps Andriashev, 1982a and P. c. kerguelensis
Andriashev, 1982b. All of these species look similar and
have very narrow tooth bands generally suggesting a trend
from biserial teeth to uniserial and finally to only a few
scattered teeth.
Discussion
Presence of a diverse liparid fauna in Australian waters is
not surprising, given their occurrence on the Pacific coast
of South America, at all benthic depths throughout Antarctic
waters and islands, and off southern Africa. Although we
might have assumed the presence of liparids in waters of
South Australia, we were amazed to find so many species in
such a small collection, suggesting that the family has more
taxonomic and ecological diversity than previously thought.
Concerned that so many species appeared to exist in our
material, we tried to reduce the number by finding characters
supporting determination of conspecificity, but instead
found clear distinctions. In our experience, these differences
are trenchant. Therefore, we describe most of these
specimens as representatives of new species. Furthermore,
we expect that as exploration of Australian slope and abyssal
depths continues, even more species will be found.
Morphology and relationships to other liparids. The
centre of taxonomic and ecological diversity and numerical
abundance of the family is in the North Pacific, where most
genera, both primitive and advanced, occur. In particular,
the primitive genera Liparis and Careproctus are very
diverse in the Bering Sea, Sea of Okhotsk, and adjacent
regions, where many monotypic genera are also present.
Some of these species reach very large size (for liparids),
up to 700 mm TL ( Liparis niger Soldatov & Lindberg, 1930;
Andriashev, unpublished). In contrast, in the Southern
Hemisphere, Liparis is absent and the approximately 45
Careproctus species are not morphologically diverse.
Southern Careproctus species have more advanced
character states than northern species, indicating their
evolutionary distance from many North Pacific relatives,
and all are medium-sized to very small. Furthermore, the
endemic Southern Hemisphere genera ( Edentoliparis
Andriashev, Eknomoliparis Stein et al., Genioliparis
Andriashev & Neelov, Notoliparis Andriashev) are clearly
derived, not primitive. Finally, the cottoids, the sister group
of the cyclopteroids (that is, the cyclopterids and liparids),
are also widely distributed and highly diverse in the North
Pacific (Yabe, 1985). Our results strongly support the North
Pacific as the centre of origin, centre of morphological
diversity, and the location of the most primitive genera.
Southern Hemisphere liparids are characterized by
radiation within genera rather than among genera
(Andriashev, 1998). Thus, there are many endemic species
but only a few endemic genera. In addition, the apomorphic
character states of Careproctus, and the absence of Liparis
from the Southern Hemisphere, strongly indicate that the
liparid fauna of the Southern Ocean is young and secondary.
The Australian Paraliparis clearly form a distinct species
group quite different from the Antarctic species. Owing to
their unusual morphological similarity we hypothesize that
they form a “species flock” in the sense of Greenwood
(1984:18), “an aggregate of several species... if its members
are endemic to the geographically circumscribed area under
consideration and are each others’ closest living relatives” and
Ribbink (1984: 24), “an assemblage of a disproportionately
high number, relative to surrounding areas, of closely related
species which apparently evolved rapidly within a narrowly
circumscribed area to which all the member species are
endemic.” We believe they typify Pianka’s (2000) statement
that “Species-rich genera are both rare and extremely
interesting because they imply that recent bouts of speciation
and niche diversification have occurred, leading to adaptive
radiation. Such events of rapid evolution promote species
diversity”. Future analyses based on specimens not yet
collected will provide the test of our hypothesis.
Paraliparis. Australian and Southern Ocean Paraliparis are
morphologically distinct, although clearly members of the
same genus. The combined range of intraspecific meristic
variability across all 28 Australian species is small; it is
similar to the range of interspecific variability within single
species from the Southern Ocean. For example, the range
in number of vertebrae in all Australian species is 7 (65-
71); moreover, 21 species (77.7%) have 65-68 vertebrae
(Table 3). In the 29 Southern Ocean species listed by
Andriashev (1986), the range is 30: number of vertebrae varies
from 51 (P. trilobodon) to 81 (P tetrapteryx ) (Andriashev, 1986:
156). In P. thalassobathyalis Andriashev, the range is 6
(Table 4). Variability in fin-ray number is similar: Australian
species have D 58-64 (v. 45-73 in Southern Ocean species),
A 52-57 (v. 41-66), P 19-24 (v. 15-31). Uniformity of the
Australian Paraliparis in caudal-fin ray number is very high
compared to the Antarctic species: 25 species have C 8, and
only 2 have C 9 (v. 4-11 in the latter). Variability in pyloric
caeca number is similar: 3-7 (v. 0-15).
Table 3. Interspecific variation: number of vertebrae in Australian
Paraliparis species.
vertebrae
65
66
67
68
69 70 71 n
number of species
6
7
4
4
2 3 1 27
Stein et al.: Australian liparids 401
Table 4. Intraspecific variation: number of vertebrae in P.
thalassobathyalis from Meteor Sea Mount (from Andriashev,
1986).
vertebrae 56 57 58 59 60 61 62 n
number of specimens 0 4118 3 1 1 28
Some important morphometric characters vary little in
Australian species. All have a short gill opening from 3.0-
4.3% SL in length; in the majority, it is less than 4% SL.
The range in Southern Ocean species is from pore-like (1-
2% SL) in the P. copei group, to 15-16% SL in P. antarcticus
and P. meganchus (Andriashev, 1986). All Australian species
have closely-set chin pores: 29% have chin pores in one
common pore or interspaced at a distance less than one pore
diameter, and 61% have them separated by a distance equal
to one pore diameter; in only 3 species is this distance equal
to two pore diameters. In Southern Ocean species, the
position of the chin pores is more variable: they are in one
common pore, are more or less closely set, or are widely
spaced so that distance pm,-pm, equals or is larger than pm-
pm 2 . In many Australian species the chin pores are not only
closely set, but also placed in a more or less developed common
pit, or have an anterior skin fold not found in Southern Ocean
species. The snout is often deep, with widely spaced nasal
pores (n, and n 2 ). Finally, most have a small mouth and
small to exceptionally small teeth (21 species, or 75%).
Pectoral girdle. Almost half the Australian species have
plesiomorphic pectoral girdle character states typical of Liparis
(notched radials and different types of fenestra between radials,
often associated with the notches), although they are not as
well developed and their occurrence is more variable than in
Liparis. In contrast, only one “Antarctic” Paraliparis species
(Paraliparis stehmanni Andriashev, 1986) has notched radials
and interradial fenestrae. In the Australian species, these
features are different with respect to their quantity, shape, size,
location, and position. Because almost all species are
represented by single specimens, it is impossible to draw
conclusions regarding intraspecific variability.
It is also worth noting the occurrence in P. coracinus of the
horseshoe-shaped R3 forming part of the margin of the pectoral
girdle. As stated in the comparative notes for that species, there
are now three species known with one or more such radials.
Despite this similarity, it is clear that the new species is not
closely related to either of the other two species, which are
Antarctic and very different in many other respects.
Occurrence of all three radials in the upper half of the
basal cartilaginous lamina is unusual in Paraliparis. It
occurs in 12 Australian species ( ater ; auriculatus,
australiensis, avellaneus, badius, gomoni, infeliciter, lasti,
piceus, plagiostomus, tasmaniensis, sp. 1). Previously
known only from the Chilean P. merodontus Stein et al.,
1991, occurrence of this character state in many more
species suggests it is another variant of radial position that
may represent a distinct evolutionary branch. This character
state, the horseshoe-shaped R3, and other characters support
Andriashev’s (1986: 187) hypothesis that “the complex
mosaic of characters suggests a probable invasion by
liparidids [sic] into the southern hemisphere repeatedly and
by species at different levels of evolutionary development .”
In other words, by species from different clades.
Epineural/epipleural ribs and costal ridge. The absence of
pleural ribs is a characteristic morphological feature of the
genus Paraliparis (Kido, 1988), in which the usual character
state is short, thin, curved epineural and epipleural ribs on
abdominal and anterior caudal vertebrae. However, in some
Australian species, we found greatly developed epineural
and epipleural ribs which, at their greatest development,
produce a distinct external keel-like costal ridge above the
abdominal cavity on each side of the fish. This is a
previously undescribed character unique to these species
among liparids.
Epineural ribs are present from the 2nd (less commonly
3rd) to 6-19th vertebrae in all Australian species. Anteriorly,
they start at the vertebral neural arches, and posteriorly are
at the parapophyses and caudally, at the haemal arches.
Epipleural ribs occur anteriorly on vertebrae 2-3,
occasionally on 4 or 5 and then to the 11—25th vertebrae,
depending upon species. Anteriorly, they are located on the
vertebral body and posteriorly on the parapophyses and
haemal arches. They probably occur in all species, although
we could not see them clearly in radiographs of P. infeliciter,
labiatus, and piceus.
Usually both epineural and epipleural ribs are slender
and short, no longer than 1.5-2 vertebrae, occasionally as
long as 3 vertebrae. However, in three species (P. costatus,
P. dewitti, and P sp. (cf. copei group) from the Indian Ocean)
the anterior 7-9 ribs (both rows) are stout and elongated,
reaching 4-5 vertebrae in length (Fig. 11). In these species,
the ribs form the support for the costal ridge, which may be
present to some extent in other species (P. lasti, Paraliparis
sp. 2, for instance), but is much less well developed. This
seems to be an apomorphic character unique to these species.
Chin pores. The Australian liparids demonstrate all
variations of chin pore character states, from fused (i.e., a
single pore), paired and touching, with an interspace from
much less than one pore diameter to two pore diameters, in
a pit, not in a pit, and finally, with a distinct skin fold anterior
to the pore pair. We have found little intraspecific variability
in this character in other species, and have therefore used it
consistently to distinguish the new species where possible.
We do not understand the functional significance of these
interspecific variations, although we assume that, because
these pores are so close to the mouth, they are related to
feeding behaviour or food habits. In studies of food-finding
by other Antarctic fishes (mainly Nototheniidae) Janssen
(1996) considered the chin pores the most important
cephalic pores.
Colour. One of the significant differences between the
Australian and Antarctic species is colour. The body and
body cavities of the Australian species are characteristically
darkly pigmented (including black, brownish-black, dark
brown, and brown). Twenty-three species (82%) are very
dark; only 5 (18%) are paler (nut brown, light brown). In
contrast, Southern Ocean species are mainly pale after
fixation (61%), and in life are pale or rosy. Only a few are
lilac-rose; this coloration persists after fixation, becoming
paler. One species, P. cerasinus, is quite dark grey after
preservation, although cherry-red in life. Brown or black
species are very rare; P. trunovi and P. leucoglossus are the
only Southern Ocean species similar in colour to those of
Australia. We suggest that these colour differences reflect
absence of close relationships between the two groups.
402 Records of the Australian Museum (2001) Vol. 53
Summary. Australian Paraliparis species are much more
similar to each other than are co-occurring Paraliparis
species in other geographic areas such as Antarctica
(Andriashev, 1986) or the northeastern Pacific (Stein,
1978b). Nevertheless, the Australian species differ distinctly
(although often subtly) from each other in many ways:
general body, snout, and opercular flap shape, degree of
development of subrostral fold (from deep to absent),
presence or absence of rudimentary rays in the pectoral-fin
notch, pectoral-fin mobility (the degree to which
corresponding pectoral-fin rays are free of gelatinous tissue
is highly variable between species), and state of chin pores
(at the surface, in a pit, protected by an anterior skin fold).
In some species the chin has a well-developed layer of
honeycomb tissue, or the chin profile is not slanted as usual
but is distinctly right-angled. The skin in some species is
quite thick and dense, but in others it is thin and
semitransparent, and the pale musculature seen through it
influences the general body colour. We suggest this is a
monophyletic group that evolved recently in the conditions
of oceanic isolation that resulted in the high endemism of
shallow-water Australian marine taxa (Wilson & Allen,
1987), and hypothesize that these distinct but small
morphological differences (relative to those evident among
Paraliparis in other regions) represent recency of evolution,
i.e., Australian Paraliparis are so similar because they
diverged recently. That all 29 species from southeastern
Australia occur at similar depths (987-1404 m, 20 species
at 987-1225 m) also supports this hypothesis. Although it
is difficult to generalize on the basis of so few specimens
of each species, their apparent sympatry is striking: one
trawl (1000-992 m) caught 5 different species (P. ater, P.
auriculatus, P. brunneus, P. delphis, and P. labiatus). In
our experience this is unusual.
Psednos. In comparison to other species of Psednos from
both the northern and southern hemispheres, the Australian
species have a large number of vertebrae (56-58 v. 39-47),
differently arranged pectoral radials (2+0+1 v. 1 + 1+1,
equally spaced), and notched radials with one or two
interradial fenestrae (v. round, unnotched, fenestrae absent).
Because these character states are plesiomorphic for the
genus, and are similar to those found in other liparid genera,
we suggest that these interpretations support the hypothesis
of Andriashev (1992, 1993) regarding a Southern Ocean
origin for the genus. However, there are a number of
Psednos species from elsewhere in the Pacific that are
undescribed. There seem to be two or three from off Mexico,
one from New Zealand, and possibly one or two more from
the central South Pacific (DLS, unpublished). Analysis of
evolution in Psednos awaits description of these species.
Distribution. Contrary to the situation in the Northern
Hemisphere, in the Southern Hemisphere shallow-water
liparids are generally absent (exceptions being the isolated
Falklands, South Georgia, South Shetland Is and the
southern tip of South America) and those present have deep¬
water ancestors (Andriashev, 1965). Deep-water species are
common in both hemispheres (Andriashev, 1998; Burke,
1930; Stein, 1978b; and others). Given the rich liparid fauna
of the Southern Ocean (Andriashev, 1998; Andriashev &
Stein, 1998; Stein & Andriashev, 1990), it is not surprising
that Australia also has a liparid fauna. Both Antarctica and
Australia are isolated “continental islands” with significant
endemic shallow water faunas. In Antarctica, the noto-
thenioids, dominant in shallower waters, are replaced at
greater depths by the secondary deep-sea fishes, especially
liparids and zoarcids (Andriashev & Stein, 1998),
suggesting that the latter arrived after the notothenioids were
already dominant. Otherwise we would expect to find
liparids abundant at shallower depths also. There is not an
obvious parallel in Australian temperate waters; data are
not yet available to provide even a hazy picture of the
distribution of Australian fishes below mid-slope depths.
Why are there no shallower liparids around Australia in
cool temperate regions such as Tasmania, considering that
some Southern Ocean islands (e.g., South Georgia), have
shallow species evolved from deeper water ones
(Andriashev, 1965)? The temperatures around Tasmania
(surface: 10.5-18.5°, Harris etal., 1987) are similar to those
in the North Pacific off Southern California (surface: 12.5-
20°, Anon., 2000), where post-larval liparids occur in the
plankton. Perhaps the answer is that the deeper water genera
do not have the same temperature tolerance as the shallow
water Liparis of the North Pacific and North Atlantic
Oceans.
Wherever deep-water liparids occur, their species
diversity is high, probably owing to their life history
characteristics. They have large benthic eggs and probably
have direct development (Kido & Kitagawa, 1986; Stein,
1980). In addition, a number of species in North and South
America are known reproductive commensals, laying eggs
in the carapaces of crabs and in other invertebrates (Able
& Musick, 1976; Andriashev & Prirodina, 1990; Balbontin
et al., 1979; Peden & Corbett, 1973; and others). For these
reasons, they are less likely to disperse or be dispersed, and
are more likely to be affected by topographic barriers such
as shallow water or deep submarine canyons. Thus, the
liparids include many genera that are closely tied to the
bottom, whose members are relatively unaffected by water
movement, and in which the species are easily isolated,
leading to “local” evolution and subsequent existence of
endemic species groups. Such a pattern is not unlike that
occurring in some terrestrial groups of isolated (but closely
related) taxa whose ranges are restricted by and to
topographic features such as mountain tops or canyons.
The odd pelagic genus Psednos is known from South
Africa and the North Atlantic, and specimens of undescribed
species are known from off Southern California, Mexico,
and New Zealand. It seems likely that representatives of
the genus occur worldwide at mesopelagic depths, but at
low population densities.
Recent exploration of deep-water areas in search of
commercially exploitable fish (Koslow et al., 1994; May
& Blaber, 1989; Williams et al., 1996) has led to a great
expansion of knowledge about fishes of the mid- and upper
Australian continental slope, for example, chondrichthyans
(Last & Stevens, 1994). Despite these collections, not
enough is known yet about the deep-water fauna to support
its biogeographic analysis. Koslow et al. (1994) analyzed
the mid-slope demersal fish community of southeastern
Australia and investigated its similarity to the mid-slope
communities of the eastern and western North Pacific and
North Atlantic and off New Zealand. At the generic and
family level, there was “substantial overlap” with North
Atlantic slope faunas, but little with that of the North Pacific.
They hypothesized this was related to the distribution of
Stein et al.: Australian liparids 403
the Antarctic Intermediate Water Mass, which does not
penetrate far into the Pacific, but which has been detected
as far as 20°N latitude in the North Atlantic, providing a
corridor for dispersal. They concluded that, therefore, the
Australian deep water fauna has a closer evolutionary
relationship with the North Atlantic fauna, supporting
Andriashev’s (1991) hypothesis of a southern transoceanic
liparid dispersal pathway. Liparids apparently dispersed
from the Antarctic into the South, and then North, Atlantic
Oceans after the opening of the Drake Passage 20-22
MYBP, possibly using midocean ridges (Andriashev,
1977). It is important to note that the Antarctic species
of the Nototheniidae (with one or two exceptions),
Artedidraconidae, Channichthyidae, and others did not
disperse northwards. Amaoka et al. (1990) surveyed fishes
off New Zealand and reached the opposite conclusion to
Koslow et al. (1994), stating that 113 genera (of 265) and
40 species (of 267) were common to New Zealand and
Japan. These results may not be directly comparable to those
of Koslow et al. (1994), however, because Amaoka et al.
(1990) included pelagic and benthic species from a wide
variety of depths, whereas Koslow et al. (1994) included
only mid-slope demersal species.
Despite the paucity of knowledge regarding development
and distribution of the Australian deep water fish fauna,
analysis of the shallow water fishes is suggestive. There is
a notably high degree of endemism in southern Australian
shallow water fishes and invertebrates without a pelagic
early life history (Wilson & Allen, 1987). Pleistocene sea
level fluctuated more than 200 m, at times creating land
barriers to dispersal, and resulting in species pairs on the
western and eastern sides of Tasmania. Some shallow water
fish families underwent great radiation, apparently because
“the temperate environment would appear to offer more
opportunity for niche specialization because of lowered
competition” (Wilson & Allen, 1987) compared to the
tropics where species diversity, and thus competition, is
much higher. “The four [fish] families exhibiting the most
speciation... are characterized by reproductive modes that
tend to discourage extensive dispersal” and “there are
several southern Australian temperate species of diverse
groups which also occur in similar latitudes in the northern
Pacific or which have close relatives there”. The presence
of at least 16 fa mi lies that also occur in the North Pacific is
presumably associated with narrowing of the tropical zone
and reduction of the barrier it presented (Wilson & Allen,
1987). All of the preceding factors may have played roles
in diversification of the liparids in Australian waters.
Conclusions. A few decades ago, we would have
confidently stated that only a few liparids exist in the
Southern Hemisphere. Now, however, it is clear that there
are probably at least as many species, but not as many genera
(about 120 species known in 7 genera v. more than 130 in
about 20 genera) in the Southern Hemisphere as in the
Northern, and that despite the work of the last two decades,
many more remain to be discovered and described, not only
from Australia, New Zealand, and Chile, but also from
Antarctica.
Although the possible derivation of some Atlantic liparids
from South Pacific or Southern Ocean ones is becoming
clearer, the origins of Australian liparids are unknown. They
are not closely related to Antarctic liparids and are unlikely
to be derived from North Atlantic taxa (Andriashev, 1998).
It is possible that Australian species are derived from those
on the west coast of Chile, which in turn seem likely to be
derived from North Pacific species.
In 1970, Giles Mead wrote in “The History of South
Pacific Fishes”, that the Order Scorpaeniformes “are
potentially a most significant group for historical analysis”,
that the history of the Suborder Cottoidei (which includes
the liparids) “will, when fully understood, form a
zoogeographical contribution of great importance”, and that
the fish fauna of cold-temperate Chile will be crucial in
this analysis. He further included the cold-temperate North
Pacific and the cottoids as one of the four sources of the
temperate South Pacific fish fauna. It is clear that when an
analysis of the historical biogeography of Southern
Hemisphere liparids is accomplished, it will provide
significant advances in knowledge of the relationships
among the secondary deep-water fish faunas of the South
Pacific and how they evolved.
Acknowledgments. We thank P. Last, J. Paxton, and M. Gomon
for the loan of specimens and hospitality to DLS, A. Graham, M.
McGrouther, and T. Bardsley for their able assistance, B. Collette
for providing space and logistic support, and V. Prirodina for her
invaluable cleared and stained preparations of pectoral girdles.
Radiographs of three species were prepared by S. Raredon. All
drawings were prepared by N. Chernova except the distribution
map, prepared by M. Ford, NODC, NOAA, and the figures from
Makushok (1958). Research was supported by National Science
Foundation Grant No. DPP-8915669 to the senior author; by the
Office of Oceanic and Atmospheric Research, NOAA, and by the
Zoological Institute of the Russian Academy of Sciences. V.
Springer was instrumental in making Chernova’s USNM visit
possible. We also used the scientific collections of the Zoological
Institute, Russian Academy of Sciences, which are financially
supported by the Ministry of Science of the Russian Federation
(Registration No. 96-03-16). Chernova and Andriashev were
supported by the subprogram “Studies and observations of
Antarctica” FGP “World Ocean” Project 16, and Grants 96-15-
97881, 99-04-49774. Chernova received additional support from
the Schauer-Dieck Liparid Foundation, for which she is deeply
appreciative.
References
Able, K.W., & J.A. Musick, 1976. Life history, ecology, and
behavior of Liparis inquilinus (Pisces: Cyclopteridae)
associated with the sea scallop, Placopecten magellanicus.
Fishery Bulletin 74(2): 409-421.
Amaoka, K., K. Matsuura, T. Inada, M. Takeda, H. Hatanaka &
K. Okada, 1990. Fishes Collected by the R/V Shinkai Maru
Around New Zealand. Tokyo: Japan Marine Fishery Resource
Research Center, pp. 410.
Andriashev, A.P, 1954. Fishes of the Northern Seas of the U.S.S.R.
Fauna of the USSR 53. Izdatel’stvo Akademii Nauk SSSR
Moscow [In Russian]. Fishes of the northern seas of the
U.S.S.R. Translated 1964. Israel Program for Scientific
Translations 836: 1-617.
Andriashev, A.P, 1965. A general review of the Antarctic fish
fauna. In Biogeography and Ecology in Antarctica, ed. P. Van
Oye & J. Van Mieghem, pp. 491-550. Monographiae
Biologicae 15. The Hague: W. Junk, pp. 762.
Andriashev, A.P, 1977. Some additions to schemes of the vertical
zonation of marine bottom fauna. In Adaptations Within
Antarctic Ecosystems: Proceedings of the Third SCAR
404 Records of the Australian Museum (2001) Vol. 53
Symposium on Antarctic Biology, ed. G.A. Llano, pp. 351-
360. Washington, DC, Gulf Publishing Co., pp. 1252.
Andriashev, A.P., 1982a. A new species and a new subspecies of
Paraliparis (Liparidae) from western Antarctica. II. Voprosy
Ikhtiologii 22(2): 179-186 [in Russian. English translation in
Journal of Ichthyology 22(3): 1-9].
Andriashev, A.P., 1982b. A review of fishes of the genus
Paraliparis Collett (Liparidae) from the Kerguelen area,
Subantarctic. Zoologicheskii Zhurnal 61(5): 716-725 [in
Russian].
Andriashev, A.P, 1986. Review of the snailfish genus Paraliparis
(Scorpaeniformes: Liparididae) of the Southern Ocean. Theses
Zoologicae 7. Koenigstein, Federal Republic of Germany:
Koeltz Scientific Books, pp. 204.
Andriashev, A.P, 1991. Possible pathways of Paraliparis (Pisces:
Liparididae) and some other North Pacific secondarily deep-
sea fishes into North Atlantic and Arctic depths. Polar Biology
11:213-218.
Andriashev, A.P, 1992. Morphological evidence for the validity
of the antitropical genus Psednos Barnard (Scorpaeniformes,
Liparididae) with a description of a new species from the eastern
North Atlantic. UO 41: 1-18.
Andriashev, A.P, 1993. On the validity of the genus Psednos
Barnard (Scorpaeniformes, Liparidae) and its antitropical area.
Voprosy Ikhtiologii 33: 5-15 [in Russian].
Andriashev, A.P, 1998. A review of recent studies of Southern
Ocean Liparidae (Teleostei: Scorpaeniformes). Cybium 22(3):
255-266.
Andriashev, A.P., & A.V. Neelov, 1979. New species of the genus
Paraliparis (Liparidae) from the western Antarctic. Voprosy
Ikhtiologii 19: 10-19. [In Russian. English translation in
Journal of Ichthyology 19: 7-15].
Andriashev, A.P, A.V. Neelov & V.P. Prirodina, 1977. On methods
of study of the morphology and systematics of the fish family
of sea snails (Liparidae). Zoologicheskii Zhurnal 50: 141-147
[in Russian].
Andriashev, A.P, & V.P. Prirodina, 1990. A review of Antarctic
species of the Genus Careproctus (Liparididae) and notes on
the carcinophilic species of this genus. Voprosy Ikhtiologii 30:
709-719. [In Russian. English translation in Journal of
Ichthyology 30: 63-76].
Andriashev, A.P., & D.L. Stein, 1998. Review of the snailfish
genus Careproctus (Liparidae, Scorpaeniformes) in Antarctic
and adjacent waters. Contributions in Science of the Natural
History Museum of Los Angeles County 470: 1-63.
Anonymous, 2000. Physical, chemical, and biological data.
CalCOFI Cruise 9908, 7-29 August 1999. CalCOFI Cruise
9910, 3-10 October 1999. SIO Reference 00-10, 29 June 2000,
pp. 104.
Balbontin, F., G.I. Campodonico & M.L. Guzman, 1979.
Description de huevos y larvas de especies de Careproctus
(Pisces: Liparidae) comensales de Paralomis granulosa y
Lithodes antarctica (Crustacea: Lithodidae). Apartado Anales
del Instituto de la Patagonia 10: 235-243.
Barnard, K.H., 1927. Diagnoses of new genera and species of
South African marine fishes. Annals and Magazine of natural
History 9(XX): 66-79.
Bunt, J.S., 1987. The Australian marine environment. Ch. 2 in
Fauna of Australia. General Articles, ed. G.R. Dyne & D.W.
Walton, pp. 17-42. Canberra: Australian Government
Publishing Service. Vol. 1A, pp. 339.
Burke, C.V., 1912. Note on the Cyclogasteridae. Annals and
Magazine of Natural History Series 8, 9: 507-510.
Burke, C.V., 1930. Revision of the fishes of the family Liparidae.
Bulletin of the U.S. National Museum 150: 1-204.
Chabanaud, P, 1951. Morphologie des arcs hemaux abdominaux
des teleosteens symmetriques et dissymmetriques. Comptes
Rendus de I’Academie des Sciences de Paris, CCXXXIII:
1393-1395.
Cohen, D.M., 1968. The cyclopterid genus Paraliparis, a senior
synonym of Gymnolycodes and Eutelichthys, with the
description of a new species from the Gulf of Mexico. Copeia
2: 384-388.
Collett, R., 1878. Fiske fra Nordhavs-expeditionens sidste togt,
sommeren 1878. Forhandlinger Videnskabs-selskabet i
Christiania 1878 14: 1-106.
Conolly, J.R., 1968. Submarine canyons of the continental margin,
east Bass Strait (Australia). Marine Geology 6: 449-461.
Duhamel, G., 1992. Descriptions d’especes nouvelles de
Careproctus et Paraliparis et donnees nouvelles sur ces genres
et le genre Edentoliparis de 1’ocean austral (Cyclopteridae,
Liparinae). Cybium 16(3): 183-207.
Exon, N.F., P.J. Hill & J.-Y. Royer, 1995. New maps of crust
off Tasmania expand research possibilities. EOS 76(20): 201,
206-207.
Ford, E., 1937. Vertebral variation in teleostean fishes. Journal of
the Marine Biological Association of the United Kingdom 22( 1):
1-60.
Garman, S., 1892. The Discoboli. Memoirs of the Museum of
Comparative Zoology 14(2): 1-96.
Garman, S., 1899. The Fishes. In: Reports on an Exploration off
the West Coasts of Mexico, Central and South America, and
off the Galapagos Islands, in Charge of Alexander Agassiz, by
the U.S. Fish Commission Steamer “Albatross” During 1891.
Lieut.-Commander Z.L. Tanner, U.S.N. Commanding. Part 26.
Memoirs of the Museum of Comparative Zoology 24: 1-431,
plates 1-84.
Gilbert, C.H., 1890. A preliminary report on the fishes collected
by the steamer “Albatross” on the Pacific coast of North
America during the year 1889, with descriptions of twelve new
genera and ninety-two new species. Proceedings of the U.S.
National Museum 13(797): 49-126.
Godfrey, J.S., I.S.F. Jones, J.G.H. Maxwell & B.D. Scott, 1980. On
the winter cascade from Bass Strait into the Tasman Sea. Australian
Journal of Marine and Freshwater Research 31: 275-286.
Goode, G.B., & T.H. Bean, 1896. Oceanic Ichthyology, a treatise
on the deep-sea and pelagic fishes of the world, based chiefly
upon the collections made by the steamers “Blake”,
“Albatross”, and “Fish Hawk” in the northwestern Atlantic,
with an atlas containing 417 figures. Bulletin of the U.S.
National Museum 2: 1-553; Atlas: 123 plates.
Greenwood, P.H., 1984. What is a species flock? In Evolution of
fish species flocks, ed. A.A. Echelle & I. Kornfield, pp. 13-19,
Orono: University of Maine Press, pp. 257.
Hamilton, L.J., 1990. Temperature inversions at intermediate
depths in the Antarctic Intermediate Water of the south-western
Pacific. Australian Journal of Marine and Freshwater Research
41:325-351.
Harris, G., C. Nilsson, L. Clementson & D. Thomas, 1987. The
water masses of the east coast of Tasmania: seasonal and
interannual variability and the influence on phytoplankton
biomass and productivity. Australian Journal of Marine and
Freshwater Research 38: 569-590.
Hubbs, C.L., 1927. Notes on the blennioid fishes of western North
America. Papers of the Michigan Academy of Sciences, Arts
and Letters Nll( 1926): 351-394.
Hubbs, C.L., & K.F. Lagler, 1949. Fishes of the Great Lakes
Region. Cranbrook Institute of Science Bulletin 26: 1-186.
Huyer, A., R.L. Smith, P.J. Stabeno, J.A. Church & N.J. White,
1988. Currents of south-eastern Australia: results from the
Australian Coastal Experiment. Australian Journal of Marine
and Freshwater Research 39: 245-288.
Janssen, J., 1996. Use of the lateral line and tactile senses in feeding
in four Antarctic nototheniid fishes. Environmental Biology of
Fishes 47: 51-64.
Jordan, D.S., & B.W. Evermann, 1898. The Fishes of North and
Middle America. Bulletin of the U.S. National Museum, Part
II, V. 47: 1241-2183.
Stein et al.: Australian liparids 405
Kido, K., 1988. Phylogeny of the Family Liparididae, with the
taxonomy of the species found around Japan. Memoirs of the
Faculty of Fisheries of Hokkaido University 35: 125-256.
Kido, K., & D. Kitagawa, 1986. Development of larvae and
juveniles of Rhinoliparis barbulifer (Liparididae). In Indo-
Pacific Fish Biology: Proceedings of the Second International
Conference on Indo-Pacific Fishes, ed. T. Uyeno, R. Arai, T.
Taniuchi & K. Matsuura, pp. 697-702. Tokyo: Ichthyological
Society of Japan, pp. 985.
Kishinouye, K., 1923. Contributions to the comparative study of
the so-called Scombroid fishes. Journal of the College of
Agriculture of Imperial University, Tokyo 8: 293-475.
Koslow, J.A., C.M. Bulman & J.M. Lyle, 1994. The mid-slope
demersal fish community off southeastern Australia. Deep-Sea
Research 1,41(1): 113-141.
Krpyer, H.N., 1862. Nogle Bidrag tel Nordisk ichtyologi.
NaturhistoriskTidsskrift Kobenhavn (3 Raekke) IB: 233-310.
Last, P.R., & J.D. Stevens, 1994. Sharks and rays of Australia.
Australia: Commonwealth Scientific and Industrial Research
Organization, pp. 612.
Leviton, A.E., R.H. Gibbs Jr., E. Heal & C.E. Dawson, 1985.
Standards in herpetology and ichthyology. Part I. Standard
symbolic codes for institutional resource collections in
herpetology and ichthyology. Copeia 1985: 802-832.
Makushok, V.M., 1958. The morphology and classification of the
northern blennioid fishes (Stichaeoidae, Blennioidei, Pisces).
Trudy Zoologicheskogo Instituta Akademia Nauk SSSR 25: 3-
129 [in Russian].
Matallanas, J., 1999. New and rare snailfish genus Paraliparis
from the Weddell Sea with the description of two new species.
Journal of Fish Biology 54: 1017-1028.
Matsubara, K., & T. Iwai, 1954. Some remarks on the family
Liparidae with description of three new species and two
interesting ones of the genus Liparis. Report of the Faculty of
Fisheries of the University of Mie 1: 425^-41.
May, J.L., & S.J.M. Blaber, 1989. Benthic and pelagic fish biomass
of the upper continental slope off eastern Tasmania. Marine
Biology 101: 11-25.
Mead, G.W., 1970. A history of South Pacific fishes. In Scientific
Exploration of the South Pacific, ed. W.S. Wooster, pp. 236-251.
Washington, D.C.: National Academy of Sciences, pp. 257.
Peden, A.E., & C.A. Corbett, 1973. Commensalism between a
liparid fish, Careproctus sp., and the lithodid box crab,
Lopholithodes foraminatus. Canadian Journal of Zoology
51(5): 555-556.
Pianka, E.R., 2000. Ctenotus web page: Phylogenetic analysis of
a major adaptive radiation. Motivation.
http ://uts. cc .utexas. edu/~ varanus/ctenotus .html
Rao, T.S.S., & R.C. Griffiths, 1998. Understanding the Indian Ocean.
Perspectives on oceanography. Paris: UNESCO, pp. 187.
Ribbink, A.J., 1984. Is the species flock concept tenable? In
Evolution offish species flocks, ed. A.A. Echelle & I. Komfield,
pp. 21-25. Orono: University of Maine Press, pp. 257.
Richards, W.J., 1966. Paraliparis wilsoni, a new liparid fish from
the Gulf of Guinea. Proceedings of the Biological Society of
Washington! 9: 171-174.
Rochford, D.J., 1975. The physical setting. In Resources of the
Sea, ed. M.R. Banks & T.G. Dix, pp. 15-27. Hobart: Royal
Society of Tasmania, pp. 119.
Smith, J.L.B., 1967. Anew liparine fish from the Red Sea. Journal
of Natural History 2: 105-109.
Soldatov, V.K., & G.U. Lindberg, 1930. A Review of the Fishes
of the Seas of the Far East. Izvestiya Tikhookeanskogo
Nauchnogo Instituta Rybnogo Khozyaistva 5: 1-576.
Stein, D.L., 1978a. The genus Psednos a junior synonym of
Paraliparis, with a redescription of Paraliparis micrurus
(Barnard) (Scorpaeniformes: Liparidae). Matsya 4: 5-10.
Stein, D.L., 1978b. A review of the deepwater Liparidae (Pisces)
from the coast of Oregon and adjacent waters. Occasional
Papers of the Califomia Academy of Sciences 127: 1-55.
Stein, D.L., 1980. Aspects of reproduction of liparid fishes from
the continental slope and abyssal plain off Oregon, with notes
on growth. Copeia 1980: 687-699.
Stein, D.L., 1986. Family Liparididae. In Smiths’ Sea Fishes, ed.
M.M. Smith & PC. Heemstra, pp. 492-494. Grahamstown,
South Africa: J.L.B. Smith Institute of Ichthyology, pp. 1047.
Stein, D.L., & A.P. Andriashev, 1990. Family Liparididae. In
Fishes of the Southern Ocean, ed. O. Gon & PC. Heemstra,
pp. 231-255. Grahamstown, South Africa: J.L.B. Smith
Institute of Ichthyology, pp. 462.
Stein, D.L., & J.E. Fitch, 1984. Paraliparis nassarum n. sp.
(Pisces, Liparididae) from off southern California with
description of its otoliths and others from north-east Pacific
liparidids. Bulletin of the southern California Academy of
Sciences 83(pt 2): 76-83.
Stein, D.L., R. Melendez C., & I. Kong U., 1991. A review of
Chilean snailfishes (Liparididae, Scorpaeniformes) with
descriptions of a new genus and three new species. Copeia
1991:358-373.
Stein, D.L., & L.S. Tompkins, 1989. New species and new records
of rare Antarctic Paraliparis fishes (Scorpaeniformes:
Liparididae). Ichthyological Bulletin of the J.L.B. Smith
Institute of Ichthyology 53: 1-8.
Tchernia, R, 1980. Descriptive Regional Oceanography. New
York: Pergamon Press, pp. 253.
Williams, A., PR. Last, M.F. Gomon & J.R. Paxton, 1996. Species
composition and checklist of the demersal ichthyofauna of the
continental slope off Western Australia (20-35°S). Records of
the Western Australian Museum 18: 135-155.
Williams, D.F., & B.H. Corliss, 1982. The South Australian
continental margin and the Australian-Antarctic sector of the
Southern Ocean. Ch. 3. In The Ocean Basins and Margins.
Vol. 6. The Indian Ocean, ed. A.E.M. Nairn & F.G. Stehli, pp.
545-584. New York: Plenum Press, pp. 776.
Wilson, B.R., & G.R. Allen, 1987. Major components and
distribution of marine fauna. Ch. 3 in Fauna of Australia.
General Articles, ed. G.R. Dyne & D.W. Walton, pp. 43-68.
Canberra: Australian Government Publishing Service. Vol. 1 A,
pp. 339.
Yabe, M., 1985. Comparative osteology and myology of the
superfamily Cottoidea (Pisces: Scorpaeniformes), and its
phylogenetic classification. Memoirs of the Faculty of
Fisheries, Hokkaido University 32(1): 1-130.
Manuscript received 10 July 2000, revised 23 February 2001 and accepted
8 March 2001.
Associate Editor: J.M. Leis.
406 Records of the Australian Museum (2001) Vol. 53
Contents
Key to genera of Australian liparids.353
Genus Careproctus Krpyer, 1862 . 353
Careproctus paxtoni n.sp. 353
Genus Psednos Barnard, 1927. 355
Key to Southern Hemisphere Psednos . 356
Psednos balushkini n.sp.356
Psednos nataliae n.sp. Stein & Andriashev.357
Psednos whitleyi n.sp.359
Psednos sp.360
Genus Paraliparis Collett, 1878. 360
Key to Australian Paraliparis . 360
Paraliparis anthracinus n.sp.364
Paraliparis ater n.sp. 365
Paraliparis atrolabiatus n.sp.366
Paraliparis auriculatus n.sp.367
Paraliparis australiensis n.sp.369
Paraliparis avellaneus n.sp.370
Paraliparis badius n.sp. 371
Paraliparis brunneocaudatus n.sp. 372
Paraliparis brunneus n.sp. 374
Paraliparis coracinus n.sp.375
Paraliparis costatus n.sp. 376
Paraliparis csiroi n.sp. 378
Paraliparis delphis n.sp.379
Paraliparis dewitti n.sp.381
Paraliparis eastmani n.sp.382
Paraliparis gomoni n.sp.383
Paraliparis hobarti n.sp. 384
Paraliparis impariporus n.sp. 386
Paraliparis infeliciter n.sp.387
Paraliparis labiatus n.sp.388
Paraliparis lasti n.sp. 389
Paraliparis obtusirostris n.sp. 391
Paraliparis piceus n.sp. 392
Paraliparis plagiostomus n.sp.393
Paraliparis retrodorsalis n.sp. 394
Paraliparis tasmaniensis n.sp.396
Paraliparis sp. 1. 397
Paraliparis sp. 2. 398
Incertae sedis.399
Paraliparis sp. (cf. copei group). 399
© Copyright Australian Museum, 2001
Records of the Australian Museum (2001) Vol. 53: 407-425. ISSN 0067-1975
Larvae and Juveniles of the Deepsea
“Whalefishes” Barbourisia and Rondeletia
(Stephanoberyciformes: Barbourisiidae, Rondeletiidae),
with Comments on Family Relationships
John R. Paxton, 1 G. David Johnson 2 and Thomas Trnski 1
1 Fish Section, Australian Museum,
6 College Street, Sydney NSW 2010, Australia
j ohnp @ au s tmu s. g o v. au
tomt @ au stmu s. go v. au
2 Fish Division, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
j ohns on. dave @ nmnh. si. edu
Abstract. Larvae of the deepsea “whalefishes” Barbourisia rufa (11: 3.7-14.1 mm nl/sl) and Rondeletia
spp. (9: 3.5-9.7 mm si) occur at least in the upper 200 m of the open ocean, with some specimens taken
in the upper 20 m. Larvae of both families are highly precocious, with identifiable features in each by
3.7 mm. Larval Barbourisia have an elongate fourth pelvic ray with dark pigment basally, notochord
flexion occurs between 6.5 and 7.5 mm si, and by 7.5 mm si the body is covered with small, non-
imbricate scales with a central spine typical of the adult. In Rondeletia notochord flexion occurs at
about 3.5 mm si and the elongate pelvic rays 2-4 are the most strongly pigmented part of the larvae.
Cycloid scales (here reported in the family for the first time) are developing by 7 mm; these scales later
migrate to form a layer directly over the muscles underneath the dermis. By 7 mm si there is a unique
organ, here termed Tominaga’s organ, separate from and below the nasal rosette, developing anterior to
the eye. Larvae of the two species of Rondeletia can be distinguished by the presence or absence of
developing spongy bone in the pectoral girdle and sphenotic by at least 9 mm and by the counts of the
vertebrae, pelvic-fin rays, and dorsal hypural bones in smaller larvae. The presence of Tominaga’s
organ in the gibberichthyid Gibberichthys suggests that “the whalefishes”, Barbourisiidae, Rondeletiidae,
and Cetomimidae, as a group are paraphyletic, and that Rondeletia and Gibberichthys are sister taxa.
Paxton, John R., G. David Johnson & Thomas Trnski, 2001. Larvae and juveniles of the deepsea “whalefishes”
Barbourisia and Rondeletia (Stephanoberyciformes: Barbourisiidae, Rondeletiidae), with comments on family
relationships. Records of the Australian Museum 53(3): 407^-25.
The deepsea “whalefish” families Rondeletiidae and
Barbourisiidae have been considered close relatives since
the description of the latter family by Parr (1945). Recent
authors have considered them part of a separate order
Cetomimiformes (Ebeling & Weed, 1973), part of a
“stephanoberycoid assemblage” (Rosen, 1973) or part of a
suborder of the Beryciformes (Rosen & Patterson, 1969;
Keene & Tighe, 1984; Moore, 1993). We follow Johnson &
Patterson (1993) and Nelson (1994) in recognizing two
orders: Stephanoberyciformes (Melamphaidae, Stephano-
408 Records of the Australian Museum (2001) Vol. 53
berycidae, Hispidoberycidae, Gibberichthyidae, Rondel-
etiidae, Barbourisiidae, Cetomimidae, Megalomycteridae,
Mirapinnidae) and Beryciformes (Holocentridae, Beryc¬
idae, Diretmidae, Anoplogastridae, Trachichthyidae,
Anomalopidae, Monocentridae), respectively sequential
sister groups to the Percomorpha. Recently Colgan et al.
(2000) questioned the monophyly of the Stephano-
beryciformes based on partial 12S and 16S rDNA sequences.
Further consideration of family relationships within the
Stephanoberyciformes is in the Discussion.
In their description of a 6.2 mm larval specimen of the
anomalopid Kryptophanaron, Baldwin & Johnson (1995)
reported that larvae of 10 of the 16 recognized stephano-
beryciform and beryciform families had been described.
They also noted that larval specimens of two additional
families, Rondeletiidae and Barbourisiidae, had been
identified from collections. The purpose of this paper is to
describe those specimens and to comment on family
relationships based on the larval characters.
Boehlert & Mundy (1992) described an 11.3 mm larva
from near Hawaii that they tentatively placed in the
Stephanoberycidae as either Malacosarcus or an undescribed
form. Body shape, meristics, and the lack of scales at that
size preclude identification as either Barbourisia or
Rondeletia.
The family Barbourisiidae is monotypic. Barbourisia
rufa was described by Parr (1945) from the Gulf of Mexico.
The species has since been collected from the Atlantic,
Pacific and Indian Oceans from >60°N to 45° S; at least
100 specimens have been collected (Kotlyar, 1995; Paxton,
unpubl.). Captures have been with both benthic nets between
350 and 1500 m and pelagic nets to at least 800-2000 m.
Barbourisia rufa attains 390 mm SL and the sexes are
separate (Paxton, unpubl.). Struhsaker (1965) figured the
distinctive scales, and osteological features of the gill arches
and caudal skeleton were described by Rosen (1973).
Ebeling & Weed (1973) also summarized selected features
of Barbourisia. In his phylogenetic analysis of the
“trachichthyiform” fishes Moore (1993) coded 25
osteological characters for Barbourisia. Johnson &
Patterson (1993) discussed cranial sensory features and
other selected aspects of the osteology, including the
intermusculars (also discussed and tabulated by Patterson
& Johnson, 1995). Kotlyar (1995) described and figured
the osteology, based primarily on a cleared and stained
specimen 212 mm si. Colgan et al. (2000) detailed partial
sequences of 12S and 16S rDNA for the species.
The Rondeletiidae includes Rondeletia bicolor Goode
& Bean (1895) and R. loricata Abe & Hotta (1963). Parr
(1929) described the osteology of R. bicolor, and Paxton
(1974) described that of R. loricata and summarized
distributional data for both species. Selected osteological
features have been described by Ebeling & Weed (1973),
Rosen (1973), Moore (1993), Johnson & Patterson (1993),
and Patterson & Johnson (1995). Bast & Klinkhardt (1990)
described specimens of R. loricata from the northeast and
southwest Atlantic. Kotlyar (1996) detailed the osteology
of R. loricata with many illustrations, and analysed the
distributions of both species. Colgan et al. (2000) detailed
partial sequences of 12S and 16S rDNA of R. loricata. The
species are meso- and perhaps bathypelagic, with captures
from 250-2000 m in open nets. Rondeletia loricata occurs
between 58°N and 48°S in all three oceans. Rondeletia
bicolor is most common in the Caribbean and western North
Atlantic between 0° and 37°N, with only one record from
the South Atlantic and two records from the South Pacific
(Paxton, 1974; unpublished). Maximum size of the genus
is 113 mm si.
Materials and methods
Institutional abbreviations follow Leviton et al. (1985). TH
is the Tokai Regional Fishery Research Laboratory, Tokyo,
the specimens of which have recently been transferred to
NSMT. Standard length = si; notochord length = nl. The
abbreviations of measurements follow Paxton (1989: 139);
P2 = pelvic fin. All measurements are in si and mm unless
otherwise indicated. Most of the larvae were found in the
Dana Collections at ZMUC (Table 1); the fishing depths
are estimated to be one third the amount of wire out
(Bertelsen, 1951:198). Most of the juveniles are from MCZ.
All measurements of larvae were made with an ocular
micrometer in a dissecting microscope. Measurements of
juveniles and adults were made with dial calipers. Meristics
of adults are mostly from xrays. Selected specimens were
stained with alcian blue for cartilage and/or alizarin for bone.
Identifications
Identification of larval Barbourisia rufa was based on the
presence of non-imbricate scales with a central spine
(Struhsaker, 1965: fig. 1) and abdominal pelvic fins, both
characteristic of adults, and was confirmed with comparative
meristics of the other families in the orders (Keene & Tighe,
1984). Adult Acanthochaenus, Hispidoberyx, and Stephano-
beryx have similar but fewer and much larger spiny scales;
their vertebral count of 30-34 (Keene & Tighe, 1984; Yang
et al., 1988) differs from the 40-44 vertebrae of Barbourisia.
Identification of larval Rondeletia (3.5-9.7 mm) was
based on fin-ray and vertebral counts and abdominal pelvic
fins. Smaller larvae were distinguished by pelvic-fin and
vertebral counts: 6 and 26-27, respectively, in R. bicolor, 5
and 24-26, rarely 27 in R. loricata (Paxton, 1974). The
largest larvae and small juveniles (over 8.5 mm) were
identified to species by the presence (R. loricata) or absence
(R. bicolor) of spongy, honeycomb-like ossifications of the
main bones of the pectoral girdle, with posterior extensions
on the posttemporal dorsally and cleithrum ventrally. This
was facilitated by comparison of the larvae with a series of
juvenile specimens (12.6 to 21.7 mm) that are recognizable
by adult features such as vertical rows of lateral-line
neuromasts, abdominal pelvic fins and brown colour.
No distinct metamorphosis from larval to juvenile stage
is present in either family, rather a gradual transition occurs.
We have arbitrarily chosen the completion of the adult
condition of the lateral-line system on the body to
distinguish larvae from juveniles. In Barbourisia the largest
larva at 14.1 mm has enlarged scales in an open lateral-line
trough, while the smallest juvenile at 30.0 mm has the
enlarged scales within a closed lateral-line canal. In
Rondeletia, lateral-line head pores and vertical rows of
papillate superficial neuromasts are visible in a 12.6 mm R.
loricata and a 14.4 mm R. bicolor, but not in a 9.7 mm R.
loricata considered the largest larva. The 13.5 mm R. bicolor
lacks visible features of the lateral-line system, but is
completely faded and in poor condition. Based on
Paxton et al .: Barbourisia and Rondeletia larvae 409
Table 1. Barbourisia rufa material examined. Abbreviations: cl, closing net; * = cleared and stained; # = drawn.
specimen
catalogue
size (mm)
location
depth (m)
day/night
date
1
AMS 129035-003
3.7
14°40'S 145°15'W
0-10
D
31 Jan 1989
2 #
AMS 129176-002
4.8
14°56'S 147°52'W
0-5
D
14 Feb 1989
3
USNM 363086
4.9
21°32'N 157°45'W
0-0.7
N
14 Dec 1985
4
AMS 124586-007
5.0
21°16'N 157°32'W
0-1
?
13 Jun 1972
5 *
ZMUC P2340802
6.2
1°15'N 136°07'E
0-33
N
14 Jul 1929
6 #
AMS 129174-002
6.6
14°56'S 147°52'W
0-5
D
14 Feb 1989
7 #
MCZ 75627
7.5
2°06'N 33°38'W
0-70
?
16 Mar 1977
8 *
USNM 363087
10.0
33°59'N 76°22'W
0-63
?
15 Sep 1994
9
USNM 305035
13.1
19°25'N 156°18'W
0-50
N
27 Sep 1988
10 *
ZMUC P2340803
13.4
10°51'S 168°40'W
0-33
N
29 Oct 1928
11 *#
ZMUC P2340804
14.1
15°56'S 172°30'W
0-66
N
7 Nov 1928
12
AMS 118823-001
30.0
21°25'N 158°25'W
825-1150 cl
17 Mar 1971
13
SIO 88-172
34.6
6°55'N 177°48'W
14 Mar 1987
14
TH 865522
45.6
29°59'N 134°H'E
0-1040
18 Jul 1986
15
AMS 126869-001
89
off Zanzibar
0-200
1965
16
AMS 127260-001
92
21°23'N 158°18'W
17 Jun 1973
17 *
AMS 118824-001
100
25°25'N 158°25'W
250-300 cl
N
23 Apr 1971
18
AMS 122812-001
114
18°08'S 116°43'E
0-800
5 Apr 1982
19 *
AMS 127261-001
133
0°08'N 154°02'W
?
2 Mar 1969
similarities of body shape and fin development with the
larger juveniles, it is assumed to be the smallest known
juvenile. Head pores and papillate neuromasts are visible
in some, but not all, of the juveniles of both species less
than 20 mm si depending on their skin condition.
Results
Barbourisia rufa Parr, 1945
Fig. 1
Eleven larvae 3.7-14.1 mm were examined, six preflexion
specimens 3.7-6.6 mm and five postflexion specimens 7.5-
14.1 mm (Table 1). The three ZMUC specimens are faded
and transparent, having been stored for decades in formalin,
and have now been stained with alizarin. The 13.1 mm
USNM specimen retains pigment, but unfortunately had
the pelvic fins removed, apparently by an overzealous
plankton sorter attempting to “clean” the specimen. In
the three largest specimens >13 mm si, the skin is inflated,
loose and balloon-like around the body, and appears to
have little connection to the underlying muscle. The four
smallest larvae (5.0 mm nl and smaller) are very slender,
distinguished by long, abdominal pelvic fins. The body is
deeper anteriorly and slender posteriorly in the two largest
preflexion specimens >6 mm nl, moderately deep in the
smallest postflexion specimen 7.5 mm si, and deep and
globose in the four largest postflexion specimens. The jaws
are relatively short and obliquely directed in all five
postflexion specimens (Fig. 1).
In the larvae, eye diameter, pectoral- and pelvic-fin
lengths and body depth (only in postflexion specimens) are
relatively greater than those of the juveniles and adults,
while the snout length is less. Nostrils are visible only as
small slits anterior to the eye in the largest larvae, although
a small nasal pit becomes apparent at 6.6 mm nl. The
distinctive elongate pelvic fin reaches the equivalent of
beyond the hypurals by 6.6 mm nl and at least the 4th ray
retains this relative length at least until the 14.1 mm larvae,
while the pelvic-fin insertion changes from closer to the
pectoral-fin origin to closer to the anal-fin origin through
the same size range (Fig. lb,d). The growth of some
elements (pectoral and pelvic fins) becomes isometric by
small juvenile size (30 mm), but the small juvenile
specimens are not in good enough condition to determine
if the 4th fin ray is longer than the other rays. The eye
diameter becomes isometric only at more than 150 mm si.
Variation in other measurements (e.g., pectoral-fin origin
to anus and body depth) is due to the flabby nature of the
specimens resulting in imprecise measurements. A
comparison of the larval shape of Barbourisia rufa (Fig.
Id) with that of the adult (Rofen, 1959: fig. 3) shows the
striking transformation in head shape, snout length, and jaw
angle. By 30 mm the shapes of these elements are similar
to those of the adult.
Pigmentation. Many of the larval specimens are faded. The
eye is solid black, except for the white lens. There are two
layers of melanophores over part of the head and anterior
half of the body by 13 mm when the skin separates from
the body; a layer of larger, lighter and more widely spaced
melanophores just under the skin and a deeper layer
overlying the viscera and part of the brain.
Small, evenly-distributed melanophores are present
dorsally and dorsolaterally on the entire head in the smallest
specimen. The density and size of melanophores vary as
the larvae develop. However in larger preflexion larvae,
melanophores tend to be stellate and more densely arranged
over the brain. In the largest specimen superficial
melanophores are present circumorbitally, on the cheek and
the upper half of the opercles. More closely spaced
melanophores are on the top of the head in the supra-
occipital-posterior frontal area. A deeper layer of darker,
more widely spaced melanophores covers the visible
lobes of the brain above and behind the orbit. A few
melanophores are present on the lower jaws throughout
larval development. The small melanophores on the dorsal
surface of the head extend posteriorly to the nape and are
distributed over the entire musculature of the trunk and tail.
410
Records of the Australian Museum (2001) Vol. 53
Figure 1. Barbourisia rufa larvae, a, AMS 129176-002,4.8 mm nl; b, AMS 129174-002, 6.6 mm nl, spinules shown
in profile only, with distribution indicated by dashed line across posterior of gut and anterior of tail; c, MCZ 75627,
7.5 mm si, note missing posterior of gut; d, ZMUC P2340804, 14.1 mm si, with enlargement of left lateral-line
scale no. 20 and adjacent body scales illustrated below caudal fin.
Paxton et al .: Barbourisia and Rondeletia larvae 411
The melanophores on the notochord tip are restricted to the
dorsal and ventral margins. In postflexion larvae, small
melanophores are distributed fairly evenly throughout the
loose, balloon-like skin that covers the body, except over
the abdominal cavity, where they extend ventrally only
about to the level of the pectoral fin. Beneath the loose skin,
the wall of the abdominal cavity is evenly covered with
somewhat larger, more closely-spaced melanophores. This
internal layer of peritoneal pigment extends almost to the
ventral margin of the body in the region behind the pelvic
girdle. Small melanophores are also found beneath the loose
skin on the epaxial musculature, where they are more
sparsely distributed and tend to concentrate along, and thus
delineate, the myosepta and horizontal septum. The dorsal,
anal, caudal and pectoral fins are unpigmented, except for
two melanophores on each side of the base of the dorsal-
most principal caudal ray only in the largest (14.1 mm)
specimen. Pelvic fins are unpigmented proximally. Small
melanophores cover both the membranes and elements, and
extend to the tips of the pelvic fins.
The three 30-46 mm juvenile specimens are faded white
in preservative and presumably were the original red-orange
colour of adults. The carotenoid pigment is alcohol soluble
and is bleached in preservation (Herring, 1976). In the
largest two specimens (the smallest is completely faded)
melanophores overlying the brain and muscle mass show
through the skin. Those on the body are in two layers, one
in a transparent sheet of tissue under the skin with light
streaks of pigment, and another of lighter, more widely
scattered streaks closely associated with the muscle bands.
The peritoneum is solidly pigmented black. The basal half
of the fourth pelvic-fin ray has large dark melanophores.
By 89-114 mm, the two layers of pigment over the
musculature are light but distinct, and a single large pigment
spot remains near the base of the fourth pelvic-fin ray under
the skin; scattered light melanophores are on the basal
portions of all pelvic-fin rays in the least faded specimen.
The posterior half of the medial side of the gill cover has a
layer of moderately dense melanophores. In fresh specimens
over 300 mm si, the sheet of tissue between the skin and
muscles is pigmented with brown blotches and a single large
black spot is visible after dissection at the base of the fourth
pelvic-fin ray. The inside of the gill cover is solid black.
Scales. Scales are present in the smallest specimen. They
are small, round and non-imbricate with a single, central
spine and appear identical in form to the adult scale
illustrated by Struhsaker (1965). They are restricted to the
dorsal surface of the trunk with at most 3-4 longitudinal
rows of scales. This dorsal shield spreads in all directions;
by late preflexion the scales extend over the trunk and tail
from the nape to just beyond the anus, over the anterior and
middle of the gut, but not onto the posterior-most portion
of the gut. Two small patches are also present on the opercle.
Scales develop progressively more posterior on the tail in
postflexion larvae, and become more extensively distributed
on the head. In the smallest postflexion specimen scales
cover the tail except for the caudal peduncle. A few rows of
scales extend over the base of the anterior-most dorsal-fin
rays. The preopercle and opercle are almost entirely covered,
and scales are also present postorbitally and on the maxilla.
In the largest larva the scales cover the entire trunk and tail,
and extend forward to cover much of the head, with the
exception of the lower jaw, snout, premaxilla and anterior
portion of the maxilla, some aspects of the frontals, the
anterior three infraorbitals, and posterior surface of the
preopercle. The gular region is scaleless anterior to the
cleithral symphysis, but there is an elongate median patch
6-7 scales wide in the 13.4 mm specimen behind the lower
jaw symphysis. An envelope of scale-bearing skin extends
about Vi of the way out the dorsal-and anal-fin rays of the
three largest specimens. In the 30 mm juvenile and larger
specimens scales extend to the tip of the snout and to the
tips of all the fin rays.
In the preflexion specimens specialized lateral-line scales
are not apparent. We cannot ascertain whether the lateral¬
line trough is scaleless at this stage or whether it is covered
with small spined scales. We do not expect that the
specialized lateral-line scales will transform from body
scales. The two smallest postflexion specimens are
somewhat damaged, and the rippled skin makes scale
distribution difficult to observe. In the three 13.1-14.1 mm
postflexion specimens enlarged scales extend along the
lateral line from the head to the base of the caudal fin, with
one good count of 33 scales. These scales are 2-4 times the
size of the body scales, have a central foramen and four
spines, two dorsal and two ventral (Fig. Id). The small body
scales are absent between adjacent lateral-line scales. In 30
and 34 mm juvenile specimens the lateral line has
invaginated to form a canal that is overgrown with skin.
The overlying skin is pierced by small pores, but is only
partially covered with small body scales in a series of narrow
strips between the pores. Enlarged lateral-line scales, each
with a central foramen, lie in the bottom of the lateral-line
canal. No enlarged spines remain, but each scale has dorsal
and ventral extensions that run laterally along the walls of
the lateral-line canal. Each extension consists of two narrow
elements that may represent the four spines present on each
scale in the larvae. There is a neuromast on each lateral¬
line scale, innervated by a branch of the lateral-line nerve
that emerges through the foramen of each scale. In a 133
mm specimen the dorsal and ventral extensions of the
lateral-line scales extend further laterally and support
approximately half of the roof of the canal. Each pair of
extensions is strengthened by a series of small cross struts
(Paxton, 1989: fig. 5a).
Head spines. No head spines are developed in the preflexion
specimens, and the infraorbitals are unossified. In the
smallest postflexion specimen the orbital rims of all six
infraorbitals (including the dermosphenotic) bear small
spines that may be on body scales. In the largest postflexion
specimen the ventral rim of each infraorbital also has a
single row of small spines. The interopercle bears 4-5 spines
along its ventral margin that may also be scale spines. The
preopercle has 2 small spines on the lateral surface and 2
small spines on the posterior margin in the smallest
postflexion larva. None of the other opercular or pectoral-
series bones bear spines. The supraorbital ridge is serrate
in the smallest postflexion larva. From 13.1 mm the
supraorbital edge of the frontal bears several longitudinal
ridges each with one or two spines resulting in a triangular¬
shaped cluster of spines, medial to which are two transverse
serrate ridges of bone forming walls for a portion of the
supraorbital commissure of the lateralis system. A narrow
upright bony strut lies medial to the anterior-most ridge.
412 Records of the Australian Museum (2001) Vol. 53
There is a single extrascapular anterior to the posttemporal
in all postflexion larvae, with the slightly raised anterior
margin bearing several spines. The nasal bone has a single
minute spine on the lateral rim in the postflexion larvae,
and there are several small spines on the ventrolateral surface
of the supramaxilla that are scale spines. There is a low
ridge with 1-2 small spines laterally on the dentary in
postflexion larvae only.
Fin formation. In the second largest preflexion specimen
6.2 mm nl, the median fins appear to be developing in both
anterior and posterior directions. There are about 18 dorsal-
and 14 anal-fin bases and approximately 14 and 11 incipient
rays, respectively (Table 2). The caudal fin has about five
dorsal and eight ventral rays. The dorsal-most pectoral-fin
rays have begun to differentiate in the largest preflexion
specimen. The pelvic-fin origin is initially slightly closer
to the head than the anus. It has four well-developed rays
in the smallest specimen, five rays by 4.8 mm and six rays
by 6.2 mm. The fourth ray is produced and up to 50% longer
than the other longest rays, but is often broken. The pelvic
fins are initially close to each other and the ventral body
margin. In postflexion larvae the pelvic fins are widely
separated from one another and located higher on the body
than in the preflexion larvae. By 30 mm and larger, the
pelvic fin is much shorter, ending far forward of the anal-
fin origin, and closer to the ventral margin of the body. All
four postflexion specimens have full fin-ray complements
in all fins (although the pelvic fins are missing in the largest
specimen). Only the three cleared and stained postflexion
larvae have visible supraneurals, with six or seven present.
Dentition. A single row of small triangular teeth is apparent
on the premaxilla and dentary of the 6.2 mm larva. By 6.6
mm, the premaxilla and dentary have two rows of widely
spaced, small, triangular teeth. Teeth increase in number as
larvae develop. By 13 mm the teeth have become conical-
triangular and are closely set in two rows, and by 34 mm
the teeth have the adult form of a broad band of small conical
teeth with about six tooth rows across the band. At 100 and
133 mm the teeth have a slightly enlarged tip, are depressible
orally and the largest teeth are in the inner row. With
increasing specimen size, the number of teeth across the
jaw increases.
Internal anatomy. In smaller preflexion specimens the gut
is narrow and folded anteriorly and straightens before
exiting near the anal-fin origin. In larger preflexion larvae
the gut is thick and folded. In the two larger ZMUC
specimens the stomach is obscured by the liver; the intestine
is considerably folded with a short straight section directed
posteroventrally to the anus. A small swimbladder is evident
in the smallest specimen, and is visible in larger specimens
until the skin thickens. In the postflexion specimens it is
present under the kidneys and extends as a space over the
intestine. The swimbladder is regressed in adult Barbourisia
(Bertelsen & Marshall, 1984: 382).
Caudal skeleton. In the 6.2 mm preflexion larva the
parhypural and at least 4 hypurals are evident on the ventral
side of the notochord posteriorly. The last several centra
are not yet fully formed. In the two largest cleared
postflexion specimens there are three epurals, two
uroneurals, two urostylar centra (the compound PU1-U1
and a separate U2), one parhypural and six hypurals. The
first epural originates over the posterior edge of the neural
crest of PU2. The parhypural and hypurals 1 and 2 articulate
with an oblong block of cartilage lying along the ventral
surface of PU1-U1. Hypurals 3 and 4 articulate with U2. In
the 100 and 133 mm specimens hypural 3 articulates with
both the base of U2 and the cartilage anterior to that centrum.
The bases of hypurals 3 and 4 are in close contact with U2,
but not fused to it. The cartilage between hypurals 2 and 3
remains unossified in the larvae and the 100 mm specimen
Table 2. Barbourisia rufa counts. Abbreviations: A, anal-fin rays; Cprin, principal caudal-fin rays; Cproc, procurrent
caudal-fin rays; D, dorsal-fin rays; LL, lateral line; Myom, myomeres; P, pectoral-fin rays; P2, pelvic-fin rays;
Supran, supraneural elements; Vert, vertebrae; t fin bases only; $ fins removed; horizontal broken line indicates
limit of preflexion and postflexion specimens; solid line indicates limit of larvae and juveniles; others as in Table 1.
specimen
size
D
A
P
P2
Cprin
Cproc
Supran
Myom/Vert
LL scales
1
3.7
_
_
_
4
_
_
41
2
4.8
—
—
—
5
—
—
42
3
4.9
—
—
—
5
—
—
42
4
5.0
—
—
—
5
—
—
42
5 *
6.2
c. 18
c. 14
—
6
c. 5+8
—
42-43
6
6.6
20 t
16 t
—
6
5+5
—
41
7
7.5
21
17
12+
6
10+9
2+3
8 *
10.0
20
17
12+
6
10+9
9+8
7
42
9
13.1
22
18
13
$
10+9
9+8
10 *
13.4
22
17
14
6
10+?
9+?
6
42
11 *
14.1
21
17
13
6
10+9
10+9
6
42
33
12-14
30.1-47.3
21
16
12-13
6
41-43
25/26
15-23
88.0-169
19-22
16-18
12-14
6
41-42
28-34
24-36
250-305
20-21
16-17
12-14
6
41-43
28-34
37-48
318-386
20-23
16-18
12-14
6
40^13
25-34
Paxton et al .: Barbourisia and Rondeletia larvae 413
and is about one-third ossified in the 133 mm specimen. In
these larger specimens the first epural originates over the
anterior end of PU2.
Rosen (1973: 492) described the caudal skeleton of
Barbourisia as sharing with Rondeletia and the cetomimids
a “complex joint of the upper hypurals with a cartilaginous
plug on the hinder end of the compound centrum”, but his
figure 120 of Barbourisia shows no cartilage in this region
and shows hypural 4 fused with the second ural centrum.
We have examined Rosen’s specimen and find both his
description and his illustration to be in error. There is no
exposed cartilage plug joint and although hypural 4
articulates tightly with PU2, it is not fused to it. We place
little significance on the presence or absence of Rosen’s
so-called cartilage plug, as it is a general pattern in teleost
fishes for the parhypural and hypurals 1 and 2 to develop
together along a single block of cartilage ventral to PU1-
U1 (Potthoff & Tellock, 1993; GDJ, pers. observ.). The
degree of exposure of their cartilage in adults is merely a
function of the extent of ossification of the bases of the
proximal portions of the three elements. Thus, the “cartilage
plug” is large and well exposed in larval Barbourisia and
juveniles, but by 133 mm is almost fully covered by the
ossified bases of the parhypural and hypurals 1 and 2.
Distribution. The 11 larvae (6 preflexion, 5 postflexion)
are distributed as follows: Pacific—Hawaii 2, 1; Tuamotus
3,0; Samoa 0, 2; Indonesia 1,0; Atlantic—USAO, 1; Brazil
0, 1 (Table 1, Fig. 2). The species is now known from all
oceans, with adult specimens from 65°N to 40°S in the
Atlantic, 50°N to 50°S in the Pacific, and 5-20°S in the
Indian Ocean (Kotlyar, 1995; Paxton, unpublished). All
larvae were caught with open nets, fishing from the surface
to a maximum depth of 70 m (Table 1).
Five of the six preflexion larvae were caught in the upper
10 m, with two of these caught at one m or less. All five
postflexion larvae were caught in nets fishing to at least 33 m.
A. S. Oil'd
■ R. ttfootof
1H?
# R loricata
1B0
Figure 2. Geographic distribution of larval Barbourisia rufa,
Rondeletia bicolor, and R. loricata, symbols may represent more
than one specimen.
Rondeletia bicolor Goode & Bean, 1895
Figs. 3, 4, 8
Only one larva 7.2 mm si is known; it is postflexion.
However, the next smallest specimen, 13.5 mm si, was
originally assumed to be a larva; it lacks pigment and has
been cleared and stained (Fig. 3). It is now considered to be
the smallest known juvenile, based on its similarity of shape
and fin formation to the next largest specimens, and the
differences in larvae and juveniles of R. loricata of similar
sizes (see Identification section above). Many of the features
of this smallest juvenile are included in the larval description
below. Four specimens 14.4-21.7 mm have the loose,
uniformly dark brown skin of adults and the smallest has
clearly developed head pores and vertical rows of papillate
superficial neuromasts of the lateral-line system; they are
here considered juveniles. Both of the smallest specimens
have been cleared and stained and the amount of connective
tissue is not apparent. In the second smallest juvenile,
considerable fibrous connective tissue is present between
the skin and muscle mass, as is typical of adults. The head
and body of the two smallest specimens are moderately
deep, with the body particularly short in the larva. The tail
region is more slender in the smallest juvenile. The jaws
are relatively short and directed obliquely in the larva and
two smallest juveniles. In a 17 mm juvenile the jaws have
lengthened to reach the level of the middle of the eye (the
adult position) and are almost horizontal. The only figures
of adult R. bicolor are that in the original description (Goode
& Bean, 1895: plate 17, fig. 1), and a painting of Bermuda
specimens (Harry, 1952: plate 1), neither of which adequately
illustrate characters considered important now. The new
illustration (Fig. 4) is based on a 60 mm si specimen from
the central Atlantic kindly provided by K. Hartel of MCZ.
Pigmentation. The two smallest specimens have faded with
80+ years storage in formalin and the only remaining
pigment is that dark brown covering the stomach of the
13.5 mm specimen. The 14.4 mm specimen is covered with
the loose, uniformly dark brown skin characteristic of
preserved adults. At this size an even layer of subdermal
melanophores is present under the gelatinous connective
tissue over the main muscle mass. At 60 mm light irregular
streaks are present on the surface of the muscles.
Scales. At 7.2 mm two parallel rows of small, circular,
cycloid scales extend from the top of the opercle to the
level of the PU1+U1 centrum of the caudal skeleton. The
scales are arranged approximately one per myomere and
number 24-25 per row. The scales of each row are separated
by a space equal to one half to one scale diameter and the
two rows are separated by an equal space. The scales overlie
the skin and are very weakly ossified, picking up much less
alizarin than the fin rays or other developed bones. Two
other rows of scales are developing on either side of the
dorsal midline, where seven smaller scales are present from
the level of the preopercle to half way to the dorsal-fin
origin. A few apparent scale primordia are present in the
area between the pectoral- and pelvic-fin bases. No other
scales are apparent on the body or head.
In the 13.5 mm juvenile the scales remain very weakly
ossified and can only be seen with certain angles of reflected
light. The scales of the two rows in the lateral-line region
414 Records of the Australian Museum (2001) Vol. 53
Figure 3. Rondeletia bicolor, a, ZMUC P2340805, 7.2 mm si larva, note outline of Tominaga’s organ anterior to
eye (dashed line); b, ZMUC P2334327, 13.5 mm si juvenile, Tominaga’s organ is extensive anterior to the eye.
have increased in size so that some scales overlap slightly
within each row. These scales are now embedded in the
skin at a slight angle with the anterior edge of each scale
deeper in the tissue. They are dorsoventrally ovoid and the
space between the two rows is only about one-tenth of a
scale diameter, with the two rows sometimes touching. The
scales extend to the urostyle and number 24 in a row. A row
of scales on either side of the dorsal midline extends from
the level of the preopercle to the dorsal-fin origin and
numbers 10-11 small circular scales. There is another group
of circular scales in a triangular area between the pectoral-
and pelvic-fin bases and the ventral midline. There is no
indication of spines on any scale of either specimen. Further
description of scales in larger specimens is presented after
the larval descriptions.
Head spines. There are no strong head spines in the two
smallest specimens. Two very weak spines are present on
the opercle of the 7.2 mm larva. In the 13.5 mm juvenile a
spine is beginning to develop on the dorsal end of a ridge
on the anterior orbital margin of the sphenotic. In the larva
the infraorbitals are just beginning to ossify and a small
amount of spongy bone is present only in the posterior
portion of the frontal. In the smallest juvenile, spongy bone
is evident on the frontal, sphenotic, parietal, supraoccipital,
epioccipital and pterotic. All elements of the pectoral girdle
lack spongy bone in both of the smallest two specimens.
Fin formation. In the larva, all of the fins have the complete
complement of rays (Table 4). None of the 6 rays of the
pelvic fin is greatly produced, with the 3rd-5th rays longest.
In both of the two smallest specimens the longest rays extend
to the base of anal-fin ray 4-5. In the 14-17 mm juveniles
the rays extend only to the anal-fin origin, whereas in the
60 mm adult the pelvic-fin rays do not reach the anal-fin
origin. In both of the two smallest specimens the pelvic-fin
origin is at about the level of the 10th vertebra, slightly
anterior to the dorsal-fin origin and about 2 A of the way
between the head and anal-fin origin. The pelvic fins are in
about the same position in the smaller juveniles (< 20 mm),
but the pelvic fin of the 60 mm specimen is closer to halfway
between the head and anal-fin origin. Both of the smallest
specimens have 6 supraneurals anterior to the dorsal-fin
origin.
Paxton et al .: Barbourisia and Rondeletia larvae 415
Figure 4. Rondeletia bicolor, AMS 118415-001, 60.0 mm si adult.
Dentition. The 7.2 mm larva has a single row of small,
triangular teeth in both jaws. The premaxillary teeth are
widely spaced, those of the dentary closely set with some
almost touching at their bases. The teeth of the 13.5 mm
juvenile are very closely set in both jaws, but still primarily
in one row. In the 17 mm juvenile the teeth are in 1-2 rows,
while in a 60 mm adult there are 5-6 small conical teeth
rows across the oral surface of each jaw.
Internal anatomy. In the 7.2 mm larva the stomach is
moderately large, occupying about % of the abdominal
cavity, and appears to have a smaller anterodorsal portion
and a larger posterior portion. The intestine emerges from
the anteroventral region of the posterior portion of the
stomach. Coiling of the intestine is not clear; the intestine
ends in a long straight section in the ventral abdominal
cavity from the level of the stomach to the anus slightly
closer to the anal-fin origin than the pelvic-fin base. A small
mass of tissue at the top of the stomach may represent a
developing, non-functional swimbladder. In the 13.5 mm
juvenile the stomach occupies about Vi the abdominal
cavity. The intestine emerges from the anteroventral arm
of the stomach with apparently some folding on the right
side of the stomach. The course of the intestine to the
anus, about midway between the pelvic-fin base and anal-
fin origin, is unclear. No pyloric caeca are apparent. The
swimbladder is not apparent. Adults also lack a
swimbladder (Parr, 1929).
A large mass of globular white tissue is present anterior
to the orbit and posterior and medial to the nostrils and
developing nasal rosette in both of the two smallest
specimens. Tominaga (1970) briefly described similar tissue
in an adult R. loricata, and we here term it Tominaga’s organ.
In the 7.2 mm larva the organ is slightly smaller than the
orbit and extends anteriorly to the posterior margin of the
developing nasal organ. In the 13.5 mm juvenile the organ
is larger than the orbit and extends to the anterior margin of
the nasal organ. The adult condition is described more fully
following the description of the larvae of R. loricata.
Caudal skeleton. All specimens have a full complement
of caudal elements and fin rays. There are three epurals
(the first originating over the dorsal crest of preural
centrum two), two uroneurals, one parhypural and six
hypurals (two ventral and four dorsal). In the two smallest
specimens ural centrum 2 is a separate, distinct
ossification that abuts against and appears to be fusing
with the base of hypural 4. The base of hypural 3
articulates along the notochord in the space between PU1-
U1 and U2. In the 7.2 mm larva the distal tips of the
parhypural and hypurals 1-5 are unossified and hypural
6 is a tiny ossification dorsal to hypural 5. Uroneural 2
is very small and epurals 2 and 3 are unossified. In the
13.5 mm juvenile all hypurals and epurals are completely
ossified. Hypurals 1 and 4 are the largest and hypural 6
remains autogenous. In both specimens the parhypural
and hypurals 1 and 2 articulate with a large oblong
cartilage below the urostylar centrum. Hypurals 1 and 2
are fused distally in both specimens, and in the larger
they have also fused proximally, similar to the condition
in our third cleared and stained specimen, a 21.7 mm si
juvenile. Parr (1929: fig. 18) figured the caudal skeleton,
presumably of an adult specimen, with little description.
His figure shows the proximal but not the distal fusion
of hypurals 1 and 2 and does not show hypural 6.
Distribution. All specimens examined for this study were
collected in the western North Atlantic, where most
specimens of this species have been collected (Table 3; Fig.
2; Paxton, 1974; Kotlyar, 1996). In an addendum, Paxton
(1974: 188) noted a single adult specimen collected in the
southeast Pacific at 25°48'S 108°46'W (near Easter Island
off Peru) that Kotlyar (1996: 220) considered most likely
based on an error in determination or labelling. The original
information was received in 1970 about a 1969 SIO
expedition to that area, and is unlikely to be a labelling
error. The 83 mm specimen was re-examined recently by
H.G. Moser and R. Rosenblatt and found to be correctly
identified, with the diagnostic bony hook over the orbit
present. In addition, a 44 mm specimen from 15°S 175°W
in the central Pacific collected in 1927 (ZMUC P2334334)
was identified by the first author and confirms the presence
of R. bicolor in the South Pacific.
The larva and juveniles were all caught with open nets,
with the larva caught in the upper 50 m and the juveniles in
nets fishing from 200 to 1100 m depth.
416 Records of the Australian Museum (2001) Vol. 53
Table 3. Rondeletia bicolor material examined. Abbreviations and symbols as in Table 1.
specimen
catalogue
size
location
depth (m)
day/night
date
1 *#
ZMUC P2340805
7.2
31°59'N 59°52'W
0-50
N
24 Oct 1913
2 *#
ZMUC P2334327
13.5
17°41'N 60°58'W
0-200
N
27 Nov 1921
3
MCZ 50681
14.4
23°13'N 44°56'W
0-1100
15 Oct 1973
4
ZMUC P2334332
17.0
19°04'N 65°43'W
0-900
DN
09 Mar 1922
5
ZMUC P2334328
18.0
19°01'N 65°23'W
0-600
N
03 Jan 1922
6 *
ZMUC P2334331
21.7
24°05'N 74°36'W
0-650
D
15 Feb 1922
7 #
AMS 118415-001
60.0
9°15'N 49°16'W
22 Sep 1973
Table 4. Rondeletia bicolor counts. Abbreviations, symbols and lines as in Table 2; D hypurals = dorsal hypurals.
specimen
size
D
A
P
P2
Cprin
Cproc
Supran
Myom/Vert
scale rows
D hypurals
1 *
7.2
15
15
10
6
10+9
5+4
6
27
24/25
4
2 *
13.5
15
14
10-11
6
10+9
5+5
6
27
24
4
3
14.4
14
14
10
6
10+9
5+4
4
17.0
15
15
10
6
10+9
5+5
27
5
18.0
15
14
10
6
10+9
5+5
25
6 *
21.7
15
14
10
6
10+9
5+5
7
27
4
Rondeletia loricata Abe & Hotta, 1963
Figs. 5, 6
Eight larvae 3.5-9.7 mm si, one flexion and seven
postflexion, were examined (Table 5). In specimens 8.8 mm
and larger, there is a moderate to large amount of gelatinous,
fibrous connective tissue between the skin and muscle mass
and the skin is loose and slightly inflated, somewhat
reminiscent of lophiiform larvae (Pietsch, 1984). Large
amounts of thick connective tissue under the skin are typical
of the adults of both species of Rondeletia. The head and
body of the smallest specimen are moderate in depth,
becoming deeper with increasing size (4.1-4.6 mm). The
head and anterior body are deepest in the 8.8-9.7 mm larvae.
The 12.7 mm juvenile R. loricata is deeper in both head
and body than the 13.5 mm R. bicolor. The eye is large and
the snout short in the smallest specimens, while by 8.8-9.7
mm the snout and eye sizes approach the ratio typical of
the juvenile and adult. In the smallest larvae (3.5-4.6 mm)
the jaws are short and moderately oblique, and almost or
just reach the level of the anterior margin of the orbit. The
jaws lengthen in the 8.8-9.7 mm larvae and become almost
horizontal by 12.7 mm. Jaw length displays allometric
growth in the juveniles (Paxton, 1974: fig. 2), with the
posterior end of the upper jaw nearing the level of the middle
of the orbit only in a 22 mm juvenile.
Pigmentation. The larva retaining the most pigment is a
4.6 mm specimen (Fig. 5a) collected in 1985. The remaining
larval specimens were collected at least 25 years ago and
the three smallest were collected more than 80 years ago
and stored for most of that time in formalin. All are faded
to a greater or lesser degree. The 4.6 mm larva has the body
and head covered with widely spaced melanophores. All of
the fin rays are unpigmented except those of the pelvic fin,
which are densely covered with melanophores that are larger
and darker than those on the head and body. Some myoseptal
pigment is present in the region of preural centra 2-3, but
the subdermal melanophores typically found on the surface
of the muscles in the larger larvae are not evident. The
stomach is dark, as in all the larvae.
In the 3.5 and 4.1 mm larvae faded melanophores are
visible on the pelvic-fin rays, and to a lesser extent under
the posterior bases of the dorsal and anal fins of the smaller
specimen. The eye is dark while all the remaining tissues
of the head and body are yellowish to light brown. In the
8.8 mm and 9.6 mm larvae all the pigment in the skin has
faded and only the pelvic-fin rays have distinct melano¬
phores. In the 9.7 mm larva (Fig. 5c), the skin of the head
and body is covered with light, closely-spaced melano¬
phores. In this specimen, small widely-spaced melanophores
are present on the surface of the muscle mass underneath
the skin and connective tissue, as in the 9.6 mm specimen
(Fig. 5b). This subdermal pigment extends from the base
of the skull back to the end of the caudal peduncle. In the
12.5-14 mm juveniles the skin is uniformly dark brown as
in adults; the pelvic fins have lost much pigment distally
and are only slightly darker than the body skin proximally.
At this size the neuromasts of the lateral-line system on the
body are visible. Pigment extends onto the bases of the rays
of all the other fins. The subdermal pigmentation also
increases and at 14.1 mm extends over the main muscle
mass and is also visible on the skull bones. In the region of
the posttemporal there are three layers of melanophores,
one in the skin, one within the spongy bone and one on the
surface of the muscles that have been overgrown by the
posterior extension of the posttemporal. By 33 mm the
subdermal pigmentation is reduced to light irregular streaks
over the muscles that are visible also in adult specimens
after the connective tissue has been removed.
Scales. Scales are visible in our specimens at 8.8 mm and
above. In the 9.6 mm specimen two rows of round, thin
Paxton et al .: Barbourisia and Rondeletia larvae 417
Figure 5. Rondeletia loricata. a, AMS 125228-001, 4.6 mm larva, note outline of Tominaga’s organ anterior to eye
(dashed line); b, LACM 36982-1, 9.6 mm larva showing internal pigment, outline of Tominaga’s organ (anterior to
eye), spongy bone of head and pectoral girdle, and supraneural bones; c, outline of b. showing predorsal scales,
trunk and tail scales, and external pigment; pigment derived from 9.7 mm larva (MCZ 50683).
scales overlie the skin and extend from above the opercle
to the caudal peduncle (Fig. 5c); each row includes 17-18
scales. Seven smaller scales are present in a row just off the
dorsal midline over the posterior portion of the head. One
large and nine smaller scales are present below and behind
the pectoral-fin base.
Head spines. Head spines are lacking in all our larval
specimens. At 4.6 mm there is a considerable amount of
spongy, sculptured bone in the supraorbital region of the
frontal. The pectoral girdle is weakly ossified with neither
spongy bone nor posterior expansions of the posttemporal
or cleithrum present. However, the dorsal portion of the
418 Records of the Australian Museum (2001) Vol. 53
cleithrum is wider than that of a 7.2 mm R. bicolor. In the
9.6 mm specimen spongy bone is well developed dorsally
on the frontals and supraoccipital, with separate lateral
patches on the parietal/pterotic and preopercle, and on most
elements of the pectoral girdle—the posttemporal,
supracleithrum and cleithrum. Posterior extensions of
spongy bone are developing dorsally and ventrally on the
posttemporal and cleithrum respectively.
Fin formation. At 3.5 mm the dorsal, anal and pelvic fins
have the full complement of rays (Table 6). The third and
fourth rays of the pelvic fin are the longest, extending
beyond the anal-fin base; these two rays are about one-third
longer than the second and fifth rays and almost twice as
long as the first pelvic-fin ray. At 8.8-9.7 mm, pelvic-fin
rays 2 and 5 are subequal to rays 3 and 4 and all extend to
anal-fin rays 2-3. Pelvic-fin rays 3 and 4 are the same
absolute length (1.5-1.6 mm) in both the 4.6 and 9.7 mm
specimens; the negative allometry is also evident in small
juveniles. In adults the pelvic-fin rays do not reach the anal-
fin origin. The pelvic-fin base is much closer to the anal-
fin origin than to the pectoral-fin base at 4.6 mm. By 9.7
mm the pelvic-fin base is closer to midway between the
two fins than to the anal-fin origin, similar to the adult
condition.
The pectoral fins are damaged in the smallest larva, but
at least three rays are visible on the right fin. By 4.6 mm all
but the last ray is ossified. In the four smallest larvae the
pectoral fin is relatively high on the side of the body. By
Figure 6 (left). Rondeletia loricata, AMS 121141-001, 73.4 mm
adult, a, position of Tominaga’s organ; dotted line—outline of
cavity of Tominaga’s organ; long dashed line—outline of lobes of
Tominaga’s organ; short dashed line—cavity of nasal organ; scale
= 1 cm. b, detail of Tominaga’s organ, anterior to left, showing
anterior ducts to cavity of nasal organ; scale = 1 mm.
Table 5. Rondeletia loricata material examined. Abbreviations and symbols as in Table 1.
specimen
catalogue
size
location
depth (m)
day/night date
1
ZMUC P2334325
3.5
26°46'N 54°14'W
0-8
N
16 Jul 1920
2
ZMUC P2334326
4.5
28°20'N 63°50'W
0-8
N
21 Jul 1920
3 *
ZMUC P2334323
4.6
28°49'N 54°10'W
0-17
N
15 Jul 1920
4 #
AMS 125228-001
4.6
14°33'S 145°36'E
0-40
D
11 Feb 1985
5
NSMT PL108
5.0
17°00'S 118°00'E
0-75
N
21 Jan 1993
6
ZMUC P2334335
8.8
11°00'S 172°37'W
0-333
DN
02 Nov 1928
7 *#
LACM 36982-1
9.6
21°23'N 158°18'W
23 Jun 1971
8
MCZ 50683(1)
9.7
23°08'S 32°22'W
0-110
09 Mar 1967
9
MCZ 50684
12.6
25°52'N 36°48'W
0-140
30 Nov 1970
10 *
MCZ 50679(1)
12.7
23°02'S 32°15'W
0-175
09 Mar 1976
11
MCZ 50679(2)
13.0
12
AMS 127620-001
13.0
21°23'N 158°18'W
11 May 1972
13
MCZ 50679(3)
13.2
14
MCZ 50683(2)
13.2
15
MCZ 50679(4)
13.3
16
MCZ 50683(3)
14.1
17
MCZ 50680
15.5
27°03'N 53°56'W
0-1000
08 Oct 1972
18 *
MCZ 50679(5)
18.3
19 *
AMS 120522-001
23.8
22°N 158°W
0-1000
N
05 Nov 1976
20 *
AMS 120314-011
37.1
33°28'S 152°33'E
0-900
D
14 Dec 1977
21 *
AMS 120307-011
60.4
33°28'S 152°25'E
0-900
DN
13 Dec 1977
22 *
LACM 9254-33
94
32°13'N 120°41.5'W
0-400
N
18 Oct 1966
Paxton et al .: Barbourisia and Rondeletia larvae 419
Table 6. Rondeletia loricata counts. Abbreviations, symbols and lines as in Tables 1 and 2. Specimen between
dashed lines is undergoing notochord flexion. + = present but accurate counts not possible.
specimen
size
D
A
P
P2
Cprin
Cproc
Supran
Myom/Vert
scale rows
D hypurals
1
3.5
13
12
>3
5
c. 5+4
—
24
—
2
4.5
13
12
?
5
10+9
_
3 *
4.6
13
12
?
5
?
—
—
24
—
?
4
4.6
13
13
9(1)
5
10+9
—
5
5.0
14
13
8+
5
?
—
6
8.8
13
13
10
5
10+9
—
+
7 *
9.6
14
13
10
5
10+9
5+4
3
24
17-18
3
8
9.7
13
12
10
5
10+9
3?+2?
9
12.6
14
13
10
5
10+9
5?+4?
10*
12.7
13
13
10
5
10+9
4-5+4
3
24
?
3
18*
18.3
13
14
11
5
9+9
4+4
3
24
?
3
19*
23.8
13
13
10
5
?
?
4
25
?
3
20*
37.1
14
13
9
5
10+9
5+4
4
25
?
?
21 *
60.4
14
13
10
5
10+9
5+4
?
25?
?
3
22*
94
14
13
5
10+9
5+5
7
26
?
3
9.7 mm the pectoral-fin base is in a lower position as in the
adults. The caudal fin is damaged in the smallest larva,
where there are approximately 5+4 incipient principal rays.
By 4.5 mm the notochord is fully flexed and the caudal fin
has the full complement of principal caudal rays. Procurrent
rays are apparent from 9.6 mm. Only the 9.6 mm larva has
visible supraneurals, with three.
Dentition. At 4.6 mm a single row of tiny triangular teeth
are present in both jaws. Those of the dentary are closely
set, the spacing of those on the premaxilla unclear. In the
smallest juveniles at 12.7 mm both jaws bear a single row
of closely set, conical teeth.
Internal anatomy. None of the four smallest larvae (3.5-
4.6 mm) is transparent enough to see details of the
internal organs. The stomach is large, occupying half or
more of the abdominal cavity. In the three largest larvae
8.8-9.7 mm the stomach is small to massive, occupying
one-third to two-thirds of the abdominal cavity,
presumably depending on the amount of stomach
contents. None of these larvae is clear enough to see other
details. In a 12.7 mm cleared and stained juvenile the
stomach fills about one half the abdominal cavity, and
the intestine exits from the anteroventral margin of the
stomach. The intestine has one loop in the dorsal portion
of the coelom to the right of the stomach and another
smaller loop further posterior, exiting through a short
straight section anterodorsal to the anus. Other organs,
such as swimbladder and pyloric caeca, are either
undeveloped or have been digested in the clearing
process.
Tominaga’s organ is visible in the 4.1-4.6 mm larvae. It
may be present in the 3.5 mm larva, but the poor condition
of the specimen makes it difficult to discern. The organ is
initially small and is located above the nasal organ. As the
snout elongates, Tominaga’s organ extends posteriorly to
fill most of the gap between the nasal organ and the eye.
The anterior margin of Tominaga’s organ is dorsomedial to
the anterior of the nasal organ in all postflexion larvae.
Caudal skeleton. The smallest cleared and stained larva,
4.6 mm si, is damaged in the caudal area. In the 9.6 mm
larva the bone is well stained with alizarin. Cartilage stained
well and bone poorly in the 12.7 mm juvenile. Both the
cleared and stained larva and 12.7 mm juvenile have the
same caudal elements: three epurals, at least one uroneural,
two ural centra (PU1-U1 and U2), five hypurals (two ventral
and three dorsal) and one parhypural. The parhypural and
hypurals 1 and 2 articulate with a large oblong cartilage
ventral to PU1-U1; the haemal spines of preural vertebrae 2-
4 also articulate with a cartilage ventral to their respective
centra. Hypural 3 articulates with the notochord at the space
between PU1-U1 and U2, while hypural 4 articulates with U2.
Distribution. Four of the eight larvae were collected in the
central and western North Atlantic, one in the North Pacific
near Hawaii, two in the South Pacific near Samoa and in
the Coral Sea, and one in the eastern Indian Ocean off NW
Australia (Table 5; Fig. 2). The species is recorded from all
oceans between 47°N and S (Paxton, 1974; Kotlyar, 1996).
The eight larvae were all taken with open nets, fishing
to a maximum of 333 m. The two smallest larvae were taken
in the upper 8 m, while the next two smallest larvae were
taken in nets fishing to 17 and 40 m. The shallowest capture
depth is 110 m for the 10 juveniles less than 20 mm si, and
eight of these were caught with open nets fishing only to
110-175 m (Table 3). The vast majority of adult specimens
over 50 mm si have been caught with nets fishing below
400 m (Bast & Klinkhardt, 1990; Paxton, unpublished).
Thus there is a clear indication of ontogenetic descent,
beginning when the larvae reach 4-5 mm si.
Scales
Scales of adult Barbourisia rufa were described and figured
by Struhsaker (1965). Scales have not been reported
previously in the family Rondeletiidae. Developing
individual scales were first seen in cleared and stained
larvae, as described above. In the cleared and stained 21.7
mm juvenile R. bicolor in poor condition, no scales are
420 Records of the Australian Museum (2001) Vol. 53
visible. However, in the cleared and stained 18.3 mm R.
loricata very thin scales are visible in two separate rows,
with scales within a row overlapping by 10-30% of scale
length. In this specimen, and confirmed by dissection in
smaller, unstained juveniles, the scales are underneath the
skin in the presumed connective tissue over the underlying
body muscles. Strands of presumed connective tissue attach
the anterior end of the scales to the underlying muscle and
the posterior end of the scales to the overlying skin. A very
thin layer of overlapping scales is visible in some specimens
(those with the best preservation?) 35-85 mm si, embedded
in presumed connective tissue between the skin and muscle
on the side of the body. These scales are so thin, and take
up stain so poorly, that they have never been identified, or
at least described previously, in larger cleared and stained
specimens.
[A
?
&
v?
S-
-
E
tz
£
c
0.15-I
0.00 4 -
0
A. ft. iitfufor ittcW.
■ ft. rtl'iWtf .
■ ft, mm
E~l ft. :r.i\-.JTJ .1tt7U.fr
60
120
slandard lenglh, mm
Figure 7. Length of Tominaga’s organ as %sl/sl; “indet.”—sex
indeterminate (larvae and juveniles) or not determined (adults).
Tominaga’s organ
Tominaga (1970) briefly described an unnamed structure
lying under the frontal between the nasal rosette and orbit
of R. loricata. The organ was described as having two
subequal lobes with no apparent external openings or ducts.
Based on histology, each lobe was comprised of multiple
globules with hollow centres and the large cells surrounding
the cavities stained well with the acidic dyes acid violet,
phloxin and light blue. Tominaga did not propose a function
for this organ.
Rondeletia loricata. Dissection of 20 specimens 31-109
mm si (not listed in the material examined) representing
both sexes confirmed the above description. The organ, here
termed Tominaga’s organ, develops in a cavity in front of
the eye that extends dorsally below the lateral shelf of the
frontal, anteriorly medial to the posterior half of the nasal
cavity and posteriorly to, or medial to the anterior portion
of, the orbit (Fig. 6a, PL la). The lining of the cavity has
sparse grey-brown pigment, as does the covering of the two
lobes of Tominaga’s organ. The lobes are light yellow or
orange-pink in colour and about equal in length, but the
lateral lobe is somewhat larger in width and therefore
volume. The lobes are posteromedial to the nasal organ and
do not reach the level of the posterior margin of the floor of
the nasal cavity in which the nasal rosette lies (PL lb). The
lateral ethmoid is greatly reduced in relation to other
stephanoberycoids, (see Kotlyar, 1996: fig. lb) with
Tominaga’s organ filling much of the space normally
occupied by that bone.
The olfactory nerve runs between the two lobes of
Tominaga’s organ to enter into the floor of the nasal cavity
and central raphe. Two internal pores at the posterior end
of the raphe of the nasal rosette open into this region, and a
thin-walled duct runs from each pore to the anteromedial
portion of each lobe of Tominaga’s organ (Fig. 6b). Each
duct appears to branch within the lobe, but these branches
could not be followed.
The globular structure of the organ is visible with a
dissecting microscope and clearly shown histologically. The
cavities of the globules or chambers are lined by a single
layer of cells, some of which are simple, squamous
epithelium, while adjacent globules may be lined with
simple cuboidal epithelium. One globule has some
flocculent material that appears granular. The histological
structure suggests a secretory function (J. Burns, pers.
comm. June 2000). In one section elongation of the cavities
and a duct lined with epithelial cells is visible. However,
this could not be traced to the main duct to the nasal cavity,
and no pattern or system of ducts could be found.
Measurements of the maximum length of individual
Tominaga’s organs of 33 R. loricata (Fig. 7) indicate the
organ reaches its maximum relative size of 11-14% si in
juvenile specimens 13-20 mm si. However, the organ
continues to grow throughout life, as the longest measured
(9.0 mm) is in a 109 mm specimen. There is no correlation
of organ size or appearance with sex. At about 60 mm si,
increasing amounts of connective tissue are found in the
cavity housing Tominaga’s organ. By 90 mm and larger the
cavity is almost filled with connective tissue, which also
appears to invade the organ.
Figure 8. Rondeletia bicolor, AMS 118415-001, 60.0 mm si adult,
showing position of Tominaga’s organ, line conventions as in Fig.
6, scale = 1 cm.
Paxton et alBarbourisia and Rondeletia larvae 421
Plate 1. Tominaga’s organ, a, Rondeletia loricata, USNM 206836, 83 mm si; b, R. loricata, same specimen; c, R.
bicolor, USNM 240130, 35 mm si; d, Gibberichthys pumilis, USNM 207512, 75 mm si, arrow indicates right
dorsal margin of Tominaga’s organ.
Rondeletia bicolor. Tominaga’s organ is similar to that
described for R. loricata, except for the following
differences. The lobes of Tominaga’s organ of R. bicolor
are semi-equal in volume, with the medial lobe notably
longer than the roughly spherical lateral lobe (Fig. 8). The
lobes are medial to the nasal organ, with the lateral lobe
extending anteriorly beyond the posterior margin of the
nasal organ to a point about one quarter along the nasal
rosette. The longer medial lobe extends anteriorly almost
to the anterior end of the nasal rosette. Consequently the
ducts from the pores at the end of the raphe enter nearer the
midpoint of each lobe, rather than at the anterior margin as
in R. loricata. The posterior floor of the nasal organ is tightly
bound by tough connective tissue to the dorsal surface of
the lobes in R. bicolor. Measurements of the few available
specimens of R. bicolor do not indicate significant differences
with R. loricata in the length of the organs in relation to standard
length (Fig. 7). However, relative to snout length, both
Tominaga’s organ and the nasal rosette are larger in R.
bicolor than in R. loricata (Figs. 6, 8; PI. lc).
Gibberichthys. A search for Tominaga’s organ in other
stephanoberyciform taxa (Table 7) revealed a similar
Figure 9 (right). Gibberichthys pumilis, CAS 14565, c. 67 mm.
a, position of Tominaga’s organ, line conventions as in Fig. 6,
Tominaga’s organ fills cavity, scale = 1 cm; b, detail of Tominaga’s
organ, anterior to left, nasal organ anterodorsal to Tominaga’s
organ, scale = 1 mm.
422 Records of the Australian Museum (2001) Vol. 53
structure only in the gibberichthyids Gibberichthys pumilis
and G. latifrons. The following description is based on two
dissected specimens of G. pumilis ; superficial dissection of
G. latifrons revealed no basic differences. Tominaga’s organ
is medial to the lacrymal and ventromedial to a shelf of the
lateral ethmoid, upon which the nasal rosette sits (Fig. 9).
There are no pores at the posterior end of the nasal rosette,
and no ducts or opening in Tominaga’s organ are evident.
The roughly pyramidal-shaped organ (PI. Id) has globular
internal structure, but is not divided into distinct lobes. The
slightly rounded dorsal surface of the organ is tightly bound
to the ventral surface of the lateral ethmoid by connective
tissue. Histology reveals globules lined with low epithelium
and filled with flocculent tissue and purple granules that
indicate secretory function. No globules lined with cuboidal
epithelium, as seen in Rondeletia loricata, were apparent
(J. Burns, pers. comm. June 2000).
Discussion
The larvae of Barbourisia and Rondeletia are easily
recognized primarily because they exhibit adult character¬
istics at an early stage in development. Neither have highly
specialized larval morphology, except for the large,
precocious pelvic fins. By 10 mm larval Barbourisia have
a few clusters of minute spines on some head bones and an
inflated, balloon-like envelope of skin. They differ further
from the adults in having relatively elongate pelvic fins, a
smaller, more oblique mouth, larger eye, and shorter snout.
Although the body is covered with the distinctive scales of
the adult by about 6 mm nl, the lateral line is represented
only by enlarged scales with no canal formation even at 13
mm. The changes that take place with attainment of the
juvenile stage include loss of the spines on the head bones,
reduction of the pelvic-fin rays, and formation and closure
of the lateral-line canal. The smallest examined juvenile is
30.0 mm si.
Larval Rondeletia are extremely precocious (flexing at
3.5 mm) and even less specialized than those of Barbourisia,
differing from the adult in having heavily pigmented and
relatively longer pelvic fins, a smaller mouth and superficial
scales, those on the lateral body arranged in two distinct
rows. By 14 mm the juveniles look like miniature adults.
The two species of Rondeletia can be distinguished in
the early larval stages by meristics and in late larvae and
juveniles by posterior extensions of spongy bone in the
posttemporal and cleithrum.
There have been conflicting descriptions of the caudal
skeleton in the past based on adult osteology, with
differences in the described number of dorsal hypurals not
corresponding to species. Parr (1929: fig. 18) figured three
dorsal hypurals with a question for R. bicolor and Kotlyar
(1996: fig. 3d) showed two dorsal hypurals for R. loricata,
while Ebeling & Weed (1973: fig. 5) illustrated four in R.
bicolor, and Rosen (1973: fig. 121) and Paxton (1974)
described three in R. loricata. Development of the bones of
the caudal skeleton have clarified the different number of
dorsal hypurals in the two species, four in R. bicolor and
three in R. loricata. Thus, Parr (1929) apparently did not
see the small, dorsal-most sixth hypural in his specimen of
R. bicolor, and Kotlyar (1996) interpreted the fusing
hypurals 3 and 4 of R. loricata as a single hypural 3.
The number of ossified supraneural elements above the
vertebrae anterior to the dorsal fin also varies. Paxton (1974)
described seven in a 94 mm R. loricata, while Kotlyar
(1996) indicated three or four in his 93 mm specimen. Five
larvae and juveniles here have three or four supraneurals,
while the count of seven in the 94 mm specimen is verified.
The larva and two juveniles of R. bicolor have six or seven
supraneurals.
There are distinct differences in the relation of
Tominaga’s organ and the nasal rosette in adults of the two
species of Rondeletia. Tominaga’s organ is entirely posterior
to the nasal organ in R. loricata, with the ducts entering the
anterior end of Tominaga’s organ. In R. bicolor, the
anterior half of Tominaga’s organ is medial to the nasal
organ and the connecting ducts enter about midway along
Tominaga’s organ. The presence of two separate lobes, as
well as ducts to the nasal cavity, in Rondeletia suggests that
Tominaga’s organ is more specialized in Rondeletia than in
Gibberichthys.
Gross structure and histology suggest a secretory function
for Tominaga’s organ, but the nature of the presumed
secretion is unknown. There is no difference in size
correlated with sex where a number of specimens are
available to measure in R. loricata, and the opening of the
ducts into the nasal cavity seems incongruous for
pheromone function. Perhaps the flap on the posterior nostril
of Rondeletia is involved in dispersal of the secretion. In
Gibberichthys any secretion would be internal, as no
external opening is discernible. There is nothing in the
structure to indicate luminescence. While magnetoreceptor
cells have been described in the same general anatomical
region, inside the nasal lamellae of the nasal organ of
rainbow trout (Diebel et al., 2000), homology with
Tominaga’s organ seems unlikely. Fresh tissue would be
needed to detect intracellular magnetite. Other possibilites,
such as a toxic repellent, are mere conjecture. Future study
of fresh or better preserved specimens is needed.
The relationships of the Barbourisiidae and Rondeletiidae
to other “beryciform” fishes remain problematic. However,
they have frequently been associated with the Cetomimidae,
sometimes as a suborder or superfamily, more recently with
the Mirapinnidae and Megalomycteridae (Harry, 1952;
Greenwood et al., 1966; Ebeling & Weed, 1973; Rosen &
Patterson, 1969; Paxton, 1989; Nelson, 1994). Parr (1929)
placed the Rondeletiidae in the Xenoberyces (= Stephano-
beryciformes), while Rofen (in Ebeling & Weed, 1973: 399)
and de Sylva & Eschmeyer (1977) commented on the
similarity of Rondeletia and Gibberichthys. Most recently
all three whalefish families have been placed with other
families Mirapinnidae, Megalomycteridae, Stephano-
berycidae, Hispidoberycidae, Gibberichthyidae and
Melamphaidae in an order or suborder (Rosen, 1973;
Moore, 1993; Johnson & Patterson, 1993).
The most recent hypothesis of relationships among these
families is that of Moore (1993, fig. 5). Based on one
character (Y-shaped pattern of frontal ridges), he placed the
Gibberichthyidae as the sister group of Stephanoberycidae +
Hispidoberycidae in one lineage, which he considered to be
the sister group of a second lineage comprising, in phyletic
sequence, Rondeletiidae, Barbourisiidae, Megalo¬
mycteridae and Cetomimidae. Placement of Rondeletiidae
within the latter lineage was again based on one character
(loss of fin spines).
Paxton et alBarbourisia and Rondeletia larvae 423
Table 7. Other specimens examined. a taken near the surface of 2743 m deep waters, b bottom trawl, c Moore & Merrett manuscript.
taxon catalogue no. (size, mm) location depth (m) day/night date
Anoplogastridae
Anoplogaster cornuta
AMS 127174-003
1(29)
Cetomimidae
Cetostoma regani
SIO 70-95
1(116)
Ditropichthys storeri
AMS 121143-001
1(84)
Ditropichthys storeri
AMS 128177-001
1(142)
Procetichthys kreffti
ISH 1188/71 holotype
1(236)
Gibberichthyidae
Gibberichthys latifrons
AMS 115999-001
1(103)
Gibberichthys pumilis
UMML 16213
1(7.8)
Kasidoron edom
CAS 14565 paratype
2(48.9-c.67)
Hispidoberycidae
Hispidoberyx ambagiosus
MNHN unregistered
1(175)
Megalomycteridae
Ataxolepis apus
MCZ 60720
1(41)
Melamphaidae
Scopelogadus mizolepis
AMS 125858-008
1(89)
Stephanoberycidae
Acanthochaenus luetkeni
AMS 128176-001
1(94)
n.gen. n.sp. c
AMS 140443-001
1(134)
Trachichthyidae
Hoplostethus latus
AMS 131163-007
1(100)
22°46'S 177°00'E
0-230
03 Sep 1987
31°37'N 120°19'W
0-C.1100
22 Mar 1970
21°25'N 158°25'W
0-3440
01 Jun 1976
29°49'S 47°24'E
N
27 Dec 1988
37°08'S 5°23'E
0-2200
N
21 Mar 1971
11°17'S 142°47'W
7-8 Feb 1969
32°46'N 64°33'W
0-0.3 a
N
03 Aug 1964
29°16'N 86°55'W
660 b
12 Feb 1970
S of New Caledonia
1350
10 Nov 1996
17°06'N 73°37'W
18 Jun 1982
54°44'N 18°23'W
0-800
D
06 Jul 1986
30°27.5'S 46°56.5'E
2680
D
26 Dec 1988
20°53'N 31°14'W
4522
04 Oct 1993
24°52'S 112°07'E
468
28 Jan 1991
The recently published DNA sequence data analysis
(Colgan et al., 2000), which did not include Gibberichthys,
placed Barbourisia and Rondeletia as sister groups. We
think that the unique presence of Tominaga’s organ in
Rondeletia and Gibberichthys, together with additional
morphological characters discussed below, belies that
hypothesis, and provides convincing evidence for a sister
group relationship between the latter two taxa.
Moore (1993) did not discuss Rosen’s (1973: 492)
assertion that “on the evidence of the lateral-line canal, jaw
musculature and pharyngobranchials Rondeletia is most
closely related to Gibberichthys Our observations confirm
the striking similarities between the two taxa in jaw
musculature (Rosen, 1973: fig. 37) and dorsal gill-arch
elements (Rosen, 1973: figs. 122-124), and the distinctive
presence in both taxa of vertical rows of free neuromasts as
lateral-line organs. Furthermore, if one allows for loss of
head and fin spines in Rondeletia, its general body form
and relative proportions (e.g., very large head, at least 40%
si) more closely resemble those of Gibberichthys than any
other stephanoberyciform. In addition, we note that the
internal, non-imbricate, cycloid scales of Rondeletia are
similar to those of Gibberichthys, as described by Parr
(1934: 35) “... the squamation, which is on trunk and tail
and consists of thin, but not excessively thin, cycloid scales,
is entirely subcutaneous, i.e., the scale pockets are
completely closed and covered by a thin, generally
transparent, continuous sheet of epidermis without openings
of any kind.”
Our investigation of the above character evidence, not
considered by Moore (1993), led us to reject his hypothesis
and to agree with Rosen’s (1973) suggestions that
Rondeletiidae and Gibberichthyidae are sister taxa. Subsequent
discovery that the complex, presumably secretory Tominaga’s
organ is shared uniquely among fishes by these two taxa,
provides remarkably cogent corroboration of this hypothesis,
even in the absence of a formal phylogenetic analysis of the
Stephanoberyciformes, a project that we plan to undertake in
the future. We do note that the apparent sister-group
relationship of the Rondeletiidae and Gibberichthyidae
indicates that “the whalefishes”, Barbourisiidae, Rondeletiidae,
and Cetomimidae, as a group are at best paraphyletic.
It is not our intention to rigorously examine relationships
of these families (that will be the subject of a future study),
but merely to compare features of their larvae. Aside from
common features that characterize the adults, such as
posterior placement of the pelvic, dorsal and anal fins, we
find no evidence in the morphology of the larvae of
Barbourisia and Rondeletia to suggest a close relationship
between these two families. The large precocious pelvic
fins found in larvae of both families are also present in larvae
of the stephanoberyciform families Gibberichthyidae,
Melamphaidae, and Stephanoberycidae.
The Gibberichthyidae have a distinctive “kasidoron”
larva characterized most notably by an elaborate arborescent
appendage that is an extension of the third pelvic-fin ray
and a papillose epithelium (Robins & de Sylva, 1965). The
figures of larval and juvenile Gibberichthys (Robins & de
Sylva, 1965: fig. 1; Thorp, 1969: figs. 2, 3; de Sylva &
Eschmeyer, 1977: figs. 1-3) indicate there is little space on
the snout anterior to the eye for the presence of Tominaga’s
organ that exceeds 10% of si in similar sized Rondeletia.
Our examination of a 7 mm Gibberichthys confirms the
short snout length at this size, but we have not dissected
this paratype specimen, de Sylva & Eschmeyer (1977) also
mentioned scale rows under the papillate lateral line of
Kasidoron (= Gibberichthys), but did not indicate which of
their four specimens (7.8, 12.1, 15.7, 21.2 mm) have them.
Neither vertical rows of papillate superficial neuromasts
nor scales are present in their 7.8 mm paratype. The
similarity of the papillate epidermis of Gibberichthys to
Mirapinna was noted by Robins and de Sylva (1965).
The elongate pelvic-fin ray of Barbourisia is simple and
it is not the third as in Gibberichthys, but the fourth.
424 Records of the Australian Museum (2001) Vol. 53
Although de Sylva & Eschmeyer (1977) discussed a
distinctive, multibranched postlarval pelvic fin in the
melamphaid genus Poromitra, they did not illustrate it and
we have not seen a detailed description of this feature nor
have we observed it in an actual specimen. None of the
described melamphaid larvae (Keene & Tighe, 1984: figs.
205-207) share distinctive characters with Barbourisia or
Rondeletia, and in all the pelvic fins are much farther
forward. Larval Acanthochaenus (the only described
stephanoberycid larva) are unremarkable with the exception
of the bright violet coloration of fresh specimens (Kotlyar
& Evseyenko, 1989). They share with both Barbourisia and
Rondeletia enlarged posterior pelvic fins that, like those of
Rondeletia , are heavily pigmented and lack elongate rays.
Scales form relatively early, between 8.7-11.2 mm, and are
spinous like those of the adult. The body is more heavily
pigmented than the larvae of either Barbourisia or
Rondeletia. Larval Cetomimidae remain unknown.
We conclude that the larval morphology of the
stephanoberyciform fishes, as presently known, provides
little evidence to elucidate the phylogenetic relationships
of this relatively diverse and highly specialized group.
Unfortunately, the larvae described to date are either
relatively unspecialized or exhibit autapomorphic
specializations. It is likely that larval and small juvenile
specimens will be useful in clarifying structural homology
of problematic characters such as the plate-like dorsal-fin
“spines” of Gibberichthys and the additional “supraneural”
elements of that genus, Barbourisia and Rondeletia.
Acknowledgments. Specimens and collection data were kindly
provided by T. Clarke (University of Hawaii), W. Eschmeyer
(CAS), E. Fujii (TH), K. Hartel (MCZ), R. Lavenberg and J. Seigel
(LACM), J. Leis and M. McGrouther (AMS), K. Matsuura
(NSMT), N. Merrett (BMNH), B. Mundy (National Marine
Fisheries Service, Honolulu), J. Nielsen (ZMUC), A. Powell
(Center for Coastal Fisheries and Habitat Resources, Beaufort,
NC), and R. Rosenblatt and H. Walker (SIO). J. Burns (George
Washington University) sectioned Tominaga’s organ and provided
interpretation of the histology. B. Washington (USNM) drew Fig.
Id; all other figures were done by T. Trnski. Y. Tominaga (TZM)
and I. Wales (AMS) translated Japanese, while W. Ivantsoff (MU)
translated a Russian article. R. Rosenblatt (SIO) and H.G. Moser
(National Marine Fisheries Service, La Jolla) examined specimens
for us. K. Clements (Auckland University) brought the reference
on the trout magnetoreceptor to our attention. Comments by two
referees and the Associate Editor improved the manuscript. Funds
for research were provided by the Australian Research Council,
the Australian Museum Trust, the Smithsonian Institution Research
Opportunity and Susan Lieber Ericson Funds. To all go our
appreciation.
References
Abe, T., & H. Hotta, 1963. Description of a new deep sea fish of
the genus Rondeletia from Japan. Japanese Journal of
Ichthyology 10(2/6): 43^-8.
Baldwin, C.C., & G.D. Johnson, 1995. Alarva of the Atlantic flashlight
fish, Kryptophanaron alfredi , (Beryciformes: Anomalopidae), with
a comparison of beryciform and stephanoberyciform larvae.
Bulletin of Marine Science 56(1): 1-24.
Bast, H.-D., & M.B. Klinkhardt, 1990. Records of the redmouth
whalefish, Rondeletia loricata Abe & Hotta, 1963
(Osteichthyes: Cetomimiformes: Rondeletiidae), from the
northeast and southwest Atlantic. Archive fur Fischerei-
wissenschaften 40(3): 249-263.
Bertelsen, E., 1951. The ceratioid fishes, ontogeny, taxonomy,
distribution and biology. Dana-Report 39: 1-276.
Bertelsen, E., & N.B. Marshall, 1984. Mirapinnatoidei:
development and relationships. In Ontogeny and Systematics
of Fishes, H.G. Moser, W.J. Richards, D.M. Cohen, M.P. Fahay,
A.W. Kendall & S.L. Richardson (eds). American Society of
Ichthyologists and Herpetologists Special Publication 1: 380-
383.
Boehlert, G.W., & B.C. Mundy, 1992. Distribution of ichthyo-
plankton around southeast Hancock Seamount, central north
Pacific, in summer 1984 and winter 1985: data report. National
Oceanic and Atmospheric Administration Technical Memor¬
andum, National Marine Fisheries Service, Southwest Fisheries
Science Center 176: 1-109.
Colgan, D.C., C.-G. Zhang & J.R. Paxton, 2000. Phylogenetic
investigations of the Stephanoberyciformes and Beryciformes,
particularly whalefishes (Euteleostei: Cetomimidae), based on
partial 12S rDNA and 16S rDNA sequences. Molecular
Phylogenetics and Evolution 17: 15-25.
de Sylva, D.P., & W.N. Eschmeyer, 1977. Systematics and biology
of the deep-sea fish family Gibberichthyidae, a senior synonym
of the family Kasidoroidae. Proceedings of the California
Academy of Sciences 49(6): 215-231.
Diebel, C.E., R. Proksch, C.R. Green, P. Nellson & M.M. Walker,
2000. Magnetite defines a vertebrate magnetoreceptor. Nature
406(20 July 2000): 299-302.
Ebeling, A.W., & W.H. Weed, 1973. Order Xenoberyces
(Stephanoberyciformes). In Fishes of the Western North
Atlantic, D.M. Cohen, A.W. Ebeling, T. Iwamoto, S.B.
McDowell, N.B. Marshall, D.E. Rosen, P. Sonoda and W.H.
Weed (eds). Sears Foundation for Marine Research Memoir
1(6): 397^478.
Paxton et alBarbourisia and Rondeletia larvae 425
Goode, G.B., & T.H. Bean, 1895. On Cetomimidae and
Rondeletiidae, two new families of bathybial fishes from the
northwestern Atlantic. Proceedings of the United States
National Museum 17(1012): 451-454.
Greenwood, P.H., D.E. Rosen, S.H. Weitzman & G.S. Myers, 1966.
Phyletic studies of teleostean fishes, with a provisional
classification of living forms. Bulletin of the American Museum
of Natural History 131(4): 339-456, figs.
Harry, R.R., 1952. Deep-sea fishes of the Bermuda Oceanographic
Expeditions, families Cetomimidae and Rondeletiidae.
Zoologica (N.Y.) 37(1): 55-72.
Herring, P, 1976. Carotenoid pigmentation of whale fishes. Deep-
sea Research 23: 235-238.
Johnson, G.D., & C. Patterson, 1993. Percomorph phylogeny: a
survey of acanthomorphs and a new proposal. Bulletin of
Marine Science 52(1): 554-626.
Keene, M.J., & K.A. Tighe, 1984. Beryciformes: development
and relationships. In Ontogeny and Systematics of Fishes. H.G.
Moser, W.J. Richards, D.M. Cohen, M.P Fahay, A.W. Kendall
& S.L. Richardson (eds). American Society of Ichthyologists
and Herpetologists Special Publication 1: 383-392.
Kotlyar, A.N., 1995. Osteology and distribution of Barbourisia
rufa (Barbourisiidae). Voprosy Ikhtiologii 35(3): 282-289. (In
Russian, English transl. Journal of Ichthyology 35(6): 140-
150)
Kotlyar, A.N., 1996. Osteology, intraspecific structure, and
distribution of Rondeletia loricata (Rondeletiidae). Voprosy
Ikhtiologii 36(2): 154-168. (In Russian, English transl. Journal
of Ichthyology 36(3): 207-221)
Kotlyar, A.N., & S.A. Evseyenko, 1989. Larvae of the pricklefish
Acanthochaenus luetkeni (Stephanoberycidae) from the
southwest Pacific. Voprosy Ikhtiologii 29(5): 848-852. (In
Russian, English transl. Journal of Ichthyology 29(8): 102-
107)
Leviton, A.E., R.H. Gibbs, E. Heal & C.E. Dawson, 1985.
Standards in herpetology and ichthyology: part I. Standard
symbolic codes for institutional resource collections in
herpetology and ichthyology. Copeia 1985(3): 802-832.
Moore, J.A., 1993. The phylogeny of the Trachichthyiformes
(Teleostei: Percomorpha). Bulletin of Marine Science 52(1):
114-136.
Nelson, J.S., 1994. Fishes of the World, ed. 3. John Wiley & Sons,
New York, pp. 1-523.
Parr, A.E., 1929. A contribution to the osteology and classification
of the orders Iniomi and Xenoberyces. Occasional Papers of
the Bingham Oceanographic Collection 2: 1-45.
Parr, A.E., 1934. Report on experimental use of a triangular trawl
for bathypelagic collecting. Bulletin of the Bingham
Oceanographic Collection 4(6): 1-59.
Parr, A.E., 1945. Barbourisidae, a new family of deep sea fishes.
Copeia 1945(3): 127-129.
Patterson, C., & G.D. Johnson, 1995. The intermuscular bones
and ligaments of teleostean fishes. Smithsonian Contributions
to Zoology 559: 1-83.
Paxton, J.R., 1974. Morphology and distribution patterns of the
whalefishes of the family Rondeletiidae. Journal of the Marine
Biological Association of India 15(1): 175-188.
Paxton, J.R., 1989. Synopsis of the whalefishes (family
Cetomimidae) with descriptions of four new genera. Records
of the Australian Museum 41(2): 135-206.
Pietsch, T.W., 1984. Lophiiformes: development and relationships.
In Ontogeny and Systematics of Fishes, H.G. Moser, W.J.
Richards, D.M. Cohen, M.P. Fahay, A.W. Kendall & S.L.
Richardson (eds). American Society of Ichthyologists and
Herpetologists Special Publication 1: 320-325.
Potthoff, T., & J.A. Tellock, 1993. Osteological development of
the snook, Centropomus undecimalis (Teleostei, Centro-
pomidae). Bulletin of Marine Science 52 (2): 669-716.
Robins, C.R., & D.P. de Sylva, 1965. The Kasidoroidae, a new
family of mirapinniform fishes from the western Atlantic
Ocean. Bulletin of Marine Science 15(1): 189-201.
Rofen, R.R., 1959. The whale-fishes: families Cetomimidae,
Barbourisiidae and Rondeletiidae (order Cetunculi). Galathea
Reports 1: 255-260.
Rosen, D.E., 1973. Interrelationships of higher euteleostean fishes.
In Interrelationships of Fishes, PH. Greenwood, R.S. Miles
& C. Patterson (eds). Zoological Journal Linnean Society of
London 53 (Supplement 1): 397-513.
Rosen, D.E., & C. Patterson, 1969. The structure and relationships
of the paracanthopterygian fishes. Bulletin of the American
Museum of Natural History 141(3): 357-474.
Struhsaker, R, 1965. The whalefish Barbourisia rufa (Cetunculi)
from waters off southeastern United States. Copeia 1965(3):
376-377.
Thorp, C.H., 1969. A new species of mirapinnaform fish (family
Kasidoroidae) from the western Indian Ocean. Journal of
Natural History 3(1): 61-70.
Tominaga, Y., 1970. On the glandular organs before the eyes of
the red-coated whalefish, Rondeletia loricata. Zoological
Magazine (Tokyo) 79(11-12): 368. (In Japanese).
Yang, Y.R., B.G. Zeng & J.R. Paxton, 1988. Additional specimens
of the deepsea fish Hispidoberyx ambagiosus (Hispido-
berycidae, Beryciformes) from the South China Sea, with
comments on the family relationships. UO 38: 3-8.
Manuscript received 5 December 2000, revised 28 February 2001 and
accepted 7 March 2001.
Associate Editor: J.M. Leis.
Records of the Australian Museum (2001) Volume 53. ISSN 0067-1975
CONTENTS
Volume 53 • Numbers 1-3 • 2001
Andriashev, Anatoly P., see under Stein.341
Barbara Baehr, see under Jocque.21
Barker, J.S.F., see under McEvey.255
Bohlke, Eugenia B., & John E. McCosker. The moray eels of Australia and New Zealand,
with the description of two new species (Anguilliformes: Muraenidae).71
p. 71 http://www.amonline.net.au/pdf/publications/1325.pdf
pp. 71-102 http://www.amonline.net.au/pdf/publications/1325_complete.pdf
Chernova, Natalia V., see under Stein.341
Crane, Martin J., James L. Sharpe & Peter A. Williams. Formation of chrysocolla and
secondary copper phosphates in the highly weathered supergene zones of some Australian
deposits.49
p. 49 http://www.amonline.net.au/pdf/publications/1323.pdf
pp. 49-56 http://www.amonline.net.au/pdf/publications/1323_complete.pdf
Donnellan, Stephen, see under Mahony.37
Edgecombe, Gregory D. Revision of Paralamyctes (Chilopoda: Lithobiomorpha: Henicopidae),
with six new species from eastern Australia.201
p. 201 http://www.amonline.net.au/pdf/publications/1328.pdf
pp. 201-241 http://www.amonline.net.au/pdf/publications/1328_complete.pdf
Foster, Ralph, see under Mahony.37
Glover, Emily A., & John D. Taylor. Systematic revision of Australian and Indo-Pacific
Lucinidae (Mollusca: Bivalvia): Pillucina, Wallucina and descriptions of two new genera
and four new species.263
p. 263 http://www.amonline.net.au/pdf/publications/1349.pdf
pp. 263-292 httpV/www.amonline.net.au/pdf/publications/l349_complete.pdf
Holmes, W.B. Keith. Equisetalean plant remains from the Early to Middle Triassic of New
South Wales, Australia.9
p. 9 http://www.amonline.net.au/pdf/publications/1320.pdf
pp. 9-20 http://www.amonline.net.au/pdf/publications/1320_complete.pdf
Horne, Michelle L. A new seahorse species (Syngnathidae: Hippocampus ) from the Great
Barrier Reef.243
p. 243 http://www.amonline.net.au/pdf/publications/1330.pdf
pp. 243-246 httpV/www.amonline.net.au/pdf/publications/l330_complete.pdf
Hosoya, Seiichi, see under Iwata.103
Iwata, Akihisa, Seiichi Hosoya & Helen K. Larson. Paedogobius kimurai, a new genus and
species of goby (Teleostei: Gobioidei: Gobiidae) from the west Pacific.103
p. 103 http://www.amonline.net.au/pdf/publications/1326.pdf
pp. 103-112 http://www.amonline.net.au/pdf/publications/1326_complete.pdf
Jocque, Rudy, & Barbara Baehr. Revisions of genera in the Asteron -complex (Araneae:
Zodariidae). Asteron Jocque and the new genus Pseudasteron .21
p. 21 http://www.amonline.net.au/pdf/publications/1321.pdf
pp. 21-36 http://www.amonline.net.au/pdf/publications/1321_complete.pdf
Johnson, G. David, see under Paxton.407
Knowles, Ross, see under Mahony.37
Kuiter, Rudie H. Revision of the Australian seahorses of the genus Hippocampus
(Syngnathiformes: Syngnathidae) with descriptions of nine new species.293
p. 293 http://www.amonline.net.au/pdf/publications/1350.pdf
pp. 293-340 httpV/www.amonline.net.au/pdf/publications/l350_complete.pdf
Larson, Helen K., see under Iwata.103
428 Records of the Australian Museum (2001)
Mahony, Michael, Ross Knowles, Ralph Foster & Stephen Donnellan. Systematics of the
Litoria citropa (Anura: Hylidae) complex in northern New South Wales and southern
Queensland, Australia, with the description of a new species.37
p. 37 http://www.amonline.net.au/pdf/publications/1322.pdf
pp. 37-48 http://www.amonline.net.au/pdf/publications/1322_complete.pdf
McAlpine, D.K. Review of the Australasian genera of signal flies (Diptera: Platystomatidae).113
p. 113 http://www.amonline.net.au/pdf/publications/1327.pdf
pp. 113-199 http://www.amonline.net.au/pdf/publications/1327_complete.pdf
McCosker, John E., see under Bohlke. 71
McEvey, Shane F., & J.S.F. Barker. Scaptodrosophila aclinata : a new Hibiscus flower-breeding
species related to S. hibisci (Diptera: Drosophilidae).255
p. 255 http://www.amonline.net.au/pdf/publications/1332.pdf
pp. 255-262 http://www.amonline.net.au/pdf/publications/1332_complete.pdf
McLay, C.L.. A new genus and two new species of unusual dromiid crabs (Brachyura:
Dromiidae) from northern Australia.1
p. 1 http://www.amonline.net.au/pdf/publications/1319.pdf
pp. 1-8 http://www.amonline.net.au/pdf/publications/1319_complete.pdf
Paxton, John R., G. David Johnson & Thomas Trnski. Larvae and juveniles of the deepsea
“whalefishes” Barbourisia and Rondeletia (Stephanoberyciformes: Barbourisiidae,
Rondeletiidae), with co mm ents on family relationships.407
p. 407 http://www.amonline.net.au/pdf/publications/1352.pdf
pp. 407-425 httpV/www.amonline.net.au/pdf/publications/l352_complete.pdf
Sharpe, James L., see under Crane .49
Stein, David L., Natalia V. Chernova & Anatoly P. Andriashev. Snailfishes (Pisces:
Liparidae) of Australia, including descriptions of thirty new species.341
p. 341 http://www.amonline.net.au/pdf/publications/1351.pdf
pp. 341-406 httpV/www.amonline.net.au/pdf/publications/l35 l_complete.pdf
Taylor, John D., see under Glover .263
Timms, Brian V. Two new species of fairy shrimp (Crustacea: Anostraca: Thamnocephalidae:
Branchinella ) from the Paroo, inland Australia.247
p. 247 http://www.amonline.net.au/pdf/publications/1331.pdf
pp. 247-254 http://www.amonline.net.au/pdf/publications/1331_complete.pdf
Trnski, Thomas, see under Paxton .407
Watson, William. Larvae of Enchelyurus ater (Gunther, 1877) and E. kraussi (Klunzinger,
1871) (Pisces: Blenniidae: Omobranchini).57
p. 57 http://www.amonline.net.au/pdf/publications/1324.pdf
pp. 57-70 http://www.amonline.net.au/pdf/publications/1324_complete.pdf
Williams, Peter A., see under Crane .49
Records of the Australian Museum 429
Records of the Australian Museum, Supplement 26 (2001)
Ahyong, Shane T., 2001. Revision of the Australian Stomatopod Crustacea. Records of the
Australian Museum, Supplement 26, pp. 326. Sydney: Australian Museum. Published
4 July 2001. ISBN 0 7347 2303 2.
p. 1 http://www.amonline.net.au/pdf/publications/1333.pdf
pp. 1-326 http://www.amonline.net.au/pdf/publications/1333_complete.pdf
Records of the Australian Museum, Supplement 27 (2001)
Anderson, Atholl, and Peter White, eds., 2001. The Prehistoric Archaeology of Norfolk
Island, Southwest Pacific. Records of the Australian Museum, Supplement 27, pp.
vi+141. Sydney: Australian Museum. ISBN 0 7347 2305 9.
p. v http://www.amonline.net.au/pdf/publications/1334.pdf
Anderson, Atholl, and Roger Green, 2001. Domestic and religious structures in the Emily
Bay settlement site, Norfolk Island. In The Prehistoric Archaeology of Norfolk Island,
Southwest Pacific, ed. Atholl Anderson and Peter White, pp. 43-51. Records of the
Australian Museum, Supplement 27. Sydney: Australian Museum.43
p. 43 http://www.amonline.net.au/pdf/publications/1338.pdf
pp. 43-51* http://www.amonline.net.au/pdf/publications/1338_complete.pdf
Anderson, Atholl, Tom Higham and Rod Wallace, 2001. The radiocarbon chronology of the
Norfolk Island archaeological sites. Ibid. pp. 33-42.33
p. 33 http://www.amonline.net.au/pdf/publications/1337.pdf
pp. 33-42* http://www.amonline.net.au/pdf/publications/1337_complete.pdf
Anderson, Atholl, Ian Smith and Peter White, 2001. Archaeological fieldwork on Norfolk
Island. Ibid. pp. 33-42.11
p. 11 http://www.amonline.net.au/pdf/publications/1336.pdf
pp. 11-32* http://www.amonline.net.au/pdf/publications/1336_complete.pdf
Anderson, Atholl, and Peter White, 2001. Approaching the prehistory of Norfolk Island.
Ibid. pp. 1-9.1
p. 1 http://www.amonline.net.au/pdf/publications/1335.pdf
pp. 1-9* http://www.amonline.net.au/pdf/publications/1335_complete.pdf
Anderson, Atholl, and Peter White, 2001. Prehistoric settlement on Norfolk Island and its
Oceanic context. Ibid. pp. 135-141 .135
p. 135 http://www.amonline.net.au/pdf/publications/1348.pdf
pp. 135-141 *http://www.amonline.net.au/pdf/publications/1348_complete.pdf
Anderson, Atholl, see under Turner.53
Anderson, Atholl, see also under Schmidt.67
Anderson, Atholl, see also under Hold away.85
Anderson, Atholl, see also under Matisoo-Smith.81
Anderson, Atholl, see also under Walter.101
Anderson, Atholl, see also under Macphail.123
Campbell, Colin R., and Lyn Schmidt, 2001. Molluscs and echinoderms from the Emily Bay
settlement site, Norfolk Island. Ibid. pp. 109-114.109
p. 109 http://www.amonline.net.au/pdf/publications/1345.pdf
pp. 109-114*http://www.amonline.net.au/pdf/publications/1345_complete.pdf
Clark, Geoffrey, see under Smith.75
Fullagar, Richard, see under Turner.53
Fullagar, Richard, see under Schmidt.67
Green, Roger, see under Anderson.43
Higham, Tom, see under Anderson.33
Hold away, Richard N., and Atholl Anderson, 2001. Avifauna from the Emily Bay settlement
site, Norfolk Island: a preliminary account. Ibid. pp. 85-100.85
430 Records of the Australian Museum (2001)
p. 85 http://www.amonline.net.au/pdf/publications/1343.pdf
pp. 85-100* http://www.amonline.net.au/pdf/publications/1343_complete.pdf
Hope, Geoffrey S., see under Macphail .123
Horsburgh, K. Ann, see under Matisoo-Smith. 81
Macphail, Mike K., Geoffrey S. Hope and Atholl Anderson, 2001. Polynesian plant
introductions in the Southwest Pacific: initial pollen evidence from Norfolk Island.
Ibid. pp. 123-134.123
p. 123 http://www.amonline.net.au/pdf/publications/1347.pdf
pp. 123-134*http://www.amonline.net.au/pdf/publications/1347_complete.pdf
Matisoo-Smith, Elizabeth, K. Ann Horsburgh, Judith H. Robins and Atholl Anderson, 2001.
Genetic variation in archaeological Rattus exulans remains from the Emily Bay settlement
site, Norfolk Island. Ibid. pp. 81-84.81
p. 81 http://www.amonline.net.au/pdf/publications/1342.pdf
pp. 81-84* http://www.amonline.net.au/pdf/publications/1342_complete.pdf
Neuweger, Diana, Peter White and Winston F. Ponder, 2001. Land snails from Norfolk Island
sites. Ibid. pp. 115-122.115
p. 115 http://www.amonline.net.au/pdf/publications/1346.pdf
pp. 115-122*http://www.amonline.net.au/pdf/publications/1346_complete.pdf
Ponder, Winston F., see under Neuweger.115
Robins, Judith H., see under Matisoo-Smith.81
Schmidt, Lyn, Atholl Anderson and Richard Fullagar, 2001. Shell and bone artefacts from
the Emily Bay settlement site, Norfolk Island. Ibid. pp. 67-74.67
p. 67 http://www.amonline.net.au/pdf/publications/1340.pdf
pp. 67-74* http://www.amonline.net.au/pdf/publications/1340_complete.pdf
Schmidt, Lyn, see also under Campbell.109
Smith, Ian, see under Anderson.11
Smith, Ian, Geoffrey Clark and Peter White, 2001. Mammalian and reptilian fauna from
Emily and Cemetery Bays, Norfolk Island. Ibid. pp. 75-79 .75
p. 75 http://www.amonline.net.au/pdf/publications/1341.pdf
pp. 75-79* http://www.amonline.net.au/pdf/publications/1341_complete.pdf
Turner, Marianne, Atholl Anderson and Richard Fullagar, 2001. Stone artefacts from the
Emily Bay settlement site, Norfolk Island. Ibid. pp. 53-66 .53
p. 53 http://www.amonline.net.au/pdf/publications/1339.pdf
pp. 53-66* http://www.amonline.net.au/pdf/publications/1339_complete.pdf
Wallace, Rod, see under Anderson.33
Walter, Richard, and Atholl Anderson, 2001. Fishbone from the Emily Bay settlement site,
Norfolk Island. Ibid. pp. 101-108.101
p. 101 http://www.amonline.net.au/pdf/publications/1344.pdf
pp. 101-108*http://www.amonline.net.au/pdf/publications/1344_complete.pdf
White, Peter, see under Anderson .v
White, Peter, see also under Anderson.1
White, Peter, see also under Anderson.11
White, Peter, see also under Smith.75
White, Peter, see also under Neuweger.115
White, Peter, see also under Anderson.135
* scheduled for release in June 2002—six months after date of publication.
Publication dates
Records of the Australian Museum 53(1): 1-112.9 May 2001
Records of the Australian Museum 53(2): 113-262. 12 September 2001
Records of the Australian Museum 53(3): 263-430. 12 December 2001
Records of the Australian Museum, Supplement 26: 1-326.4 July 2001
Records of the Australian Museum, Supplement 27: 1-141 ....28 November 2001
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