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Memoirs of
Museum Victoria

Volume 67 31 December 2010

Cover images

Front top left : Enhanced colour photo of preserved specimen of Taeniogyrus australianus (Stimpson, 1855), showing wheel cluster papillae
(large) and hook clusters (small) (27 mm long; Australia, Sydney Harbour, 1968; AM J16377).

Front top right : SEM image for specimen of Taeniogyrus australianus (Stimpson, 1855) (Long Reef, Collaroy, Sydney; RBINS IG 31 459 ex
AM J20086).

Front lower: Metacrangon spinidorsalissp. nov., holotype, female (cl 10.4 mm), WAM C45115, entire animal in lateral view. Scale bar = 5 mm.
Described by Tomoyuki Komai and Joanne Taylor.

Back top left: Hydrobiosella nandawar sp. nov., lateral view. Described by David I. Cartwright.

Back top right: A new cucumberfish of the genus Paralaupus (family Paraulopidae) from the Tasman Sea, described in this issue by Martin
Gomon.

Back lower: Osteichthyan lower jaw. Gyroptychius specimen NRM PZ PI 409.

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MELBOURNE AUSTRALIA

Volume 67
31 December, 2010

Chief Executive Officer
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Robin Hirst

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Richard Marchant

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Martin F. Gomon
David J. Holloway
Gary C. B. Poore
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Mi

MUSEUMVICTORIA

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Museum Victoria 47: 201-206.

Last, PR., and Stevens, J.D. 1994. Sharks and rays of Australia.

CSIRO Publishing: Melbourne. 513 pp.

Wilson, B.R., and Allen, G.R. 1987. Major components and
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Contents

Volume 67 2010

1 > Studies of Australian Hydrobiosella Tillyard : a review of the Australian species of the Hydrobiosella
bispina Kimmins group (Trichoptera: Philopotamidae)

David I. Cartwright

1 5 > A new species of Paraulopus (Aulopiformes: Paraulopidae) from seamounts of the Tasman Sea
Martin F. Gomon

19 > Taxonomic revision of the genus Ratabulus (Teleostei: Platycephalidae), with descriptions of
two new species from Australia
Martin F. Gomon and Hisashi Imamura

35 > Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia
Timothy Flolland

45 > Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda:
Caridea) from Australia
Tomoyuki Komai and Joanne Taylor

61 > A revision of Antarctic and some Indo- Pacific apodid sea cucumbers (Echinodermata:
Holothuroidea: Apodida)

P. Mark O’Loughlin and Didier VandenSpiegel

Memoirs of Museum Victoria 67: 1-13 (2010)

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Studies of Australian Hydrobiosella Tillyard: a review of the Australian species of
the Hydrobiosella bispina Kimmins group (Trichoptera: Philopotamidae)

David I. Cartwright

13 Brolga Crescent, Wandana Heights, Victoria, 3216, Australia (cartwright@hotkey.net.au)

Abstract Cartwright, D.1. 2010. Studies of Australian Hydrobiosella Tillyard: a review of the Australian species of the Hydrobiosella

bispina Kimmins group (Trichoptera: Philopotamidae). Memoirs of Museum Victoria 67: 1-13.

Descriptions and keys are provided for males of 12 philopotamid caddis fly species in the genus Hydrobiosella
Tillyard, H. bispina group. Among these are ten new species from Australia: Hydrobiosella bilga, H. dugarang, H. gurara,
H. moorda, H. mundagurra, H. nandawar, H. thurawal, H. unispina, H. woonoongoora and H. yokunna. Females of six
of the species are also described. Group separation is based on male genitalic characteristics, the key features for the
H. bispina group being inferior appendages with an elongate terminal segment, fringed ventrally by a row of dark setae,
and nearly all species with paired, hooked lateral processes on segment ten. Of the 12 species treated here, all are endemic
to the east coast region of the Australian mainland.

Keywords Trichoptera, caddis flies, Philopotamidae, Hydrobiosella, Australia

Introduction

Records of Hydrobiosella Tillyard in Australia are relatively
recent; the genus was first described in 1924 for a New Zealand
species, H. stenocerca Tillyard. Only in 1953 were the first
Australian species recognised in the genus, with the transfer of
the southern Western Australian species H. michaelseni
(Ulmer 1908) from Dolophilus and description of H. arcuata
Kimmins from southeastern Queensland, H. bispina Kimmins
from New South Wales, and H. cognata Kimmins, H.
tasmanica Mosely and H. waddama Mosely all from Tasmania
(in Mosely and Kimmins, 1953). Subsequently, the following
species were added: H. letti Korboot (1964) from New South
Wales, H. armata Jacquemart (1965), H. anasina, H. cerula,
H. corinna, H. orba and H. sagitta all from Tasmania (Neboiss,
1977), and H. amblyopia Neboiss (1982) from southern
Western Australia. Neboiss (2003) added six more Tasmanian
species, H. anatolica, H. disrupta, H. otaria, H.propinqua, H.
scalaris and H. tahunense, bringing the total Australian
species of Hydrobiosella to 20.

Ross (1956) recognised Hydrobiosella as a subgenus of
Sortosa Navas, but during his work on Tasmanian caddis flies,
Neboiss (1977) reinstated it to full species status. At the same
time, he placed the Tasmanian species in three groups — the
H. corinna group and the H. tasmanica group each with four
species; and H. waddama with only one species. In 2003,
Neboiss added a further six species to the H. tasmanica group,
bringing the Tasmanian Hydrobiosella total to 15 species.
Although the Tasmanian Hydrobiosella have been well
studied, until now, little effort has been expended on the

mainland Australian members of the genus — only five species
are presently recorded. Among them, two in the group are
being studied here: H. arcuata and H. bispina, and one of the
new species described below was cited in a checklist as
Hydrobiosella sp. nov. PT-2029 (Walker et al., 1995).

Henderson (1983), in his systematic study of New Zealand
philopotamids, discussed the wider relationships between
New Caledonian and Australian Hydrobiosella species. He
found that synapomorphies of the New Zealand species of
Hydrobiosella are not consistent in the known Australian
species, thus concluding that the Australian species cannot be
placed as a monophyletic group in his classification. Finally,
he commented ‘too little is known at present for a full
understanding of the relationships of this family’, especially
‘the lack of detailed information on the Australian and South
American fauna . . This present revision is the first of a series
in which all the Australian Hydrobiosella groups will be
revised. Once key characters of the different groups are
identified, relationships of the Australian groups with other
philopotamid groups, particularly species in New Zealand and
New Caledonia, can be assessed.

In this taxonomic revision of the Australian Hydrobiosella
bispina group, about 250 male and female specimens were
examined and referred to 12 species. The most common
species, H. woonoongoora, represented about 34% of the total
specimens examined, H. unispina about 22% and H. bispina
about 21%. Six of the 12 species are known from fewer than
five specimens. The 12 species, including the ten new species,
were collected from southern and eastern Australia. All except

2

D.l. Cartwright

two species (from northeastern Queensland) are from the
Bassian region, which is suggestive of a ‘southern’ origin, a
thesis supported by the fact that the genus is known only from
Australia, New Zealand and New Caledonia.

Methods and abbreviations

Among Hydrobiosella species — size, body and wing colour
can be useful characters, but are variable. Colour can be a
useful character in live or freshly preserved material but, with
time, it often fades in alcohol. All the H. bispina group
specimens were stored in alcohol, and many for over 20 years.
Most of the material studied was on loan from Museum
Victoria and made available by Dr Arturs Neboiss. Depositories
for specimens are abbreviated as follows: Museum Victoria,
Melbourne (NMV), Australian National Insect Collection,
Canberra (ANIC) and the Natural History Museum, London
(BMNH). All specimens mentioned in the text, including
types, are lodged in the NMV unless stated otherwise.

Males of each species are readily distinguished by genitalic
features, but often require clearing of the abdomen in
potassium hydroxide. Females were paired with respective
males on the basis of similarities in size and colouration, and
on locality.

Figured specimens are identified by the notebook numbers
of Dr Arturs Neboiss (prefix PT-) or the author (prefix CT-).
Terminology generally follows that of Neboiss (1977, 1982),
Blahnik (2005) and Holzenthal et al. (2007). However, past
authors have used a variety of names for the same structures as
outlined by Munoz-Quesada and Holzenthal (2008, p. 8). For
example, in this study, ‘harpago’ is used instead of apical or
terminal segment of the inferior appendages, and ‘preanal
appendage’ is used instead of superior appendage or cercus.
Abbreviations for genitalic parts are indicated on selected
figures . Typically, setae or spines are illustrated only on the right
side of the figure (as viewed) to enable a better depiction of the
underlying structures . Length and width measurements generally
refer to the maximum length divided by maximum width.

Descriptions

Hydrobiosella Tillyard

Hydrobiosella Tillyard 1924: 288; Mosely and Kimmins 1953:
387; Neboiss 1977: 45.

Type species. Hydrobiosella stenocerca Tillyard by monotypy.

Generic descriptions are given by Tillyard (1924: 288), Mosely
and Kimmins (1953: 387) and Neboiss (1977: 45).

Key to males of known Australian groups (or ungrouped
species) of Hydrobiosella Tillyard

1. Phallus without pair of parameres (Neboiss 1986, figs pp.
99; H. amblyopia, 101; H. armata, H. tasmanica, 102; H.
orb a, H. corinna) 2

- Phallus with pair of parameres (Neboiss 1986, figs pp. 99;
H. michaelseni, H. waddama, 101; H. letti, 102; H.bispina,
103; H. arcuata ) 4

2. Preanal appendages present, usually small (Neboiss 1977,
figs 204-211; Neboiss 1986, figs p. 102 H. orba, H.
corinna ; Neboiss 2003, figs 8a-h); Tas H. corinna group

- Preanal appendages absent (Neboiss 1986, figs pp. 99 H.

amblyopia, 101 H. armata, H. tasmanica ) 3

3. Phallus apically with downward projecting spine(s)
(Neboiss 1977, figs 216-221, 225, 226; Neboiss 1986, figs
p. 101 H. armata, H. tasmanica-, Neboiss 2003, figs lOa-g,

lla-g, 1 2a— f): Tas

H. tasmanica group

- Phallus apically without downward projecting spine(s)
(Neboiss 1982, fig. 12; Neboiss 1986, figs p. 99 H.

amblyopia ); S-WA

H. amblyopia (ungrouped)

4. Inferior appendages with harpago with dark row of setae
forming fringe along ventral margin (figs 2-4; Neboiss
1986, figs pp. 102 H.bispina, 103 H. arcuata ); E-Vic,

E-NSW, E-Qld

Hydrobiosella bispina group

- Inferior appendages with harpago without dark row of

setae forming fringe along ventral margin (Neboiss 1986,
figs pp. 99 H. michaelseni, H. waddama, 101 H. letti ) 5

5. Parameres elongate and sinusoidal, attached ventrally to
base of phallus (Neboiss 1977, fig. 233; Neboiss 1986, figs
p. 99 H. waddama-, Neboiss 2003, figs 12g, h); Tas, SE

Aust.

r r rj H. waddama (group — only one species described)

- Parameres not elongate and sinusoidal, not attached
ventrally to base of phallus (Neboiss 1982, figs 9, 10;
Neboiss 1986, figs pp. 99 H. michaelseni, 101 H. letti ) .... 6

6. Parameres curved strongly and crossed (Neboiss 1982,
figs 9, 10; Neboiss 1986, figs p. 99 H. michaelseni ); S-WA

H. michaelseni (ungrouped)

- Parameres not curved strongly and crossed (Neboiss
1986, figs p. 101 H. letti)-, CE-NSW . //. letti (ungrouped)

Hydrobiosella bispina group

Diagnosis. Key features of males in the group are inferior
appendages with the harpago elongate, often angled near
middle, with a dark row of setae forming a fringe along ventral
margin; segment X with a pair of lateral lobes, which usually
end in small hooks.

Description. Head and nota dorsally brown to dark brown with
setal warts and scutellum pale, abdomen brownish dorsally and
ventrally, paler laterally; wings light brown to brown. Medium
sized adults. Forewing length, males: 5.9-8. 8 mm; females:
6. 1-8.7 mm; forewing length about 2.9-3.0 times maximum
width, wing venation similar to the type species H. stenocerca,
R1 simple, forks 1, 2, 3, 4 and 5 present; forks 1 and 2 sessile;
fork 2 with nygma, about 1.6-1 .7 times length fork 1; fork 3
shorter, length 0.6-0.7 times length fork 2, fork 3 length about

A review of the Australian species of the Hydrobiosella bispina group

2.0 times length footstalk, cross-veins r-m and m contiguous or
nearly meeting at fork 3; fork 4 similar length to fork 3, fork
length about 4 times length footstalk; fork 5 very long, length
between 1.7-1 .8 times length fork 4; discoidal cell closed,
length about 4.5 times maximum width. Hind wing length
about 2. 5-2 .6 times maximum width, with forks 1, 2, 3 and 5
present; fork 1 usually sessile, occasionally with very short
footstalk; fork 2 sessile, nygma present, fork 2 length between
1.3—1 .4 times fork 1 length; fork 3 length about 0.5-0.6 times
fork 2 length, fork 3 similar length to footstalk; fork 5 very
long, length between 2. 1-2.2 times length fork 3; discoidal cell
closed, length between 4.5 times maximum width; with three
anal veins (fig. 1).

Male. Segment IX usually with a small rounded notch
medially on ventrodistal margin (figs 7, 13), rarely without
(figs 10, 19). Preanal appendages absent. Segment X mainly
sclerotised with a central pale, mostly membranous mesal
lobe, with one or two pairs of short hairs subapically (figs
5-6); with a pair of more pigmented lateral lobes, which
usually end in small hooks (figs 8-9). Phallus generally tube-
like, slightly dilated subapically, with a pair of slender, straight
or slightly curved parameres arising from the phallus
basolaterally (figs 2-3, 5-6). Inferior appendages two
segmented, in lateral view, basal segment robust, harpago
more slender, straight to sharply angled near middle (figs 3, 6).

Female. Genitalia typical of genus, sometimes with a
small projection, which can be diagnostic, on sternite VIII
mesodistally (figs 38-49).

Larva. Confirmed larvae are unknown, although
Hydrobiosella spp AV8 and AV15 (Cartwright, 1997) almost
certainly belong to this group. Hydrobiosella sp. AV8 larvae
have been recorded mainly in riffle habitats from small to
medium streams 2-13 m wide at low to moderate altitudes
between 70-1200 m (Suter et al. 2006). These larvae have the
forecoxa with two sclerotised processes on the anterior margin
and the anterior margin of the frontoclypeus convex.

Remarks. The 12 species in this group are known from eastern
mainland Australia, ranging from northeastern Queensland to
eastern Victoria (latitudinal range 16°35'-37°18'S). Females of
only six species have been associated.

Key to males of species of the Australian Hydrobiosella
bispina group

1. Inferior appendages with harpago straight or with ventral

margin forming a weakly obtuse angle (figs 3, 6) 2

- Inferior appendages with harpago neither straight nor

with ventral margin forming a weakly obtuse angle, but
forming almost a right angle (figs 21, 24, 36) 7

2. Segment X with a dorsal spine (Fig. 3); NE-Qld

H. unispina

- Segment X without a dorsal spine (figs 6, 9) 3

3. Segment X without subapical, lateral pair of hooks (figs

5-6); CE-NSW H. gurara

3

- Segment X with subapical, lateral pair of hooks (figs 8-9,

11-12) 4

4. Segment X in dorsal view with apex rounded, slightly

bulbous and dorsoventrally flattened (figs 8-9); NE-NSW
H. nandawar

- Segment X in dorsal view with apex not rounded and

dorsoventrally flattened, slender and laterally compressed
(figs 11-12, 14-15) 5

5. Segment X with robust subapical, lateral pair of hooks,

apex slender in lateral view (figs 11-12); NE-Qld

//. dugarang

- Segment X without robust subapical, lateral pair of hooks,

apex not slender in lateral view (figs 14-15, 17-18) 6

6. Inferior appendages with harpago dilated slightly in

apical half (fig. 15); E-Vic H. bilga

- Inferior appendages with harpago not dilated slightly in

apical half (fig. 18); C-Qld H. mundagurra

7. Parameres very long, reaching tip of inferior appendages

(figs 20-21); E-NSW H. bispina

- Parameres not very long, not reaching tip of inferior

appendages (figs 23-24) 8

8. Inferior appendages with harpago curved strongly in

distal half so apex is pointing downwards (fig. 24); SE-
Qld H. arcuata

- Inferior appendages with harpago not curved strongly in

distal half so apex is pointing posteriorly (figs 27, 30) 9

9. Segment X in dorsal view with apex tapered gradually,

not constricted sub-apically (fig. 26); CE-NSW

H. moorda

- Segment X in dorsal view with apex not tapered gradually,

constricted subapically (figs 29, 32) 10

10. Segment X in lateral view with apex slightly bulbous, not

curved downwards; subapical, lateral pair of hooks directed
outwards (figs 29-30); SE-Qld H. woonoongoora

- Segment X in lateral view with apex not slightly bulbous,

curved slightly downwards; subapical, lateral pair of hooks
directed downwards (figs 32-33, 35-36) 11

11. Inferior appendages with basal segment tapered strongly

distally; harpago not dilated slightly in distal half (fig. 33);
CE-NSW H.thurawal

- Inferior appendages with basal segment not tapered

strongly distally; harpago dilated slightly in distal half
(fig. 36); CE-NSW H. yokunna

Hydrobiosella unispina sp. nov.

Figures 2-4, 38-39

Holotype. Male, Queensland, Mt Spec State forest. Camp Ck trib.,

18°57'S, 146° 10'E, 760 m, 11 Jun 1994, A. L. Sheldon (NMV,T- 20893).
Paratypes. Queensland. 1 male (specimen PT-2029 figured), Mt

4

D.l. Cartwright

Spec, at light, 11 May 1975, R. Storey and D. Hancock; the following
sites all Mt Spec State forest, 18°57'S, 146°10'E, A. L. Sheldon; 1 male.
Birthday Ck above weir, 820 m, 6 Dec 1993; 1 male. Camp Ck proper,
760 m, 11 Jun 1994; 1 male, 1 female (specimen CT-635 figured).
Camp Ck trib., 760 m, 15 May 1994; 1 male, same site, 5 Dec 1993; 1
male, 1 female, same site, 15 Mar 1994; 1 male, same site, 12 Dec
1993; 2 males, 1 female, same site, 6 Jul 1994, 1 male, same site, 6 Nov
1993; 1 male, same site, 23 Apr 1994; 2 males, same site, 21 Nov 1993;
1 male, same site, 20 Dec 1993; 1 male, same site, 15 Oct 1993; 1 male,
same site, 4 Mar 1994 (NMV).

Other material examined. Queensland. 1 female. Upper Little
Mossman R., Mt Lewis, 10 Dec 1974, M. S. Moulds; 1 male, ‘top of the
range’, 19 Butler Drive, Kuranda, 335 m, 16°48'S, 145°38'E, 1-15 Leb
2007, D. C. L. Rentz (ANIC); the following sites all Mt Spec State
forest, 18°57’S, 146°10'E, A. L. Sheldon: 1 male, 3 females, unnamed
ck, Paluma Dam Rd, 860 m, 17 Jan 1994; 1 male, same site, 11 Jun
1994; 1 male, same site, 6 Jul 1994; 1 male, unnamed ck ‘cascade’, 920
m, 17 May 1994; 2 males, ‘Confusion’ Ck, trib. to unnamed ck, Paluma
Res., 17 May 1994; 2 females. Birthday Ck above weir, 820 m, 20 Dec
1993; 2 males, 1 female, same site, 22 Oct 1993; 1 male, same site, 6
Nov 1993; 2 females, same site, 13 Nov 1993; 1 female, same site, 21
Nov 1993; 1 male. Birthday Ck below falls, 760 m, 11 Jun 1994; 1
male. Birthday Ck, Iron Cabin, 790 m, 12 Leb 1994; 1 male. Birthday
Ck, 870 m, 16 Jan 1994; 2 males, 1 female, same site, 31 Oct 1993; 2
males, Williams Ck trib., 745 m, 13 Nov 1993; 2 males, same site, 15
May 1994; 1 female. Camp Ck trib., 760 m, 13 Dec 1993; 1 female.
Echo Ck trib., 735 m, 7 Nov 1993 (NMV).

Diagnosis. Hydrobio sella unispina can be separated from other
species in the group by the dorsal spine on segment X and segment
IX produced into a triangular point medially on distal margin.

Description. Wings similar to those of H. arcuata (fig. 1),
length of forewing: male 6. 7-8.0 mm, female 7.2-8 .7 mm.

Male. Segment IX without a noticeable notch on mesodistal
margin, instead produced into a triangular point (fig. 4).
Segment X with mesal lobe broadbased, dorsoventrally
compressed in distal two-thirds, with a mesodorsal spine (figs
2-3); in dorsal view, subtriangular or tongue shaped, tapered
distally, length about 2.3 times maximum width, with a pair of
lateral lobes, without projecting hooks (fig. 2); in lateral view
slender, downcurved in distal two-thirds (fig. 3). Inferior
appendages in lateral view, with basal segment subrectangular,
length about 1.8-1 .9 times maximum width; harpago nearly as
long as basal segment, more slender, length about 3.5 times
width, weakly (obtusely) angled near middle (fig. 3).

Female. Genitalia typical of genus, with a small triangular
projection on sternite VIII mesodistally (figs 38-39).

Etymology. Unispina — Latin for ‘one spine’ (spine on tergum

xj.

Remarks. Hydrobiosella unispina is a common species and has
been collected mainly from the Mt Spec area of northeastern
Queensland (latitudinal range 16°35'-18°57'S).

Hydrobiosella gurara sp. nov.

Figures 5-7

Holotype. Male (specimen CT-577 figured). New South Wales,
Jerusalem Falls near Karuah (about 32°39'S, 151°57'E), 6 Dec 1988, G.
Theischinger (NMV, T-20913).

Diagnosis. Hydrobiosella gurara can be separated from other
species in the group by the absence of lateral subapical hooks
on segment X and from H. unispina by the absence of a dorsal
spine on segment X.

Description. Wings similar to those of H. arcuata (fig. 1),
length of forewing: male 6.5 mm.

Male. Segment IX with a deep notch on mesodistal margin
(fig. 7). Segment X with robust mesal lobe, broadbased, with a
pair of pigmented lateral lobes, without hooks (figs 5-6); in
dorsal view subtriangular, tapered slightly distally with a
rounded apex (fig . 5) . Phallus robust (fig . 6) . Inferior appendages
in lateral view, with basal segment subquadrate, length about
1 .5 times maximum width; harpago longer than basal segment,
length about 1 .3-1 .4 times length basal segment, slender, length
about 5 .5 times width, narrowed and weakly (obtusely) angled
near middle, slightly dilated in apical third (fig. 6).

Female. Unknown.

Etymology. Gurara — Australian Aboriginal (New South
Wales) word for ‘long’ or ‘tall’ (harpago on inferior appendages).

Remarks. The holotype male is the only specimen of
Hydrobiosella gurara collected from the type locality in
central -eastern New South Wales (latitude 32°39'S).

Hydrobiosella nandawar sp. nov.

Figures 8-10

Holotype. Male, New South Wales, Mt Kaputar, 30°16'S, 150°10'E, 3
Jan 1986, G. Theischinger (NMV, T-20914)

Paratype. New South Wales. 1 male (specimen CT-428 figured),
Mt Kaputar Nat. Pk, Dawson Springs, 9 Oct 1973, A. Neboiss (NMV).

Diagnosis. Hydrobiosella nandawar can be separated from
other species in the group by the rounded, slightly bulbous and
dorsoventrally flattened apex on segment X.

Description. Wings similar to those of H.arcuata (fig. 1), length
of forewing: male 7.6-8.0 mm.

Male. Segment IX without a noticeable notch medially on
distal margin (fig. 10). Segment X mesal lobe with distal third
slightly dorsoventrally compressed (figs 8-9); in lateral view,
distal third slender, straight (fig. 9); in dorsal view, distal third
narrowed subapically, slightly bulbous and rounded apically
(fig. 8), with a pair of pigmented lateral lobes, which terminate
in small, slightly downward projecting hooks (figs 8-9).
Inferior appendages in lateral view, with basal segment length
about twice maximum width, broadest in basal third, tapered
slightly distally; harpago shorter than basal segment, length
about 0.8 times length basal segment, slender, length about 4.5
times width, weakly (obtusely) angled near middle (fig. 9).
Female. Unknown.

Etymology. Nandawar — Australian Aboriginal name for ‘Mt
Kaputar’ (type locality).

Remarks. Two male specimens of Hydrobiosella nandawar
have been collected from the Mt Kaputar National Park in
northeastern New South Wales (latitude 30°16'S).

A review of the Australian species of the Hydrobiosella bispina group

5

barpago

Figures 1-7. Hydrobiosella spp.; 1, Hydrobiosella arcuata Kimmins, wings; 2-7, Hydrobiosella spp., male genitalia in dorsal, lateral and part ventral
views; 2-4, Hydrobiosella unispina sp. nov.; 2, dorsal; 3, lateral; 4, ventral, mesodistal margin of segment IX; 5-7, Hydrobiosella gurara sp. nov.; 5,
dorsal; 6, lateral; 7, ventral, mesodistal margin of segment IX.

6

D.l. Cartwright

Figures 8-16. Hydrobiosella spp. male genitalia in dorsal, lateral and part ventral views; 8-10, Hydrobiosella nandawar sp. nov.; 8, dorsal; 9,
lateral; 10, ventral, mesodistal margin of segment IX; 11-13, Hydrobiosella dugerang sp. nov.; 11, dorsal; 12, lateral; 13, ventral, mesodistal
margin of segment IX; 14-16, Hydrobiosella bilga sp. nov.; 14, dorsal; 15, lateral; 16, ventral, mesodistal margin of segment IX.

A review of the Australian species of the Hydrobiosella bispina group

Hydrobiosella dugarang sp. nov.

Figures 11-13

Holotype. Male (specimen CT-560 figured), Queensland, Dalrymple
Ck near Eungella, 21°02'S, 148°43'E, 3 Apr 1993, G. Theischinger
(NMV, T-20915).

Diagnosis. Hydrobiosella dugarang can be separated from
other species in the group by the robust subapical, lateral pair
of hooks and slender apex, both on segment X in lateral view.

Description. Wings similar to those of H.arcuata (fig. 1), length
of forewing: male 6.2 mm.

Male. Segment IX with a small notch medially on distal
margin (fig. 13). Segment X with slender mesal lobe (figs
11-12); in lateral view, slightly downcurved apically (fig. 12);
in dorsal view, slender, not narrowed subapically (fig. 11), with
a pair of more pigmented lateral lobes, which terminate in
robust, downward projecting hooks (figs 11-12). Inferior
appendages in lateral view, with basal segment length about
twice maximum width, broadest in basal half, tapered slightly
distally; harpago about same length as basal segment, slender,
straight, length about 5 .7-5 .9 times width (fig. 12).

Female. Unknown.

Etymology. Dugarang — Australian Aboriginal word for
‘straight’ (inferior appendages).

Remarks. The male holotype is the only known specimen of
Hydrobiosella dugarang from the type locality in northeastern
Queensland (latitude 21°02'S).

Hydrobiosella bilga sp. nov.

Figures 14-16

Holotype. Male, New South Wales, Nungatta Ck, Yambula State
Forest (about 37°08'S, 149°29'E), 16-17 Feb 2000, J. Miller (ANIC).

Paratypes. New South Wales. 6 males, collected with holotype
(ANIC). Victoria. 1 male (specimen CT-571 figured). Beehive Ck, 30
km N of Cann River, (about 37°18'S, 149°12'E), 21 Mar 1977, A.
Neboiss (NMV).

Diagnosis. Hydrobiosella bilga can be separated from other
species in the group by the combination of the slightly dilated
apices and weakly obtuse angle on ventral margin of the
harpago and the pair of subapical hooks on segment X.

Description. Wings similar to those of H.arcuata (fig. 1), length
of forewing: male 73-8.4 mm. Forewing fork 2 long, length
fork 2 about 1.5 times length of fork 1; length fork 3 about 1.9
times length footstalk; fork 4 length about 7 times length
footstalk. Hind wing fork 1 sessile; fork 3 length about twice
length of footstalk.

Male. Segment IX with a small shallow notch medially on
distal margin (fig. 16). Segment X mesal lobe with a pair of
short hairs/bristles subapically, slightly laterally compressed; in
dorsal view slender, not narrowed subapically (fig. 14), with a
pair of more pigmented lateral lobes, which terminate in small,
slender, slightly outward and downward projecting hooks (figs
14-15). Inferior appendages in lateral view, with basal segment
length about 1 .8 times maximum width, broad basally, tapered

7

slightly distally; harpago more slender, straight with slightly
convex ventral margin, slightly dilated in apical third (fig. 15).
Female. Unknown.

Etymology. Bilga — Australian Aboriginal word for ‘bee’s
nest’ (type locality — Beehive Creek).

Remarks. Eight male specimens of Hydrobiosella bilga have
been collected from the two localities in southeastern New South
Wales and eastern Victoria (latitudinal range 37°08'-37°18'S).

Hydrobiosella mundagurra sp. nov.

Figures 17-19, 40-41

Holotype. Male, Queensland, Carnarvon Gorge Nat. Pk, 25°15‘S,
148°24‘E, 12 Nov 1990, G. Theischinger (NMV, T-20917).

Paratypes. Queensland. 5 males (specimen CT-575 figured), 15
females (specimen CT-607 figured), collected with holotype (NMV).

Other material examined. Queensland. 16 females, collected with
holotype (NMV).

Diagnosis. Hydrobiosella mundagurra can be separated from
other species in the group by the combination of harpago, which
is straight and not dilated in distal half, and segment X with
slender lateral pair of hooks and slender apex in dorsal view.

Description. Wings similar to those of H. arcuata (fig. 1),
length of forewing: male 5.9-67 mm, female 6.5-87 mm.
Forewing fork 2 long, length fork 2 about 1.5 times length of
fork 1; fork 3 length about 1.6 times length footstalk; fork 4
length about 4.6 times length footstalk. Hind wing fork 1 with
short footstalk; fork 3 length about 1.9 times length of footstalk.

Male. Segment IX without a noticeable notch medially on
distal margin (fig. 19). Segment X mesal lobe with a pair of
short hairs/bristles subapically, slightly laterally compressed; in
dorsal view slender, not narrowed subapically (Fig. 17), with a
pair of more pigmented lateral lobes, which terminate in small,
slender, slightly outward and downward projecting hooks (figs
17-18). Inferior appendages in lateral view, with basal segment
length about 1 .9 times maximum width, broad basally, tapered
slightly distally; harpago more slender, nearly straight (fig. 18).

Female. Genitalia typical of genus, with a small rounded
projection on sternite VIII mesodistally (figs 40-41).

Etymology. Mundagurra — named for the Australian
Aboriginal dreaming rainbow serpent believed to have created
Carnarvon Gorge.

Remarks. Six males and many females of Hydrobiosella
mundagurra have been collected from the type locality in
central-eastern Queensland (latitude 25°15'S).

Hydrobiosella bispina Kimmins
Figures 20-22, 42-43

Hydrobiosella bispina Kimmins in Mosely and Kimmins, 1953:
394, fig. 270,-Neboiss, 1986: 102.

Type material (not seen). Holotype. Male, New South Wales, Stanwell
Park, 23 Apr 1916, R. J. Tillyard (BMNH).

Paratype (not seen). New South Wales. 1 male, collected with
holotype (BMNH).

8

D.l. Cartwright

Figures 17-25. Hydrobiosella spp. male genitalia in dorsal, lateral and part ventral views; 17-19, Hydrobiosella mundagurra sp. nov.; 17, dorsal;
18, lateral; 19, ventral, mesodistal margin of segment IX; 20-22, Hydrobiosella bispina Kimmins.; 20, dorsal; 21, lateral; 22, ventral, mesodistal
margin of segment IX; 23-25, Hydrobiosella arcuata Kimmins; 23, dorsal; 24, lateral; 25, ventral, mesodistal margin of segment IX.

A review of the Australian species of the Hydrobiosella bispina group

Material examined. New South Wales. 3 males, 6 females, Wilson R.,
Wilson R. Reserve, 11 Feb 2008, R. St Clair; 1 male, Wilson R., Bobs
Ridge Rd, 31°15'S, 152°31'E, 4 Dec 2007, A. Glaister, J. Dean and R.
St Clair; 1 male (specimen PT-579 figured), Tubrabucca, Barrington
Tops, 15 Nov 1953, A. Neboiss (NMV); 2 males, 16 females, Dilgry
R., Banksia camp ground, 31°53'S, 151°32'E, 2 Dec 2007, A. Glaister,
J. Dean and R. St Clair; 3 males, 4 females, Gloucester R., Gloucester
R. camping area, 32°03'S, 15F41'E, 1 Dec 2007, A. Glaister, J. Dean
and R. St Clair; 1 male, 1 female (specimen CT-605 figured),
Gloucester Tops, el. 1280 m. Malaise, 19 Nov to 4 Dec 1988, D. Bickle;
3 males, 1 female, Gloucester Tops, 32°04'S, 151°34'E, el. 1300 m, 2-3
Dec 1988, Theischinger and Mueller; 1 male, 1 female, Jerusalem
Falls near Karuah, 6 Dec 1988, G. Theischinger; 1 male, 1 female,
Wilson R. near Bellangry, 5 Dec 1988, G. Theischinger; 1 male,
Wollomi Brook, The Basin, Olney State Forest, 33°06'S, 151 0 14'E, 26
Nov 2007, A. Glaister, J. Dean and R. St Clair (NMV).

Diagnosis. Hydrobiosella bispina can be separated from other
species in the group by the very long parameres, which reach
the tip of the inferior appendages.

Description. (Revised after Kimmins in Mosely and Kimmins,
1953.) Wings similar to those of H.arcuata (fig. 1), length of
forewing: male 6.3-83 mm, female 7.2-8.6 mm. Forewing
fork 2 long, length fork 2 about 1.5 times length of fork 1;
length fork 3 about twice length footstalk; fork 4 length about
8 times length footstalk. Hind wing fork 1 sessile; fork 3 length
about 1.5 times length of footstalk.

Male. Segment IX with small, shallow notch medially on
distal margin (fig. 22). Segment X with a slender mesal lobe,
with a pair of short hairs/bristles subapically, in lateral view
slightly downturned distally (fig. 21); in dorsal view slightly
narrowed subapically (fig. 20); with a pair of pigmented lateral
lobes, which end in small, slightly downward and outward
projecting rounded hooks (figs 20-21). Phallus generally
slender, slightly dilated subapically; with a pair of very slender
and elongate parameres arising from the phallus near the apex
(fig. 21). Inferior appendages in lateral view, with basal
segment length about twice maximum width, broadest near
middle, tapered strongly distally; harpago more slender, with
ventral margin sharply angled at about 90 degrees near middle,
tapered slightly distally (fig. 21).

Female. Genitalia typical of genus, with a small, shallow,
triangular projection on sternite VIII mesodistally (figs
42-43).

Remarks. Males and females of Hydrobiosella bispina have
been collected from nine sites in addition to the type locality,
all in eastern New South Wales (latitudinal range 31° 15'-
33°06'S).

Kimmins’ (in Mosely and Kimmins 1953) and Neboiss’
(1986) figures have been redrawn to allow direct comparisons
and to accompany the description that is revised in light of
new interpretations of Hydrobiosella genitalic structures.

Hydrobiosella arcuata Kimmins
Figures 1, 23-25, 44-45

Hydrobiosella arcuata Kimmins in Mosely and Kimmins, 1953:
397, fig. 271. -Neboiss, 1986: 103.

9

Type material (not seen). Holotype. Male, Queensland, Montville, 3
Oct 1912, R. J. Tillyard (BMNH).

Material examined. Queensland. 1 male, Booloumba Ck, 8 km
SW Kenilworth, 26°39’S, 152°39'E, 12 Dec 1984, G. Theischinger; 1
male (specimen CT-573 figured), Booloumba Ck, Mary R. catchment,
26°41'S, 152°37’E, 26 Oct 1993, collector unknown; 1 male, 1 female.
Branch Ck, Brisbane R. catchment, 26°52’S, 152°41'E, 26 Apr 1993;
collector unknown; 1 male, 2 females (specimen CT-603 figured).
Stony Ck, Brisbane R. catchment, 26°52’S, 152°43’E, 18 Aug 1992;
collector unknown (NMV).

Diagnosis. Hydrobiosella arcuata can be separated from other
species in the group by the shape of the harpago, where the
ventral margin is curved or arched strongly so that the apex
points downwards.

Description. (Revised after Kimmins in Mosely and Kimmins,
1953). Wings similar to other species in the group (fig. 1), length
of forewing: male 6.3-73 mm, female 6. 1-8.0 mm. Forewing
fork 2 long, length fork 2 about 1 .6 times length of fork 1 ; length
fork 3 about twice length footstalk; fork 4 length about 4.5 times
length footstalk. Hind wing fork 1 sessile or with very short
footstalk; fork 3 length about 1 .0-1 .3 times length of footstalk.

Male. Segment IX with shallow notch medially on distal
margin in between a pair of small knobs (fig. 25). Segment X
with a slender mesal lobe, with a pair of short hairs/bristles
subapically, in lateral view slightly downturned distally (fig.
24); in dorsal view not narrowed subapically (fig. 23), with a
pair of more pigmented lateral lobes, which terminate in small,
slightly backward and outward projecting hooks (figs 23-24).
Phallus truncate apically, with a pair of robust parameres arising
from the phallus subapically (fig. 24). Inferior appendages in
lateral view, with basal segment length about 1 .7 times maximum
width, broad basally, rounded distally; harpago more slender,
with ventral margin sharply angled near middle, curved in distal
half with downward pointing acute apex (fig. 24).

Female. Genitalia typical of genus, with a small, shallow,
rounded projection on sternite VIII mesodistally (figs 44-45).

Remarks. Males and females of Hydrobiosella arcuata have
been collected from five sites in addition to the type locality, all
in southeastern Queensland (latitudinal range 26°39'-26°52'S).

Kimmins’ (in Mosely and Kimmins 1953) and Neboiss’
(1986) figures have been redrawn to allow direct comparisons
and to accompany the description that is revised in light of
new interpretations of Hydrobiosella genitalic structures.

Hydrobiosella moorda sp. nov.

Figures 26-28

Holotype. Male, New South Wales, (about 33°41'S, 150°17'E), Pulpit
Hill Ck, Megalong Valley, 8 Oct 1985, A. Neboiss (NMV, T-20938).

Paratypes. New South Wales. 2 males (specimen PT-1421 figured),
collected with holotype (NMV).

Diagnosis. Hydrobiosella moorda can be separated from other
species in the group by the combination of segment X in dorsal
view with a robust, gradually tapered apex, not constricted
subapically, and the ventral margin of the harpago sharply
angled at about 90 degrees near middle with apex pointing
posteriorly.

10

D.l. Cartwright

Figures 26-34. Hydrobiosella spp. male genitalia in dorsal, lateral and part ventral views; 26-28, Hydrobiosella moorda sp. nov.; 26, dorsal; 27,
lateral; 28, ventral, mesodistal margin of segment IX; 29-31, Hydrobiosella woonoongoora sp. nov.; 29, dorsal; 30, lateral; 31, ventral, mesodistal
margin of segment IX; 32-34, Hydrobiosella thurawal sp. nov.; 32, dorsal; 33, lateral; 34, ventral, mesodistal margin of segment IX.

A review of the Australian species of the Hydrobiosella bispina group

Description. Wings similar to those of H.arcuata (fig. 1), length
of forewing: male 8.0-8.8 mm. Forewing fork 2 long, length
fork 2 about 1.5 times length of fork 1; length fork 3 about 1.5
times length footstalk; fork 4 length about 5.3 times length
footstalk. Hind wing fork 1 sessile; fork 3 length about 1.3— 1.4
times length of footstalk.

Male. Segment IX with a small shallow notch medially on
distal margin (fig. 28). Segment X with a robust mesal lobe,
with a pair of short hairs/bristles subapically, in lateral view
not downturned distally (fig. 27); in dorsal view, tapered
slightly distally, not narrowed subapically (fig. 26); with a pair
of more pigmented lateral lobes that end in small, slightly
downward and outward projecting hooks (figs 26-27). Inferior
appendages in lateral view, with basal segment length about

l. 9 times maximum width, broadest near middle, rounded
distally; harpago more slender, ventral margin sharply angled
at about 90 degrees near middle (fig. 27).

Female. Unknown.

Etymology. Moorda — Australian Aboriginal word for ‘blue
mountain’ (type locality — Blue Mountains).

Remarks. Three male specimens of Hydrobiosella moorda
have been collected from the type locality in central-eastern
New South Wales (latitude 33°41'S).

Hydrobiosella woonoongoora sp. nov.

Figures 29-31, 46, 47

Holotype. Male, New South Wales, Upper Crystal Ck at Crystal Ck
rainforest retreat, 28°15'S, 153°18'E, 25 Dec 2006, A. Wells (ANIC).

Paratypes. New South Wales. 20 males, 16 females (specimen CT-
636 figured), collected with holotype (ANIC). Queensland. 4 males
(specimen CT-567 figured) Coomera Ck, Lamington Nat. Pk, 8 Feb
1961, F. A. Perkins (NMV).

Other material examined. New South Wales. 12 males, collected
with holotype (ANIC); 1 male, 4 females, same site and collector, 26
Dec 2006 (ANIC); 1 male, 1 female, same site and collector, 24 Dec
2006 (ANIC). Queensland. 4 females, Coomera Ck, Famington Nat. Pk,
8 Feb 1961, F. A. Perkins; 4 males, 1 female, Famington Nat. Pk, 28 Jan
1963, G. Monteith; 1 male, 1 female, Famington Nat. Pk, 27 May 1959,
collector unknown; 1 male (damaged), Binna Burra, 22 May 1964, B.
Genn (NMV); 1 male, 1 female, Binna Burra, Famington Nat. Pk, 750

m, 28°11'S, 153° 1 l’E, 10 Nov 1988, E. S. Nielsen and M. Horak
(ANIC) ; 4 males, 3 females, Binna Burra, 28° 1 2'S, 1 53° 1 1 'E, Famington
Nat. Pk, 3-10 Nov 1984, E. D. Edwards; 1 male, ‘Gwingamma’, 6 km
SW of Tallebudgera, 28°11'S, 153°23'E, 18-23 Apr 1994, Malaise trap,
Rentz, Fee, Upton (ANIC); 2 males. Redwood Park, Toowoomba,
27°35'S, 151°59'E, 8 Nov 1988, E. S. Nielsen and M. Horak (ANIC).

Diagnosis. Hydrobiosella woonoongoora can be separated
from other species in the group by small differences in segment
X; the mesal lobe in has a slightly bulbous apex and the lateral
lobes have apical hooks directed outwards.

Description. Head, body and wings brownish, some specimens
paler. Wings similar to those of H.arcuata (fig. 1), length of
forewing: male 6.4— 8.1 mm, female 6. 6-8 .2 mm. Forewing fork

2 long, length fork 2 about 1 .6 times length of fork 1 ; length fork

3 about 1.9 times length footstalk; fork 4 length about 4.6-5 .4
times length footstalk. Hind wing fork 1 with very short
footstalk; fork 3 length about twice length of footstalk.

11

Male. Segment IX with small shallow notch medially on
distal margin (fig. 31). Segment X with mesal lobe slightly
bulbous apically, with a pair of short hairs/bristles subapically,
in lateral view not downturned distally (fig. 30); in dorsal view,
narrowed slightly subapically (fig. 29), with a pair of pigmented
lateral lobes, which terminate in small, outward projecting
hooks (figs 29-30). Inferior appendages in lateral view, with
basal segment length about twice width, broad basally, rounded
distally; harpago more slender, with ventral margin sharply
angled at about 90 degrees near middle, very slightly dilated
in apical third (fig. 30).

Female. Genitalia typical of genus, with a small, rounded
projection on sternite VIII mesodistally (figs 46-47).

Etymology. Woonoongoora — Australian Aboriginal word for
the Lamington National Park ranges (type locality —
Lamington National Park).

Remarks. Many male and female specimens of Hydrobiosella
woonoongoora have been collected from five sites near the
type locality in northeastern New South Wales and southeastern
Queensland (latitudinal range 27°35'-28°15'S).

Hydrobiosella thurawal sp. nov.

Figures 32-34, 48-49

Holotype. Male, New South Wales, Minnamurra Falls (about 34°38'S,
150°44'E), ?12 Aug 1967, N. Hynes and joint collector unknown
(NMV, T-20945).

Paratype. New South Wales. 1 male (specimen PT-580 figured), 1
female (specimen CT-604 figured), Minnamurra Falls, W of Kiama,
25 Mar 1973, A. Neboiss (NMV).

Diagnosis. Hydrobiosella thurawal can be separated from
other species in the group by small differences in segment X
and the inferior appendages; segment X in lateral view, with
apex not bulbous, curved slightly downwards; lateral lobes
with apical hooks directed downwards and not outwards;
inferior appendages with basal segment tapered strongly
distally; harpago with ventral margin sharply angled at about
90 degrees, not dilated in distal half.

Description. Wings similar to those of H.arcuata (Fig. 1),
length of forewing: male 6.9-8. 5 mm, female 8.7 mm. Forewing
fork 2 long, length fork 2 about 1.6 times length of fork 1;
length fork 3 about twice length footstalk; fork 4 length about
5.8 times length footstalk. Hind wing fork 1 with very short
footstalk; fork 3 length about 2.4 times length of footstalk.

Male. Segment IX with small, shallow notch medially on
distal margin (fig. 34). Segment X mesal lobe, with a pair of short
hairs/bristles subapically, in lateral view slightly downturned
distally (fig. 33); in dorsal view, narrowed subapically, very
slightly bulbous apically (fig. 32), with a pair of pigmented
lateral lobes, which terminate in small, downward projecting
hooks (figs 32-33). Inferior appendages in lateral view, with
basal segment length about 1 .8 times maximum width, broadest
near middle, tapered distally, harpago more slender, ventral
margin sharply angled at about 90 degrees near middle (fig. 33).

Female. Genitalia typical of genus, with a small, shallow
projection on sternite VIII mesodistally (figs 48-49).

12

D.l. Cartwright

Figures 35-49. Hydrobiosella spp.; 35-37, Hydrobiosella yokunna sp. nov. male genitalia in dorsal, lateral and part ventral views; 35, dorsal; 36,
lateral; 37, ventral, mesodistal margin of segment IX; 38-49, Hydrobiosella spp. female genitalia (part segment VIII) in lateral and (segment
VIII) ventral view; 38-39, Hydrobiosella unispina sp. nov.; 38, lateral; 39, ventral; 40-41, Hydrobiosella mundagurra sp. nov.; 40, lateral; 41,
female genitalia, ventral; 42-43, Hydrobiosella bispina Kimmins; 42, lateral; 43, ventral; 44-45, Hydrobiosella arcuata Kimmins; 44, lateral;
45, ventral; 46-47, Hydrobiosella woonoongoora sp. nov.; 46, lateral; 47, ventral; 48-49, Hydrobiosella thurawal sp. nov.; 48, lateral; 49, ventral.

A review of the Australian species of the Hydrobiosella bispina group

Etymology. Thurawal — Australian Aboriginal name for the
area around the type locality.

Remarks. Two males and one female specimen of Hydrobiosella
thurawal have been collected from the type locality in central-
eastern New South Wales (latitude 34°38'S).

Hydrobiosella yokunna sp. nov.

Figures 35-37

Holotype. Male (specimen CT-574 figured). New South Wales, Tuckers
Knob, Orara West State Forest, 29°41'S, 152°48'E, 22 Nov 1990, G.
Theischinger (NMV, T-20948).

Diagnosis. Hydrobiosella yokunna can be separated from
other species in the group by small differences in segment X
and the inferior appendages; segment X in lateral view, with
apex not bulbous, curved slightly downwards; lateral lobes
with apical hooks directed downwards and not outwards;
inferior appendages with basal segment not tapered strongly
distal ly; harpago with ventral margin sharply angled at about
90 degrees, dilated slightly in distal half.

Description. Wings similar to those of H.arcuata (fig. 1), length
of forewing: male 6.9-8. 5 mm. Forewing fork 2 long, length
fork 2 about 1.5 times length of fork 1; length fork 3 about 1.5
times length footstalk; fork 4 length about 7.8 times length
footstalk. Hind wing fork 1 with very short footstalk; fork 3
length about 1.2 times length of footstalk.

Male. Sternite IX with shallow notch medially on distal
margin in between a pair of small knobs (fig. 37). Tergum X
mesal lobe, with a pair of short hairs/bristles subapically, in
lateral view slightly downturned distally (fig. 36); in dorsal
view, narrowed subapically, slightly bulbous apically (fig. 35);
with a pair of pigmented lateral lobes, which terminate in
small, downward projecting hooks with slightly truncate
apices (figs 35-36). Inferior appendages in lateral view, with
basal segment length about 1.7 times maximum width,
broadest near middle, rounded distally; harpago more slender,
narrowed and with ventral margin angled at about 90 degrees
near middle, slightly dilated in apical third (fig. 36).

Female. Unknown.

Etymology. Yokunna — Australian Aboriginal word for
‘crooked’ or ‘bent’ (terminal segment of inferior appendages).

Remarks. The holotype male is the only specimen of
Hydrobiosella yokunna collected from the type locality in
central -eastern New South Wales (latitude 29°41'S).

Acknowledgements

I thank the Australian Government Department of the
Environment, Water, Heritage and the Arts, in particular
Australian Biological Resources Study (ABRS) National
Taxonomy Research Grant Program, for providing a grant to
undertake this work; Drs Arturs Neboiss and Alice Wells for
providing access to the specimens and together with John

13

Dean, for helpful advice on earlier drafts of this manuscript;
and the referee(s), for constructive comments on this
manuscript. I am indebted to John Dean and Ros St Clair for
technical assistance with scanning the figures and moral
support during the project.

References

Blahnik, R. J. 2005. Alterosa, a new caddisfly genus from Brazil
(Trichoptera: Philopotamidae). Zootaxa 991: 1-60.

Cartwright, D. I. 1997. Preliminary guide to the identification of late
instar larvae of Australian Ecnomidae, Philopotamidae and
Tasimiidae (Insecta: Trichoptera). Identification guide no. 10.
Cooperative Research Centre for Freshwater Ecology, Albury. 33 pp.
Henderson, I. M. 1983. A contribution to the systematic of New
Zealand Philopotamidae (Trichoptera). New Zealand Journal of
Zoology 10: 163-176.

Holzenthal, R. W., Blahnik, R. J„ Prather, A. L. and Kjer, K. M. 2007.
Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. Zootaxa
1668: 639-698.

Jacquemart, S. 1965. Contribution a la connaisance de la faune
Trichopterologique de la Tasmanie et de la Nouvelle-Zealande.
Bulletin de ITnstitut Royal des Sciences Naturelles de Belgique
41: 1-47.

Korboot, K. 1964. Four new species of caddis-flies (Trichoptera) from
eastern Australia. Journal of the Entomological Society of
Queensland 3: 32-41.

Mosely, M. E. and Kimmins D. E. 1953. The Trichoptera (caddis-flies)
of Australia and New Zealand. London: British Museum (Natural
History). 550 pp.

Munoz-Quesada, F. J. and Holzenthal, R. W. 2008. Revision of the
Neartic species of the caddisfly genus Wormaldia McLachlan
(Trichoptera: Philopotamidae). Zootaxa 1838: 1-75.

Neboiss, A. 1977. A taxonomic and zoogeographic study of Tasmanian
caddis-flies (Insects: Trichoptera). Memoirs of the National
Museum of Victoria 38: 1-208.

Neboiss, A. 1982. The caddis-flies (Trichoptera) of south-western
Australia. Australian Journal of Zoology 30: 271-325.

Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific -Australian
Region. Dr W. Junk Publishers, Dordrecht. 286 pp.

Neboiss, A. 2003. New genera and species, and new records, of
Tasmanian Trichoptera (Insecta). Papers and Proceedings of the
Royal Society of Tasmania 136: 43 -82.

Ross, H. H. 1956. Evolution and Classification of the Mountain
Caddisflies. University of Illinois Press, Urbana. 213 pp.

Suter P, Dean J, Cartwright D, Sutcliffe K, Davies P, Pinder A and
Bryce, C. 2006. Habitat Profiles of Selected Australian Aquatic
Insects. Australian Biological Resources Study, Australian
Government Department of the Environment, Water, Heritage and
the Arts, Canberra. Accessed 30 July 2010 at www. environment.
gov.au/biodiversity/abrs/publications/electronic-books/aquatic-
insects.html

Tillyard, R J. 1924. Studies of New Zealand Trichoptera or caddis flies
no. 2. Descriptions of new genera and species. Transactions of the
New Zealand Institute 55: 285-314.

Ulmer, G. 1908. Trichopteridae und Ephemeridae. Pp. 25-46 in
Michaelsen, W. and Hartmeyer. R. (eds). Die Fauna Sud west-
Australiens vol. 2, part 3.

Walker, K., Neboiss, A., Dean, J. and Cartwright, D. 1995. A preliminary
investigation of the caddis-flies (Insecta: Trichoptera) of the
Queensland Wet Tropics. Australian Entomologist 22: 19-3 1 .

Memoirs of Museum Victoria 67: 15-18 (2010)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

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A new species of Paraulopus (Aulopiformes: Paraulopidae) from seamounts of the
Tasman Sea

Martin F. Gomon

Sciences Department, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (mgomon@museum.vic.gov.au)

Abstract Gomon, M.F. 2010. A new species of Paraulopus (Aulopiformes: Paraulopidae) from seamounts of the Tasman Sea.

Memoirs of Museum Victoria 67: 15-18.

A new species of the Paraulopus nigripinnis species complex of the family Paraulopidae is described from four
specimens taken on seamounts and rises in the western, central and eastern parts of the Tasman Sea between 30° and 35°S.
It is distinguishable from other members of the complex in having 8-9 anal fin rays; 19 pectoral fin rays; 48 vertebrae;
19-24 predorsal scales; 5.5 scales above the lateral line; a large pelvic fin (length 22.8-28.4% SL), distal margin of pelvic
fin deeply concave, separating the fin into inner and outer lobes, with the inner lobe much shorter than the outer (the ratio
of the lengths of the outer to the inner is 1 .7-2.1); two prominent, broad brown bands on the side of the body posterior to
the dorsal fin; a broad black marginal stripe covering the distal third of the dorsal fin, with a distinct broad white submarginal
stripe; a white marginal band and a black submarginal band on the distal third of the upper lobe of the caudal fin; a black
marginal band on the ventral lobe of the caudal fin; and the buccal cavity almost entirely black.

Keywords Paraulopidae, Paraulopus, sp. nov., Tasman Sea

Introduction

Scrutiny of museum specimens referrable to the recently
described genus Paraulopus (Sato and Nakabo, 2002a) has
revealed a surprising diversity in Australian and New Zealand
waters (Sato and Nakabo, 2002b; Gomon and Sato, 2004; Sato et
al., 2010). Members of this Indo- West Pacific genus are separable
into two complexes: the Northern Hemisphere and tropical
Paraulopus oblongus complex, and a Paraulopus nigripinnis
complex — so far known only from cool tropical and temperate
Australasian localities. Species of the latter were distinguished
by Sato et al. (2010). Species in the Tasman Sea separating
Australia and New Zealand usually occur, at least in part, at deep-
shelf or upper-slope depths of one of the two countries. Several
specimens of a previously undescribed species in the P.
nigripinnis complex, first collected near Lord Howe Island, are
known from seamounts and rises in the Tasman Sea, but
apparently do not occur in Australian continental waters, and so
far have not been taken within New Zealand’s Exclusive
Economic Zone. A description of that species is presented here.

Materials and methods

Terminology and methodology mostly follow Sato et al.
(2010). The pelvic fin in species with a deeply concave distal
margin are separated into distinct inner and outer lobes; the
lengths of the lobes are measured from the base of the first
pelvic fin ray to the tip of the longest ray of each lobe.

Institutional codes are those of Leviton et al. (1985).

Paraulopus balteatus sp. nov. Banded cucumberfish

Figure 1; table 1

Material examined. Holotype: AMS 1.44606-001 (320), Tasman Sea,
Australia, New South Wales, Browns Mount off Botany Bay 34°02'S,
151°39'E (estimated), 430 m, 17 June 2008, drop line, FV Blue Eye,
collected by Jurgen Konrad and retained by Pascal Geraghty,
Department of Primary Industries, NSW Fisheries.

Paratypes: NMNZ P. 10455 (246), Three Kings Ridge, 30°45.00'S,
173°57.00'E, 537-677 m, 6 July 1962, RV Tui, beam trawl; NMNZ P.
35686 (278), Lord Howe Rise, 34°09.20'S, 162°51.80'E, 365-793 m,
16 September 1998, FV Arrow; NMV A22071 (305), Lord Howe Rise,
33°38'S-33°38'S, 162°2rE-162 0 28'E, 300-750 m, 22 March 2001,
demersal trawl, Ken Smith, MAFRI.

Diagnosis. Anal fin rays 8-9; pectoral fin rays 19; vertebrae 48;
predorsal scales 19-24; scales above lateral line 5.5; pelvic fin
large, length 22.8-28.4% SL, larger in males than females,
inner lobe much shorter than outer lobe, ratio of lengths of
outer lobe to inner 1.7-2. 1; sides with two prominent broad
brown bands posterior to dorsal fin; broad black marginal
stripe covering distal third of dorsal fin with distinct broad
white submarginal stripe; distal third of upper lobe of caudal
fin with white marginal band and black submarginal band and
ventral lobe with black marginal band; buccal cavity pigmented
black forward to jaws; males with black anal fin margin
anteriorly and distally, anal fin of females lacking dark margins.

16

M.F. Gomon

Figure 1. Paraulopus balteatus sp. nov., holotype, AMS 1.44606-001, 320 mm SL, male, Tasman Sea, Australia, New South Wales, Browns
Mount off Botany Bay 34°02'S, 151°39'E (est.), 430 m, photo by S. Humphreys (AMS).

Description. Dorsal fin rays 11; anal fin rays 9; caudal fin rays
1+8+1-I-9-I-8-I-1-I-8-I-1; pectoral fin rays 19; pelvic fin rays 9;
vertebrae 19+29; lateral line scales 48-49; scales above lateral
line 5.5; scales below lateral line 3.5; predorsal scales 19-24;
gill rakers 8-9+18-20 = 27-28. (See table 1 for morphometric
values).

Cigar-shaped body, tapering evenly to narrow caudal
peduncle; anus about midway between pelvic-fin base and anal-
fin origin. Head bluntly pointed, rather cylindrical, not depressed;
dorsal outline of head and nape nearly straight in lateral profile;
snout short. Nostrils ovoid, positioned midway between eye and
tip of snout, subdivided by transverse flap of skin. Superocular
ridge on either side above central half of eye. Eye large,
positioned dorsolaterally, on dorsal profile of head. Posterior
edge of preopercle smooth, curved at angle. Mouth terminal;
dorsoposterior corner of maxilla below centre of eye. Teeth on
jaws fine, in broad strip extending anteriorly onto lateral surfaces
of premaxilla and dentary, tapering to narrow strip posteriorly.
Vomerine teeth fine, in narrow transverse band, continuous with
posteriorly tapering band of teeth on exposed edge of palatine;
hyoid teeth fine, in ovoid patch on each side, axis angled
anteromesially at anterolateral comer of tongue; teeth on lateral
periphery slightly enlarged. Gill rakers on upper arm of first arch
short; those on lower limb moderately long and slender, with one
or two rudimentary rakers at both dorsal and ventral ends of arch.

Scales large, cycloid. Predorsal scales extending forward
to vertical through posterior extent of eye. Cheek scales large,
covering cheek and preopercle, in about three poorly defined
rows. Lateral line positioned midlaterally on side, anterior end
slightly elevated.

Dorsal fin moderately tall with short base, second ray
longest but only slightly longer than first, subsequent rays

decreasing in length; first two rays unbranched, subsequent
rays branched; vertical through origin of fin closer to origin of
pelvic fin than origin of pectoral fin; adipose fin small but
obvious, positioned just in advance of vertical through
posterior end of anal fin base. Anal fin short based, of moderate
height, first ray shortest, length of subsequent rays subequal,
first two unbranched, others branched; anal fin origin closer to
base of tail than to origin of pelvic fin. Caudal fin distinctly
forked, upper lobe slightly longer than lower. Posterior tip of
pectoral fin reaching beyond origin of pelvic fin but not to
vertical through centre of longest ray; fourth ray longest; first
ray simple, others branched. Posterior tip of depressed pelvic
fin reaching about halfway between pelvic fin origin and anal
fin origin; posterior margin distinctly concave; inner ray
distinctly shorter than fin length; first ray unbranched, others
branched; tip of outer lobe of pelvic fin expanded into a fleshy,
pad-like structure.

A large species, largest specimen examined 320 mm SL.

Pigmentation in alcohol. Body dusky dorsally, underside pale,
with two broad brown bands encircling body except ventrally,
first posterior to dorsal fin, second posterior to anal fin, and
several broad brown blotches dorsally on side, one below
posterior half of dorsal fin, second between bands (third just
prior to caudal fin in paratypes). Snout, dorsal part of cheek and
operculum very dark. Buccal lining of mouth black; tongue
black with white tip and lateral margins. Dorsal fin dark brown
basally with broad black marginal stripe occupying distal half
(to third) of fin anteriorly, and broad white submarginal stripe.
Adipose fin dusky. Anal fin white with narrow black margin
anteriorly and distal ly in males; entirely white in females.
Caudal fin dark basally with broad pale vertical intermediate

New Tasman Sea Paraulopus

17

Table 1. Selected proportional measurements and counts for types of Paraulopus balteatus sp. nov.

Holotype

Paratypes ( n

Range

= 3)

Mean ± SD

Standard length (mm)

320

246-305

% SL

Body depth

21.6

17.8-19.7

18.7+1.0

Body width

18.3

15.8-21.5

17.9+3.2

Head length

30.0

30.6-32.6

31.5+1.0

Caudal peduncle depth

8.1

5.8-7.6

6.9+0.9

Caudal peduncle length

19.6

20.2-21.8

20.8+0.8

Predorsal length

37.8

39.2-40.0

39.7+0.4

Preanal length

75.0

71.6-74.4

73.3+1.5

Prepectoral length

30.4

31.3-32.9

31.9+0.9

Prepelvic length

42.5

40.7-43.6

41.8+1.6

Preanus length

55.0

56.1-59.3

58.0+1.7

Pelvic fin origin to anus

15.2

16.4-19.1

17.6+1.4

Anus to anal fin origin

18.5

15.6-16.2

16.0+0.3

Dorsal fin base

15.1

12.8-15.0

14.1+1.1

Dorsal fin height

24.9

21.7-28.3

25.7+3.6

Dorsal fin last ray

8.0-17.0

12.5+6.4

Anal fin base

7.6

6. 7-8 .3

7.4+0. 8

Anal fin height

7.5

5.9-9.7

8. 4+2. 2

Pectoral fin length

22.0

20.8-21.6

21.2+0.4

Pelvic fin length

26.5

22.8-28.4

26.2+3.0

Pelvic fin inner lobe length

15.1

12.6-14.9

13.8+1.2

Interpelvic width

15.0

13.3-16.1

14.2+1.6

% HL

Head depth

57.8

49.7-53.1

51.8+1.8

Orbit diameter

34.9

36.1-38.8

37.7+1.4

Postorbital length

43.2

38.0-40.5

39.3+1.3

Head width

55.9

55.8-57.0

56.5+0.6

Interorbital width

10.9

9.5-11.0

10.2+0.8

Upper jaw length

44.5

43.1-43.8

43.5+0.4

Snout length

26.0

23.2-26.6

24.7+1.8

Adipose fin length

5.7

6. 2-7.4

6. 6+0.7

% pelvic fin length

Pelvic fin inner lobe length

57.0

48.7-55.4

52.7+3.6

Meristic values

Holotype

Range

Dorsal-fin rays

11

11

Anal -fin rays

9

8-9

Pectoral-fin rays

19

19

Pelvic-fin rays

9

9

Gill rakers

8+19-27

8-9 + 18-20 -

27-28

Pored lateral-line scales

47

48-49

Scales above lateral line

5.5

5.5

Scales below lateral line

3.5

3.5

Predorsal scales

24

19-21

Vertebrae

19 + 29

19 + 29

HL = head length; SD = standard deviation; SL = standard length

18

M.F. Gomon

band and narrow white marginal band with black submarginal
band on upper lobe and narrow black marginal band on dorsal
two-thirds of lower lobe. Pectoral fin dusky. Pelvic fin rather
dark with white margin on distal edge and distally along
anterior edge; fine black distal edge (in males only).

Fresh colour. Body bluish-grey above with pearlescent white
underside and iridescent blue sheen; bands and blotches
brownish. Black markings on fins intensely so; other dark areas
(described above) greyish. Pectoral and outer parts of pelvic
fins shaded yellow.

Etymology. The name balteatus, from the Latin for ‘belted or
banded’, in reference to the distinctive broad brown bands on
the sides of the body in this species.

Distribution. Known only from the Tasman Sea at 30-35°S,
from Browns Mount southeast of Botany Bay, New South
Wales, to the Three Kings Ridge just north of the northern
boundary of New Zealand’s EEZ, in depths no shallower than
300 m and no greater than 800 m.

Comments. This species was initially confused with Paraulopus
okamurai (Sato and Nakabo, 2002b), a closely related congener
that occurs in the same area. It attains a similarly large size and
has equally distinctive black-and-white -patterned dorsal and
caudal fins. The pattern of the fins in P. okamurai is the basis
for its New Zealand vernacular name, ‘magpie cucumberfish’.
The two are separable by details of the fin patterns, as well as
the presence in the new species of prominent broad brown
bands on the side of the body of adults (versus, at most, much
smaller brown blotches midlaterally and a smaller brownish
saddle behind the adipose fin), 8-9 anal fin rays (versus
9-11 rays, rarely 9), 19 pectoral fin rays (versus 16-17), and an
entirely black interior of the mouth, except for the tongue,
which has white at the tip and along the lateral edges (versus
entirely pale). The same characteristics also separate
P. balteatus from most other species in the complex. Its pectoral
fin count overlaps only with P. novaeseelandiae (Sato and
Nakabo, 2002b), which rarely has 19 rays; only P. melanostomus
(Sato et al., 2010) has the buccal cavity completely lined with
black, although the throat of some of the others is darkly
pigmented. Many of the species, especially at a small size, have
brownish spots or small blotches along the side, often
horizontally aligned midlaterally, and like P. novaeseelandiae
have a brownish saddle under or behind the adipose fin. None
of those blotches, however, are expanded to broad bands that
nearly encircle the side as in P. balteatus. The new species also
has among the largest pelvic fins of the genus, the length of
which is only matched by P. longianalis (Sato et al., 2010), and
to some extent P. novaeseelandiae, but the inner lobe of the
pelvic fin in P. balteatus is much shorter relative to the outer
lobe than in the others (length of outer lobe relative to inner
lobe 1. 7-2.1 versus 1.3 -1.7).

As in most other members of the P. nigripinnis complex,
sexual dimorphism is quite apparent in this species, involving
the size and colouration of the pelvic fins and pigmentation of
the dorsal and anal fins. Males have a larger pelvic fin (length
26.5-28.4% SL, versus 22.8% SL in females), and the
pigmentation of these fins is much darker in males with a fine

black margin distally. Males also have a fine black margin
anteriorly and distally on the anal fin that is missing in females,
while the dorsal fin has a fine, stark white distal margin that
does not appear to feature in females.

Despite the recent increase of collecting in central latitudes
of the Tasman Sea, this species is known only from four large
specimens, one of which was taken on hook and line. Its
absence from trawl collections may infer a preference by the
species for a hard-bottom habitat, which is usually avoided by
trawl fishers. The fact that P. nigripinnis, P. novaeseelandiae
and P. okamurai have also been taken with hook and line
(Roberts, 1997, 2004; Stewart, 2006; Struthers, National
Museum of New Zealand, pers. comm., March 2010) suggests
that species of this genus are carnivorous. Individuals probably
rest on their substantial pelvic fins waiting for potential prey to
come to them — a behaviour common to a number of other
members of the order.

Acknowledgments

Thanks to K. Graham, who recognised the significance of the
specimen designated as holotype and secured it for the AMS
collection; and K. Smith, MAFRI, Vic DPI, who similarly
conveyed a paratypic specimen to NMV. Assistance in
examining specimens came from M. McGrouther and A. Hay
(AMS), C. Struthers (NMNZ) and D. Bray (NMV). The
photograph of the holotype was taken by S. Humphreys
(AMS). Helpful comments on the manuscript were provided
by C. Roberts. Travel to New Zealand where NMNZ specimens
were examined was funded, in part, by Biosystematics of NZ
EEZ Fishes project (NZ Foundation for Research Science and
Technology contract C01X0502/I02-BBCF).

References

Gomon, M.F. and Sato, T. 2004. A new cucumberfish (Paraulopidae)
of the Paraulopus nigripinnis complex from Central Eastern
Australia. Records of the Australian Museum 56:195-199.
Leviton, A.E., Gibbs, R.H., Jr, Heal, E. and Dawson, C.E. 1985.
Standards in herpetology and ichthyology, part 1. Standard
symbolic codes for institutional resource collections in
herpetology and ichthyology. Copeia 1985:802-832.

Roberts, C.D. 1997. Cucumber fish. New Zealand Fishing News, Dec.
20(12): 48, 1 fig.

Roberts, C.D. 2004. New cucumberfishes described. New Zealand
Fishing News, Mar. 27(3): 44, 1 fig.

Sato, T. and Nakabo, T. 2002a. Paraulopidae and Paraulopus, a new
family and genus of aulopiform fishes with revised relationships
within the order. Ichthyological Research 49:25-46.

Sato, T. and Nakabo, T. 2002b. Two new species of Paraulopus
(Osteichthys: Aulopiformes) from New Zealand and Eastern
Australia, and comparisons with P. nigripinnis. Species Diversity
7:393-404.

Sato, T., Gomon, M.F. and Nakabo, T. 2010. Two new Australian
species of the Paraulopus nigripinnis complex (Aulopiformes:
Paraulopidae). Ichthyological Research 57:254-262.

Stewart A.L. 2006. Duskyfin cucumberfish. New Zealand Fishing
News, Nov. 29(11): 36, 1 fig.

Memoirs of Museum Victoria 67: 19-33 (2010)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

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Taxonomic revision of the genus Ratabulus (Teleostei: Platycephalidae), with
descriptions of two new species from Australia

Hisashi Imamura 1 and Martin F. Gomon 2

1 Laboratory of Marine Biology and Biodiversity (Systematic Ichthyology), Faculty of Fisheries Sciences, Hokkaido
University, 3-1-1 Minato-cho, Hakodate, Hokkaido 041-8611, Japan (imamura@fish.hokudai.ac.jp)

2 Ichthyology, Sciences Department, Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia (mgomon@
museum.vic.gov.au)

Abstract Gomon, M.F. and Imamura, H. 2010. Taxonomic revision of the genus Ratabulus (Teleostei: Platycephalidae), with

descriptions of two new species from Australia. Memoirs of Museum Victoria 67: 19-33.

The platycephlid genus Ratabulus Jordan and Hubbs, 1925 is reviewed taxonomically. The genus is defined by the
long, slender canines on its upper jaw, palatine and vomer, the presence of a small free spine between the two dorsal fins,
the iris lappet broad and simple dorsally, the suborbital ridge with numerous spines, the suborbitals and preopercle lacking
sensory tubes in the cheek region, and lateral line scales with only a single pore posteriorly. Although the genus has been
regarded as comprising only a single species, Ratabulus diversidens (McCulloch, 1914), this study presents descriptions
of four: R. megacephalus (Tanaka, 1917) in southern Japan to the South China Sea, R. diversidens in eastern Australia, R.
fulviguttatus sp. nov. in northwestern Australia and R. ventralis sp. nov. in northeastern Australia. R. megacephalus, having
been regarded as a junior synonym of R. diversidens, is easily separable from that species in having more anteroventrally
slanted oblique scale rows above the lateral line (94—112 versus 80-93). R. fulviguttatus sp. nov. is similar to R.
megacephalus in having small dark spots dorsally on the body, but differs from it in having a shorter snout (30.4—34.8%
HL versus 31 .2-35.7% HL), longer pelvic fin (20.9-25.7% SL versus 19.5-23.1% SL), and a pale brown head and body
(versus dark brown) . Although R. ventralis sp. nov. resembles R. diversidens in having the nasal bone with tubercles, the
former is distinguished from the latter and R. megacephalus by its longer pectoral fin (15.8-18.6% SL versus 13 .9-17.0%
SL). R. diversidens also differs from its three congeners in having larger brownish spots on the pelvic fin.

Keywords Ratabulus, revision, Ratabulus fulviguttatus sp. nov., Ratabulus ventralis sp. nov.

Introduction

Jordan and Hubbs (1925) proposed the genus Ratabulus for
Thysanophrys megacephalus Tanaka, 1917 (the spelling
Rutabulus also appeared in this publication, as described below),
based on its possession of characters, such as canine-like upper
jaw teeth. Matsubara and Ochiai (1955) redefined the genus
using a greater variety of characters, including osteology, as seen
in the configuration of the urohyal and pelvic bones. Insidiator
diversidens, described by McCulloch in 1914, was subsequently
referred to this genus (e.g. Sainsbury et al., 1985; Paxton et al.,
1989; Hoese et al., 2006). Knapp (1999) synonymised R.
megacephalus with R. diversidens, but some authors did not
agree (e.g. Nakabo, 2002; Hoese et al., 2006). After examining
specimens collected from the West Pacific Ocean and Australia
in detail, we concluded that the genus comprises four species,
including the northwest Pacific R. megacephalus, two new
species from northwestern, and northeastern Australia, as well as
R. diversidens, confined to southeastern Australia. We provide
descriptions for all four and a key to distinguish between them.

Materials and methods

Counts and measurements were made according to Hubbs and
Lagler (1958), and were routinely taken from the left side,
except for gill rakers that were counted on the right side. A
small detached spine at the origin of the first dorsal fin and
another between the dorsal fins are expressed by separating
the values with a V, and were not included in the length of the
first dorsal fin base. Pectoral fin counts follow the formula:

dorsal unbranched
+

intermediate branched = total rays.

+

ventral unbranched rays

The number of oblique scale rows above the lateral line is
the number of diagonally angled scale rows slanting downward
and forward (anteroventrally), or downward and backward
(posteroventrally), which was counted just above the lateral
line. Measurements of less than 100 mm were made with

20

H.lmamura & M.F. Gomon

calipers to the nearest 0.1 mm; those 100 mm or more were
recorded to an accuracy of three significant figures.
Terminology of head spines follows Knapp et al. (2000).
Institutional acronyms are from Eschmeyer (1998), except for
Hokkaido University Museum, Hakodate (HUMZ) and
National Museum of Nature and Science, Tokyo (NSMT).
Standard and head lengths are abbreviated as SL and HL,
respectively. In species descriptions, meristic and
morphometric data for primary types are presented first,
followed by the range in secondary or nontype material
enclosed by parentheses where variations that deviate from the
primary type value were observed. Color comparisons
between species are based on preserved specimens. Collection
localities of the four species of Ratabulus are shown in fig. 1.

Genus Ratabulus

(Figures 2-8)

Ratabulus Jordan and Hubbs, 1925: 286 (original description,
type species: Thysanophrys megacephalus Tanaka, 1917).

Rutabulus Jordan and Hubbs, 1925: 93 (incorrect original spelling
and unavailable name; see ‘Remarks’).

Diagnosis. A genus of Platycephalidae with I + VII to IX + 0 or
1-10 to 12 dorsal fin rays (usually I + VIII + 1-12); 11 or 12 anal
fin rays (usually 12); 18-21 pectoral fin rays (usually 19 or 20);
52-56 pored lateral line scales; 1 + 5 to 8 gill rakers (usually 1
+ 6 or 7); body depressed and elongate, mostly covered with
ctenoid scales, some cycloid scales on undersurface; head
flattened; postorbital region, opercle, interorbit and nape
scaled; upper surface of eye without papillae or flaps; iris lappet

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usually broad and simple dorsally; interorbit moderately
narrow and slightly concave; posterior margin of orbit lacking
distinct pit; interopercular flap absent; moderate to long and
slender canines on upper jaw, palatine, and in two separate
patches on vomer; tooth band on upper jaw without distinct
notch; lip margins without papillae; suborbital and preopercular
sensory tubes absent on cheek; pored lateral line scales with a
single exterior opening posteriorly; first dorsal fin originating
slightly posterior to opercular margin; pectoral fin rounded
posteriorly, its posterodorsal corner weakly pointed; innermost
pelvic fin ray unbranched, others branched; fourth pelvic fin
ray longest; and posterior margin of caudal fin slightly oblique,
upper lobe longer.

Remarks. The genus Ratabulus includes the following four
species: R. diversidens from eastern Australia, R. megacephalus
from the Northwest Pacific, R. julviguttatus sp. nov. from
northwestern Australia and i?. ventralis sp . nov. from northeastern
Australia. Ratabulus can be easily distinguished from other
genera of Platycephalidae in usually having one small free spine
between the dorsal fins, iris lappet usually broad and simple
dorsally, suborbital ridge with many small to large spines,
moderate to long and slender canines on the upper jaw, palatine
and in two separate patches on the vomer, no sensory tubes from
the suborbitals and preopercle in the cheek region, and pored
lateral line scales with one posterior exterior opening.

Although Jordan and Hubbs (1925: 286) proposed the
name Ratabulus for this genus, Rutabulus appeared in the list
of new genera on an earlier page (96) of the same publication.
As no-one has dealt with this discrepancy, we, as first revisers,
consider Ratabulus to be the correct spelling, since it has been
used by most subsequent authors (e.g. Matsubara and Ochiai,
1955; Paxton et al., 1989; Shao and Chen, 1993; Imamura,
1996; Kim et al., 2005; Knapp, 1999; Hoese et al., 2006).
Rutabulus, therefore, is regarded as an incorrect spelling and
an unavailable name (ICZN, 1999: Arts. 24.2.3, 32.4).

Key species of Ratabulus

1 Anteroventrally slanted oblique scale rows above lateral

line 80-93; dorsal surface of head with large and pale to
dark-brown spots of irregular shapes; pelvic fin with large
brown spots R. diversidens

— Anteroventrally slanted oblique scale rows above lateral

line 91-113; dorsal surface of head with small, round,
brown spots; pelvic fin with small brown spots 2

2 Nasal bone with tubercles (fig. 4d); pelvic fin 19.5-25.7%

SL; dorsal surface of body without spots

R. ventralis sp.nov.

— Nasal bone without tubercles (figs 4b-c); pelvic fin 25.3-

28.2% SL; dorsal surface of body with brownish or dark-
brownish spots 3

3 Snout 30.4-34.8% HL; dorsal surface of head and body

pale brown R. Julviguttatus sp. nov.

Figure 1. Map of the West Pacific and Australia with collection
localities for specimens of four species of Ratabulus.

Snout 31.2-35.7% HL; dorsal surface of head and body
dark brown R. megacephalus

Revision of Ratabulus

21

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Figure 2. Lateral views of four species of Ratabulus : a, R. diversidens, AMS 1.40494-001, 391 mm SL; b, R. megacephalus, HUMZ 200048, 344 mm
SL; c, R.julviguttatus sp. nov., CSIRO H4031-79, holotype, 262 mm SL; d, R. ventralis sp. nov., CSIRO H61 16-02, holotype, 304 mm SL.

Ratabulus diversidens (McCulloch, 1914)

English name: Freespine flathead
(Figures 2a, 3a, 4a, 5a, 6a, 7, 8a)

Insidiator diversidens McCulloch, 1914: 148, fig. 13, pi. 31-fig. 1
(original description, type locality: 11 km northeast of Port Stephens
Lighthouse, New South Wales, Australia); McCulloch, 1929: 403 (list
and distribution. New South Wales, Australia)

Ratabulus diversidens : Paxton et al., 1989: 470 (list and
distribution, Queensland to off Sydney, New South Wales, Australia)
(in part); Knapp, 1999: 2410, unnumbered fig. (description, eastern
Australia) (in part); Hoese et al., 2006: 944 (list and distribution,
Queensland to off Sydney, New South Wales, Australia) (in part).

Lectotype (designated here). AMS E.2103, 232 mm SL, 11 km
northeast of Port Stephens Lighthouse, New South Wales, 87 m, 10
November 1910, FIS Endeavour.

Paralectotypes. Two specimens. AMS E.1566, 212 mm SL and
AMS 1.11254, 245 mm SL, collected with lectotype.

Nontypes. Twenty-three specimens (54.4-391 mm SL) eastern
Australia. AMS 1.15523-010, 190 mm SL, off Brisbane, Queensland
(26°31'S, 153°28'E), 137 m, 26 July 1968; AMS 1.23993-003, 205 mm
SL, Ballina-Tweed Heads, New South Wales (28°13'S, 153°52'E), 201
m, 17 August 1978, FRY Kapala ; AMS 1.25097-007, 2: 118-140 mm

SL, east of Brunswick Heads, New South Wales (28°24’S, 153°51'E),
155-174 m, 3 June 1978, FRV Kapala ; AMS 1.25804-019, 278 mm SL,
just north of Townsville, Queensland (17°51'S, 147°01'E), 260 m, 9
January 1986, RV Soela; AMS 1.31332-001, 370 mm SL, off
Wollongong, New South Wales (34°25'S, 152°00'E), 18-109 m,
February 1991; AMS 1.39088-001, 369 mm SL, 5-6 km offshore.
North Head, New South Wales (33°17’S, 151°35'E), 65 m, 31 January
1999; AMS 1.40494-001, 391 mm SL, off Crowdy Head, New South
Wales (31°51'S, 152°45'E), 60-70 m, 2000; AMS 1.45084-009, 2 of 5:
182-230 mm SL, northeast of Arrawarra Headland, New South Wales
(29°30'S, 153°48'E), 7 May 1971, FRV Kapala ; CSIRO H630-29, 273
mm SL, south of Saumarez Reef, Queensland (22°36’S, 153°50'E),
345-350 m depth, 17 November 1985, FRV Soela ; CSIRO H630-30,
308 mm SL, collaboration with CSIRO H630-29; CSIRO H698-22,
295 mm SL, east of Bowen, Marian Plateau, Queensland (19°29.2'S,
150°16.5'E - 19°29.8'S, 150°17.8'E), 324-328 m, 15 November 1985,
FRV Soela ; CSIRO H4268-01, 204 mm SL, east of Pambula, New
South Wales (36°54'S, 149°58'E - 36°55’S, 149°57'E), 42-43 m, 28
April 1996, FRV Southern Surveyor ; NMV A15248, 290 mm SL, off
Lakes Entrance, Victoria (38°07’S, 147°45'E), January 1995; NMV
A 15249, 283 mm SL, collaboration with NMV A15248; NMV A19471,
349 mm SL, off Lakes Entrance, Victoria (38°17'52 2 S, 148°33'34 2 E),
90-156 m, 22 October 1997; QM 1.2117, 313 mm SL, east of Tweed
Heads, New South Wales (28°12'S, 154°54'E), 235 m, 27 July 1982;

Revision of Ratabulus

23

Figure 3 . Dorsal views of four species of Ratabulus : a, R. diver sidens, AMS 1.40494-001, 391 mm SL; b, R. megacephalus, HUMZ 200048, 344 mm
SL; c, R.fulviguttatus sp. nov., CSIRO H4031-79, holotype, 262 mm SL; d, R. ventralis sp. nov., CSIRO H61 16-02, holotype, 304 mm SL.

QM 1.18698, 215 mm SL, Queensland (25°27'S, 153°46'E - 25°17’S,
153°43'E), 183-230 m, 14 September 1980; QM 1.18813, 2: 203-252
mm SL, Queensland (23°50'S, 152°36'E - 23°46'S, 152°32’E), 238-274
m, 23 September 1980; QM 1.26624, 274 mm SL, east of Fraser Island,
Queensland (26°S, 153.3°E), 30 m, 16 May 1990; QM 1.34240, 2:
54.4-144 mm SL, southeast of Cape Moreton, Queensland (28°12’S,
154°54'E), 235 m, 27 July 1982.

Diagnosis. A species of Ratabulus with 80-93 anteroventrally
slanted oblique scale rows above lateral line; snout length 29.5-
32.3% HL, slightly decreasing proportionally with growth;
pectoral fin length 13.9-15.8% SL; pelvic fin length 22.1-28.5%
SL; nasal bone with tubercles in larger specimens; dorsal surface
of head with large, pale to dark-brown irregularly shaped spots,
body without spots dorsally; pelvic fin with large brown spots.

24

H.lmamura & M.F. Gomon

Figure 4. Dorsal views of anterior head region in four species of Ratabulus: a, R. diversidens , AMS 1.11523-010, 190 mm SL; b, R. megacephalus,
BSKU 87405, 216 mm SL; c, R. fulviguttatus sp. nov., CSIRO H4031-79, holotype, 262 mm SL; d, R. ventralis sp. nov., CSIRO H6116-02,
holotype, 304 mm SL. Arrows show tubercles on nasal bone. Scale bar = 10 mm.

Figure 5. Iris lappet (left eye) of four species of Ratabulus: a, R. diversidens , AMS 1.45084-009, 230 mm SL; b, R. megacephalus, BSKU 87405,
216 mm SL; c, R. fulviguttatus sp. nov., CSIRO H4031-79, holotype, 262 mm SL; d, R. ventralis sp. nov., CSIRO H6116-02, holotype, 304 mm
SL. Scale bar = 3 mm.

Revision of Ratabulus

25

Figure 6. Dorsolateral view of pelvic fin in four species of Ratabulus : a, R. diversidens, CSIRO H630-29, 273 mm SL; b, R. megacephalus,
HUMZ 200048, 344 mm SL; d, R. fulviguttatus sp. nov., CSIRO H4031-79, holotype, 262 mm SL; d, R. ventralis sp. nov., CSIRO H61 16-02,
holotype, 304 mm SL.

Figure 7. Dorsal (upper) and lateral (lower) views of Ratabulus diversidens, AMS E.2103, lectotype, 232 mm SL, 11 km northeast of Port
Stephens Lighthouse, New South Wales.

Description. Dorsal fin rays I+VIII+I-11 (I+VIII+I-10 or 11, 10 in
one ); anal fin rays 12; branched caudal fin rays 12 (11-13, usually
12 or 13, smallest specimen with 11 rays); pectoral fin rays 2+10
+ 8 - 20 (2 or 3 + 10 - 12 + 6 to 8 - 19 or 20); pelvic fin rays 1, 5;
scales in lateral line 54 (53 or 54), anterior 6 (3-7) scales with
spine; posteroventrally slanted oblique scale rows above lateral
line 75 (71-78); anteroventrally slanted oblique scale rows above
lateral line 84 (80-93); gill rakers 1 + 6 - 7 (1 + 5-8 - 6-9).

See table 1 for selected proportional measurements. Head
length 2.6 (2.4— 2.7) in SL. Snout rather robust, its length 3.2 in
HL (3. 1-3 .3 in HL, ratio as % HL slightly decreasing
proportionally with growth) (fig. 9). Iris lappet broad and simple
dorsally, absent ventrally (rarely absent dorsally, and usually
broad and simple ventrally) (fig. 5a). Interorbital width 14.1
(10.3-22.2) in HL. Nasal bone without distinct spines, but with
tubercles (smallest specimen, 54.4 mm SL, with single spine

26

H.lmamura & M.F. Gomon

Figure 8. Three species of Ratabulus : a, R. diversidens, off Newcastle, New South Wales (33°08'S, 151°56'E), 125-130 m, 7 September 1978,
specimen discarded, scale bar = 5 cm, photo by K. Graham; b, R. megacephalus, HUMZ 199844, 264 mm SL, East China Sea (26°35.68'N,
125°04.57'E - 26°36.73'N, 125°05.78'E), 192-186 m, 5 May 2007, photo by Hokkaido University; c, R. fulviguttatus sp. nov., upper specimen
CSIRO H1505-15, 189 mm SL, lower specimen H1505-06, 219 mm SL, north of Nickol Bay, Western Australia (19°07'S, 117°06'E - 19°07'S,
117°04'E), 177-184 m depth, 5 October 1988, FRY Soela , photo by CSIRO.

Revision of Ratabulus

27

and without tubercles) (fig. 4a). Lachrymal with four (2-7)
anterolaterally directed spines. Single (rarely two) preocular
spine in front of eye, its base with tubercles (with small spines,
or without small spines and tubercles in several specimens).
Suborbital ridge roughly serrated by many small to large spines;
anteriormost (preorbital) spine small (rarely absent) . Supraorbital
ridge serrated, except anteriorly. Single postocular spine present.
Pterotic with serrated ridge ending in strong spine. Parietal with
single spine, followed posteriorly by three small spines on left
side and four on right (by 1-5 spines). Supratemporal with
serrated ridge (with serrated or smooth ridge) ending in spine.
Posttemporal with two spines on left side and one on right (with
1—4 spines, usually one). Preopercle with three (two or three)
spines; uppermost longest, not reaching posterior margin of
opercle, bearing two small spines on left side and one on right on
base laterally (usually one). Ridge of lower opercular spine
smooth on left side and serrated on right. Posterior end of
maxilla below anterior margin of eye (just beyond it in several
specimens) . Anterior part of upper jaw with conical teeth (short
canines in several specimens) anteriorly, followed by long,
slender canines; middle and posterior parts of upper jaw with
villiform teeth, but innermost row with small, slender conical
teeth. Lower jaw with narrow tooth band containing small
conical teeth anteriorly, becoming smaller posteriorly, followed
by villiform teeth at end of jaw; innermost row conical (conical
teeth to moderately long canines). Palatine with moderately
broad tooth band; anterior part of palatine with short canines
laterally and moderately long canines mesially (moderate to
long and slender canines mesially); posterior part of palatine
with small conical teeth. Vomer with about four (about three or
four) tooth rows centrally; anterior part of vomer with small
conical teeth, middle and posterior parts with moderate to long
and slender canines. Posterior margin of caudal fin mostly
straight (rarely slightly concave); caudal fin length 5.6 (4 .8-6.1)
in SL. Pectoral fin length 7.0 (63-7.2) in SL. Posterior tip of
pelvic fin reaching second (first to third) anal fin ray; pelvic fin
length 4.0 (3 .5-4.5) in SL.

Color in alcohol (lectotype, fig. 7). Head and body mostly
faded. First dorsal fin with dark brown submarginal stripe;
second dorsal with scattered small dark brown spots. Caudal fin
with five longitudinal black stripes along fin membranes
posteriorly. Upper part of pectoral fin with about two dark brown
bands. Pelvic fin with large dark-brown spots posteriorly.

Other specimens with head and body pale to dark-brown
dorsally, pale yellow ventrally; dorsal surface of head with large
irregular pale to dark-brown spots; body without spots, but
sometimes with several narrow dark-brown bands dorsally, side
with longitudinal stripe formed by continuous gray spots below
lateral line. First dorsal fin with dark brown to black submarginal
stripe, base clear anteriorly, with scattered small dark brown to
black spots; second dorsal with dark brown spots. Anal fin pale
or with melanophores along rays. Caudal fin with 4—6 narrow
black longitudinal stripes posteriorly; lower stripes tending to
merge in some specimens; upper part of fin with several brown
spots. Pectoral fin with small dark-brown spots tending to form
bands. Pelvic fin with large brown spots (fig. 6a).

Color when fresh based on photograph (fig. 8a). Similar to
those in alcohol.

Standard length (mm)

Figure 9. Comparison of snout length (% HL) with standard length
(mm) in specimens examined of four species of Ratabulus-, solid
square, R. diversidens; open circle, R. megacephalus; open square, R.
fulviguttatus sp. nov.; solid circle, R. ventralis sp. nov.; 1, lectotype of
R. diversidens-, 2, two paralectotypes of R. diversidens-, 3, holotype of
R. fulviguttatus-, 4, holotype of R. ventralis.

Distribution. Eastern Australia from Townsville, Queensland
(17°51'S) to Lakes Entrance, Victoria (38°17'52 2 S), recorded at
depths of at least 30-345 m (McCulloch, 1914; Paxton et al:^
1989) (fig. 1).

Remarks. Ratabulus diversidens is easily separable from R.
megacephalus and R. fulviguttatus in having 80-93
anteroventrally slanted oblique scale rows above the lateral
line (versus 94-112 in R. megacephalus and 99-113 in R.
fulviguttatus ), the snout length 29.5-32.3% HL, becoming
slightly shorter proportionally with growth (versus 31.2-35.7%
HL in R. megacephalus and 30.4-34.8% HL in R. fulviguttatus)
(fig. 9), nasal bone with tubercles in larger specimens (versus
lacking tubercles in R. megacephalus and R. fulviguttatus )
(fig. 4), the dorsal surface of the head with large, irregularly
shaped, pale to dark-brown spots, the body without spots
dorsally (versus dorsal surface of head and body with small,
round, pale or dark-brown spots in R. megacephalus and R.
fulviguttatus) (fig. 3). R. diversidens resembles R. ventralis in
having tubercles on the nasal bone, but differs from it in having
a shorter pectoral fin (13.9-15.8% SL in R. diversidens versus
15.8-18.6% SL in R. ventralis ) (fig. 10) and large, irregular
spots on the head (versus a head with small, round, brown
spots in R. ventralis ) (fig. 3). This species can be separated
from the other three by the large brown spots on its pelvic fin
(versus having small spots) (fig. 6).

28

H.lmamura & M.F. Gomon

Standard length (mm)

Figure 10. Comparison of pectoral fin length (% SL) with standard
length (mm) in specimens examined of four species of Ratabulus;
solid square, R. diversidens; open circle, R. megacephalus; open
square, R. fulviguttatus sp. nov.; solid circle, R. ventralis sp. nov.; 1,
lectotype of R. diversidens-, 2, two paralectotypes of R. diversidens-, 3,
holotype of R. fulviguttatus-, 4, holotype of R. ventralis.

McCulloch (1914) recorded the collection locality of the
type specimens of R. diversidens as ‘seven miles S. 21°W. off
Port Stephens Lighthouse, New South Wales’, but data with the
specimens indicate they were captured 11 km northeast of Port
Stephens Lighthouse (S. Reader, pers. comm., 16 May 2006).

Ratabulus megacephalus (Tanaka, 1917)

Japanese name: Haname-gochi
(Figures 2b, 3b, 4b, 5b, 6b, 8b)

Thysanophrys megacephalus Tanaka, 1917: 11 (original

description, fish market in Tokyo); Tanaka, 1931: 37 (list, Japan);
Kamohara, 1952: 70 (list, Kochi, Japan).

Thysanophrys {Ratabulus) megacephalus-. Kamohara, 1964: 77
(list, Kochi).

Ratabulus megacephalus : Jordan and Hubbs, 1925: 287 (English
translation of original description published in Japanese); Okada and
Matsubara, 1938: 334 (key and distribution, Tokyo to Kagoshima,
Japan); Mori, 1952: 159 (list and distribution, Tongyeong, southern
South Korea); Matsubara and Ochiai, 1955: 95, pi. 3 (description,
southern Japan and East China Sea); Matsubara, 1955: 1122 (key, short
description and distribution, Tokyo Bay, south from Kasumi, Hyogo
Prefecture, Korea and East China Sea); Anonymous, 1962: 926, fig.
730 (description, Hainan Island, South China Sea); Ochiai, 1984: 322,
pi. 289-E (short description and distribution, southern Japan to East
China Sea); Yatou, 1985: 599, 729, pi. 371 (description and distribution,
Okinawa Trough); Shao and Chen, 1987: 86, fig. 20 (short description,
Taiwan); Shao and Chen, 1993: 258, pi. 65-4 (short description.

Taiwan); Imamura, 1996: 207, fig. 62 (list); Imamura, 1997: 220, fig. 5
in 221 page (short description); Lee and Joo, 1998: 220, fig. 4
(description, Pusan); Knapp, 2000: 608 (list. South China Sea);
Nakabo, 2002: 618, unnumbered figs (pictorial key and limited
meristic values. Pacific coast of southern Japan and East China Sea);
Shinohara et al., 2001: 318 (list, Tosa Bay, Japan); Youn, 2002: 259,
570, unnumbered fig. (pictorial key, Tongyeong, southern South
Korea); Kim et al., 2005: 237, unnumbered fig. (short description,
Tongyeong, southern South Korea).

Ratabulus diversidens (nee McCulloch, 1914): Knapp, 1999: 2410
(description. East and South China seas and northern Philippines) (in
part).

Nontypes (location of holotype unknown, Eschmeyer et al., 1998).
Twenty-three specimens (83.4-344 mm SL), Northwest Pacific. BSKU
9503, 186 mm SL, fish market, Mimase, Kochi Prefecture, 1 December
1950; BSKU 36141, 192 mm SL, fish market, Mimase, Kochi
Prefecture, 9 December 1981; BSKU 36270, 228 mm SL, fish market,
Mimase, Kochi Prefecture, 26 January 1982; BSKU 51453, 205 mm
SL, fish market, Mimase, Kochi Prefecture, 9 April 2000; BSKU
52257, 160 mm SL, Irino fishing port, Hata-gun, Kochi Prefecture, 10
August 2000; BSKU 54460, 252 mm SL, fish market, Mimase, Kochi
Prefecture, 24 November 2000; BSKU 59497, 83.4 mm SL, Irino
fishing port, Hata-gun, Kochi Prefecture, 18 March 2002; BSKU
63578, 188 mm SL, Saga fishing port, Hata-gun, Kochi Prefecture, 20
November 2002; BSKU 73629, 88.7 mm SL, Irino fishing port, Hata-
gun, Kochi Prefecture, 5 April 2002; BSKU 85012, 164 mm SL, Tosa
Bay, Kochi Prefecture, 125 m depth, 30 September 1997, R/V Kotaka-
maru; BSKU 87405, 216 mm SL, fish market, Mimase, Kochi
Prefecture, 30 March 2000; FAKU 12168, 270 mm SL, East China
Sea, February 1949, K. Matsubara and R. Ishiyama; FAKU 14957,
14959-14960, 3: 283-295 mm SL, Maisaka, Shizuoka Prefecture;
HUMZ 37397, 111 mm SL, no collection data (perhaps Kochi
Prefecture); HUMZ 49396, 187 mm SL, fish market, Mimase, Kochi
Prefecture, 15 November 1975; HUMZ 49470, 187 mm SL, fish
market, Mimase, Kochi Prefecture, 17 November 1975; HUMZ
200048, 344 mm SL, East China Sea (27°06.31’N, 125°47.32’E -
27°06.11’N, 125°46.24’E), 192-186 m, 2 June 2007; HUMZ 200050,
296 mm SL, collaboration with HUMZ 200048; NSMT-P 828, 103
mm SL, Enoura, Izu Peninsula, Shizuoka Prefecture (35°03'N,
138°54'E); USNM 329510, 239 mm SL, South China Sea (19°06'30 2 N,
112°23'E), 203-218 m, 22 July 1958; USNM 383571, 370 mm SL, fish
market, Bolinao, Luzon, Philippines, 7-9 October 1995.

Other material. One specimen. HUMZ 199844, 264 mm SL, East
China Sea (26°35.68'N, 125°04.57'E - 26°36.73’N, 125°05.78'E), 192-
186 m, 5 May 2007 (used for description of color when fresh).

Diagnosis. A species of Ratabulus with 94-112 anteroventrally
slanted oblique scale rows above lateral line; snout length 31.2-
35.7% HL, markedly decreasing in length proportionally with
growth; pectoral fin length 13.9-17.0% SL; pelvic fin length
19.5-23.1% SL; nasal bone without tubercles; dorsal surface of
head and body dark brown, with small, round, dark-brown
spots; and pelvic fin with small brown to black spots.

Description. Dorsal fin rays I + VIII + 1-11 or 12 or I + IX + 0-11
(I + IX + 0-11 in one, I + VIII + 1-12 in one); anal fin rays 12;
pectoral fin rays two or three (usually two) + 9-11 + 6-8 = 19
or 20; pelvic fin rays I, 5; branched caudal fin rays 12 or 13;
scales in lateral line 53-55, anterior three or four scales with
spine; posteroventrally slanted oblique scale rows above lateral
line 71-78; anteroventrally slanted oblique scale rows above
lateral line 94-112; gill rakers 1 + 6-8 = 7-9.

Revision of Ratabulus

29

See table 1 for selected proportional measurements. Head
length 2.4— 2.7 in SL. Snout rather long, its length 2.8-3 .2 in
HL, ratio as %HL markedly decreasing proportionally with
growth (fig. 9). Iris lappet broad and simple dorsally and absent
ventrally (fig. 5b). Interorbital width 12.8-19.6 in HL. Nasal
bone with 0-2 spines, and without tubercles (fig . 4b) . Lachrymal
with 2^4 spines directed anterolaterally. Preocular spine in
front of eye, its base without spines or tubercles. Suborbital
ridge with many small to large spines; anteriormost (preorbital)
spine usually present. Supraorbital ridge serrated medially and
posteriorly or just posteriorly. Postocular spine present. Pterotic
ridge with 1-3 spines. Parietal with single spine, followed
posteriorly by 0-2 additional spines. Supratemporal usually
with a spine, rarely with two. Posttemporal with one or two
spines. Preopercle with twp or three spines; uppermost longest,
not reaching posterior margin of opercle, bearing one small
spine on base laterally. Ridge of lower opercular spine without
serrations. Posterior end of maxilla below anterior margin of
eye or just posterior to it. Anterior part of upper jaw with
conical or small canine teeth in front of long, slender canines,
followed by villiform teeth, innermost row with small, slender
conical teeth. Lower jaw with narrow band of small conical
teeth anteriorly, smallest teeth posteriorly, followed by villiform
teeth, innermost a row of moderately long canines. Palatine
with moderately broad tooth band; those anterolaterally canines
of short to moderate length with moderate to long and slender
canines mesially; posterior part of palatine with small conical
teeth. Vomer with about 2-5 tooth rows medially; anterior teeth
conical or short canines, those medially and posteriorly slender
canines of moderate length. Posterior margin of caudal fin
mostly straight, slightly concave or slightly rounded in some;
caudal fin length 5.2-6 .0. Pectoral fin length 5.9-1 2 in SL.
Posterior tip of pelvic fin not reaching anal fin origin; pelvic fin
length 4. 3-5.1 in SL.

Color in alcohol. Head and body dark brown dorsally, pale
yellowish ventrally. Dorsal surfaces of head and body with
small, round, dark-brown spots; body usually without bands
dorsally, but occasionally with indistinct darker bands;
longitudinal stripe formed by continuous series of gray spots
on side below lateral line. First dorsal fin with dark brown to
black submarginal stripe, base clear anteriorly with scattered
small dark-brown to black spots; second dorsal with brown to
black spots. Anal fin with melanophores along rays. Caudal fin
with 5-9 narrow longitudinal black stripes posteriorly; several
dark-brown to black spots anteriorly and dorsally. Pectoral and
pelvic fins with small dark-brown to black spots (fig. 6b).

Color when fresh from photographs of HUMZ 199844 (fig.
8b). Similar to those in alcohol.

Distribution. East and South China seas, and Northwest Pacific,
including southern Japan, Korea, Taiwan, Hainan Island and
northern Philippines, at depths of 192-218 m (Anonymous,
1962; Ochiai, 1984; Shao and Chen, 1987, 1993; Knapp, 1999;
Kim et al„ 2005) (fig. 1).

Remarks. R. megacephalus is most similar to R. fulviguttatus in
having the nasal bone devoid of tubercles (versus tubercles
present in R. diversidens and R. ventralis ) (fig. 4) and the dorsal
surface of the body with small, round spots (versus without

Standard length (mm)

Figure 11. Comparison of pelvic fin length (% SL) with standard
length (mm) in specimens examined of four species of Ratabulus;
solid square, R. diversidens; open circle, R. megacephalus; open
square, R. fulviguttatus sp. nov.; solid circle, R. ventralis sp. nov.; 1,
lectotype of R. diversidens; 2, two paralectotypes of R. diversidens; 3,
holotype of R. fulviguttatus; 4, holotype of R. ventralis.

spots) (fig. 2). R. megacephalus differs from R. fulviguttatus in
having a proportionally longer snout (31.2-35.7% HL in R.
megacephalus versus 30.4—34.8% HL in R. fulviguttatus) and
shorter pectoral fins (13.9-17.0% SL compared to 14.6-16.5%
SL) (figs 9-10) at comparable sizes, and the dorsal surface of the
head and body dark brown (versus pale brown) (fig. 2).
R. megacephalus is also separable from R. ventralis in having
shorter pectoral fins (13.9-17.0% SL versus 15.8-18.6% SL) and
pelvic fins (19.5-23.1% SL versus 25.3-28.2% SL) (figs 10-11).

Sadovy and Cornish (2000) reported Ratabulus
megacephalus from Hong Kong, but their photograph depicts
a specimen of Inegocia ochiaii Imamura, 2010, previously
known incorrectly as ‘ Inegocia guttata ’ (e.g. Matsubara and
Ochiai, 1955; Ochiai, 1984), a species that has an interopercular
flap, and a long and branched iris lappet.

Ratabulus fulviguttatus sp. nov.

English name: Orangefreckled flathead
(Figures 2c, 3c, 4c, 5c, 6c, 8c)

Ratabulus diversidens (nee McCulloch, 1914): Gloerfelt-Tarp and
Kailola, 1984: 123, unnumbered fig. (short description, northwestern
Australia); Sainsbury et al., 1985: 114, unnumbered fig. (description,
northwestern Australia); Paxton et al., 1989: 470 (list and distribution,
Northwestern Shelf, Australia) (in part); Knapp, 1999: 2410 (description,
Northwestern Shelf and Timor Sea) (in part); Hutchins, 2001: 28 (list.
Western Australia); Hoese et al., 2006: 944 (list and distribution, off
North West Cape to off Port Hedland, Western Australia) (in part).

30

H.lmamura & M.F. Gomon

Holotype. CSIRO H4031-79, 262 mm SL, north of Cape Lambert,
Western Australia (18°57’S, 117°14’E), 248 m, 30 August 1995, FRV
Southern Surveyor.

Paratypes. 16 specimens (140-266 mm SL), from northwestern
Australia. AMS 1.21621-006, 206 mm SL, northwest shelf. Western
Australia (11°49'S, 124°17'E), 195-200 m, 10 June 1979; AMS
1.22805-013, 3: 177-188 mm SL, 170 km north of Port Hedland,
Western Australia (18°28'S, 118°15'E), 150-156 m, 28 March 1982,
FRV Soela ; AMS 1.22807-023, 3: 188-227 mm SL, 175 km north of
Port Hedland, Western Australia (18°32'S, 118°17'E), 200-204 m, 2
April 1982, FRV Soela ; AMS 1.22828-012, 3: 248-265 mm SL, 190
km north of Port Hedland, Western Australia (19°01'S, 117°12'E),
200-202 m, 14 April 1982, FRV Soela ; CSIRO CA3624 (voucher of
Sainsbury et al., 1985), 233 mm SL, northwest of Nichol Bay, Western
Australia (19°15'S, 116°40'E), 172 m, 25 January 1983, FRV Soela ;
CSIRO CA4091, 229 mm SL, north of Bathurst Island, Arafura Sea,
Northern Territory (10°02'S, 130 o 01'E), 216 m, 8 July 1980; CSIRO
H1035-23, 251 mm SL, north of Dampier Archipelago, Western
Australia (19°08'S, 116°54'E), 196 m, 24 October 1996, FRV Soela ;
CSIRO H1512-04, 140 mm SL, north of Monte Bello Islands, Western
Australia (19°39’S, 115°36'E), 180 m, 11 October 1988, FRV Soela-,
CSIRO H4031-79, 262 mm SL, north of Cape Lambert, Western
Australia (18°57'S, 117°14'E), 248 m, 30 August 1995, FRV Southern
Surveyor, CSIRO H4631-03, 198 mm SL, north of Dampier
Archipelago, Western Australia (19°11'S, 116°35'E), 196 m, 11 August
1997, FRV Southern Surveyor, WAM P.32204-001, 208 mm SL,
Western Australia, Timor Sea (12°57'S, 124°20'E), 8 June 1979.

Nontypes. Two specimens. WAM P.9351-001, 76.0 mm SL,
Western Australia (21°49’S, 113°56'E), 121-126 m, 1 February 1964;
WAM P. 9352-001, 77.1 mm SL, coll, with WAM P.9351-001.

Other material. Two specimens. CSIRO HI 505- 06, 219 mm SL,
north of Nickol Bay, Western Australia (19°07'S, 117°06'E - 19°07'S,
117°04'E), 177-184 m, 5 October 1988, FRV Soela-, CSIRO H1505-06,
189 mm SL, collected with CSIRO H1505-06 (used for description of
color when fresh).

Diagnosis. A species of Ratabulus with 99-113 anteroventrally
slanted oblique scale rows above lateral line; snout length
30.4-34.8% HL, markedly decreasing proportionally with
growth; pectoral fin length 14.6-16.5% SL; pelvic fin length
20.9-25.7% SL; nasal bone without tubercles; dorsal surface of
head and body pale brown, with small, round brown spots; and
pelvic fin with small brown spots.

Description. Dorsal fin rays I + VIII + 1-11 (I + VII to IX + 1-11
or 12, VII in one and IX in one, 12 in one); anal fin rays 12 (11
or 12, 11 in one); pectoral fin rays 2 + 12 + 6 = 20 (1 or 2 + 9-12
+ 6-8 = 19-21, usually 20); pelvic fin rays I, 5; branched caudal
fin rays 13; scales in lateral line 56 (54-56), anterior three
(three or four) scales with spine; posteroventrally slanted
oblique scale rows above lateral line 80 (71-80); anteroventrally
slanted oblique scale rows above lateral line 107 (99-113); gill
rakers 1 + 7 = 8 (1 + 6-8 = 7-9, usually 7).

See table 1 for selected proportional measurements. Head
length 2.6 (2.4— 2.7) in SL. Snout rather slender, its length 3.3
(3 .0-3 .3) in HL, markedly decreasing proportionally with growth
(fig. 9). Iris lappet broad and simple both dorsally and ventrally
(fig. 5c). Interorbital width 16.3 (14.5-20.0) in HL. Nasal bone
with small spine, but without tubercles (fig. 4C). Two spines on
lachrymal of right side, three on left, spines directed
anterolaterally. Single preocular spine in front of eye, its base
without spines or tubercles (with tubercle on left side of one

paratype, CSIRO H4631-03). Suborbital ridge with many small
to large spines, anteriormost (preorbital) distinct. Supraorbital
ridge serrated except anteriorly. Single postocular spine present.
Pterotic with one spine (one to four spines). Parietal with single
spine, without accompanied spines posteriorly (followed by one
or two spines in several paratypes). Supratemporal with one
spine (two to three spines in several paratypes). Posttemporal
with one spine. Preopercle with two spines (three in several
paratypes); upper longer, not reaching posterior margin of
opercle, bearing one small spine on base laterally (rarely two
spines or spines absent) . Ridge of lower opercular spine without
serrations. Posterior end of maxilla below anterior margin of
pupil (not reaching to anterior margin of pupil in several
paratypes). Anterior part of upper jaw with short canines at front,
followed by long and slender canines posteriorly; middle and
posterior parts of upper jaw with villiform teeth, innermost row
comprising small, slender, conical teeth. Lower jaw with two
tooth rows (one to about three rows); outer row with moderately
long conical teeth anteriorly, and two rows of villiform teeth
anterolaterally (outer one to two rows of villiform teeth, or small
to moderately long conical teeth), teeth becoming smaller
posteriorly, teeth villiform posteriorly; inner row with conical
teeth (canines of small to moderate size in several paratypes).
Palatine with moderately broad tooth band; anterior part of
palatine with canines of moderate length (long canines in several
paratypes) laterally, and long and slender canines mesially;
posterior part of palatine with small to moderately long conical
teeth. Vomer with about three tooth rows medially (with two to
four rows), small canines anteriorly (canines of moderate length
in several paratypes), followed by long, slender canines. Posterior
margin of caudal fin slightly concave (mostly straight in several
paratypes); fin length 6.0 (5. 0-6 .2) in SL. Pectoral fin length 6.1
(6. 1-6 .8) in SL. Posterior tip of pelvic fin not reaching anal fin
origin (just reaching anal fin origin or just beyond it in several
paratypes); pelvic fin length 4.3 (3 .9^4.5) in SL.

Color in alcohol. Head and body pale brown dorsally, pale
yellowish ventrally; dorsal surface of head and body with
small, round, brown spots, but no bands; lateral side of body
below lateral line with pale purple longitudinal stripe (or stripe
formed by continuous series of purple spots). First dorsal fin
with one dark grayish submarginal stripe, basal area clear
anteriorly with scattered small dark-brown spots; second
dorsal fin with brown (or dark brown) spots. Anal fin pale.
Caudal fin with many black longitudinal narrow stripes and
spots posteriorly; upper part of fin with several brown spots.
Pectoral fin slightly dusky, with small brown spots. Pelvic fin
mostly faded on left side, with few small indistinct brown
spots on right (with several small distinct brown spots in
several paratypes) (fig. 6c).

Color when fresh from photographs of CSIRO H505-15
and H505-16 (fig. 8c). Head and body with small, round,
reddish brown and brown spots dorsally. Body with two broad
brown bands dorsally. Other colors similar to those in alcohol.

Distribution. Northwestern Australia, from North West Cape,
Western Australia (21°49'S) to Bathurst Island, Northern
Territory (10°02'S), at depths of at least 126-248 m (Sainsbury
et al., 1985; Paxton et al., 1989) (fig. 1).

Revision of Ratabulus

31

Etymology. The specific name julviguttatus, from the Latin
word meaning ‘orange or brown spots’, refers to this species’
characteristic spots on the head and body.

Remarks. R. julviguttatus differs from R. ventralis in having
the nasal bone without tubercles (versus with tubercles) (fig. 4),
shorter pelvic fins (20.9-25.7% SL versus 25.3-28.2% SL) (fig.
11), and body with small, round, brown spots (versus without
spots) (fig. 2). It is easily separable from R. diversidens, with
which it has been confused, by the small, round, brown spots
scattered over the dorsal surface of the head and body (versus
without spots). The spots in R. julviguttatus are reddish and/or
brown when fresh, which explains the Australian standard
name. The character is clearly evident in colour photos and
description provided by Gloerfelt-Tarp and Kailola (1984) and
Sainsbury et al. (1985).

Ratabulus ventralis sp. nov.

New English name: Longfin flathead
(Figures 2d, 3d, 4d, 5d, 6d)

Ratabulus diversidens (nec McCulloch, 1914): Paxton et al., 1989:
470 (list and distribution, off Brisbane) (in part); Knapp, 1999: 2410
(description. Coral Sea) (in part); Hoese et al., 2006: 944 (list and
distribution, off Brisbane) (in part).

Holotype. CSIRO H61 16-02, 304 mm SL, east of Townsville,
Queensland (18°39.3'S, 148°03.4'E - 18°36.4'S, 147°59.5'E), 244-248
m, 8 December 1985, FRV Soela.

Paratypes. Eleven specimens (172-328 mm SL) from Queensland,
northeastern Australia. AMS 1.25804-019, 278 mm SL, just north of
Townsville (17°51'S, 147°01'E), 260 m, 9 January 1986, FRV Soela-
AMS 1.25823-002, 328 mm SL, north of Townsville (17°58'S,
147°02'E), 260 m, 16 January 1986, FRV Soela] AMS 1.25832-007-
006, 284 mm SL, north of Townsville (17°58'S, 147°03'E), 260 m, 19
January 1986, FRV Soela ; CSIRO H690-03, 232 mm SL, Swain Reefs
(21°31'S, 152°58'E), 247 m, 20 November 1985, FRV Soela ; QM

l. 18546, 284 mm SL, off Swain Reefs (22.54°S, 152.12°E - 22.59°S,
152.12°E), 347-384 m, 3 October 1980; QM 1.19276, 255 mm SL, east
of Capricorn Group (23.1 1°S, 153.00°E- 23.01°S, 1 52 ,55°E), 366-392

m, 20 September 1980; QM 1.20934, 254 mm SL, east of Swain Reefs
(22.03°S, 153.05°E), 170 m, 28 August 1983; QM 1.20939, 316 mm SL,
east of Bunker Group (23.59°S, 152.51°E), 340 m, 27 August 1983;
QM 1.21624, 221 mm SL, southeast of Swain Reefs (22.4°S, 153.35°E),
310 m, 6 September 1983; QM 1.23088, 260 mm SL, off Swain Reefs
(20.49°S, 151.52°E), 288 m, 20 September 1986; QM 1.34327, 172 mm
SL, east of Noosa (26.25°S, 153.4°E), 119-120 m, 19 July 2002.

Diagnosis. A species of Ratabulus with 91-104 anteroventrally
slanted oblique scale rows above lateral line; snout length 30.9-
32.4% HL, markedly decreasing proportionally with growth;
pectoral fin length 15.8-18.6% SL; pelvic fin length 26.2-
28.2% SL; nasal bone with tubercles; dorsal surface of head
with small, round, brown spots, body without dark spots; pelvic
fin with small brown spots.

Description . Dorsal fin rays I + VIII + 1- 1 1 (I + VII or VIII + 1-11,
or I + IX + 0-11, VII in one, IX in one); anal fin rays 12; pectoral
fin rays 2 + 11 + 7 = 20 (2 + 10 or 11 + 6-9 = 19-21); pelvic fin
rays I, 5; branched caudal fin rays 13 (12 or 13); scales in lateral
line 55 (52-55), anterior four (3-5) scales with spine;

posteroventrally slanted oblique scale rows above lateral line 83
(71-76); anteroventrally slanted oblique scale rows above lateral
line 98 (91-104); gill rakers l+7 = 8(l+7or8 = 8or9).

See table 1 for selected proportional measurements. Head
length 2.7 (2 .4-2 .6) in SL. Snout rather robust, its length 3.2 in
HL (3. 1-3 .2 in HL, slightly decreasing proportionally with
growth, fig. 9). Iris lappet broad and simple both dorsally and
ventrally (fig. 5d). Interorbital width 13.9 (12.5-16.7) in HL.
Nasal bone without distinct spines, but with tubercles (fig. 4d).
Lachrymal with three anterolaterally directed spines on left
side, right side partly damaged (2-5, with short serrations
posterior to spines in some paratypes). Single preocular spine
in front of eye, its base with tubercles (with small spines in
several paratypes). Suborbital ridge roughly serrated by many
small to large spines; anteriormost (preorbital) spine small.
Supraorbital ridge serrated except anteriorly. Single postocular
spine present. Pterotic with serrated ridge ending in strong
spine. Parietal with single spine, lacking spines posteriorly
(with small spines or tubercles in many paratypes).
Supratemporal with smooth ridge (with serrated ridge in many
paratypes) ending in spine. Posttemporal with one spine
(usually with 1-3, rarely with serrated ridge ending in one
spine). Preopercle with two (two or three) spines; upper longer,
not reaching posterior margin of opercle, with one small spine
on base laterally. Ridge of lower opercular spine without
serrations (with weak serrations or with spine in some
paratypes). Posterior end of maxilla reaching just beyond
anterior margin of eye. Front of upper jaw with short canines
(with conical teeth in some paratypes) anteriorly, followed by
long and slender canines; middle and posterior parts of jaw
with villiform teeth, one or two inner rows (innermost row)
having small, slender conical teeth. Lower jaw with narrow
tooth band of small conical teeth anteriorly, teeth smaller
posteriorly, followed by villiform teeth; innermost row with
short to moderately long (short to long and slender) canines.
Palatine with moderately broad tooth band; anterior part with
moderately long (short to moderately long) canines laterally,
and long, slender (moderately long to long and slender) canines
mesially; posterior part with moderately long to long conical
teeth. Vomer with about three (about 3-5) tooth rows medially;
short canines anteriorly, followed by long, slender canines.
Posterior margin of caudal fin slightly concave (mostly straight
in some paratypes); caudal fin length 6.0 (5.3-6.0) in SL.
Pectoral fin length 5.8 (5.4-6.3) in SL. Posterior tip of pelvic
fin reaching second (third to fourth) anal fin ray; pelvic fin
length 3.7 (3. 5-3 .8) in SL.

Color in alcohol. Head and body pale brown dorsally, pale
yellowish ventrally; dorsal surface of head with small, round,
brown spots; body without spots and bands dorsally; side
below lateral line with one pale grayish longitudinal stripe
(grayish stripe formed by continuous series of spots in some
paratypes). First dorsal fin with one blackish submarginal
stripe, base clear anteriorly with small scattered black spots;
second dorsal fin with brown (dark brown in some paratypes)
spots. Anal fin pale. Caudal fin with several black longitudinal
narrow stripes and spots posteriorly; upper part of caudal fin
with several brown spots. Pectoral and pelvic fins with small
brown spots (fig. 6d).

32

H.lmamura & M.F. Gomon

Distribution. Northeastern Australia from Townsville (17°51'S)
to Noosa, Queensland (26.25°S), at depths of at least 120-366
m (fig. 1).

Etymology. The specific name ventralis from Latin, meaning ‘of
the belly’, refers to this species’ characteristic long pelvic fin.

Remarks. R. ventralis is most similar to R. diversidens, from
which it can be distinguished as discussed in ‘Remarks’ in the
above treatment of the latter. This species is poorly represented
in museum collections and has been mostly overlooked in the
literature. It has been considered by authors, which have treated
specimens, as simply northern records of R. diversidens.

Acknowledgments

We are indebted to D. Bray (NMV), H. Endo (BSKU), A.
Graham (CSIRO), J. Johnson (QM), Y. Kai (FAKU), M.
McGrouther and S. Reader (AMS), S. Morrison (WAM), G.
Shinohara (NSMT) and J. T. Williams (USNM) for making
material available and conveying information about collection
specimens. We also express sincere thanks to L. W. Knapp and
S. Raredon (USNM), for providing data and photographs,
respectively, of a USNM specimen of Ratabulus megacephalus
from the Philippines. We are grateful to K. Graham and
CSIRO for allowing us to reproduce their color photographs.

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Francisco.

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Hoese, D.F., Bray, D.J., Paxton, J.R. and Allen, G.R. 2006. Fishes. Pp.
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taxa (Pisces: Scorpaeniformes). Speces Diversity 1: 123-233.
Imamura, H. 1997. Platycephalidae, Bembridae, Plectrogeniidae (in
part), Platycephalidae, Hoplichthyidae. Pp. 218-221 in: Okamura,
O., and Amaoka, K. (eds). Sea fishes of Japan. Yama-Kei
Publishers Co., Ltd.: Tokyo (in Japanese).

Imamura, H. 2010. A new species of the flathead genus Inegocia
(Teleostei: Platycephalidae) from East Asia. Bulletin of National
Museum of Natural Sciences, Series A, Supplement 4: 21-29.

Jordan, D.S. and Hubbs, C.L. 1925. Record of fishes obtained by David
Starr Jordan in Japan, 1922 .Memoirs of the Carnegie Museum 10:
93-346, pis. 5-12.

Kamohara, T. 1952. Revised descriptions of the offshore bottom-fishes
of Proc. Tosa, Shikoku, Japan. Reports of the Kochi University,
Natural Science (3): 1-122.

Kamohara, T. 1964. Revised catalogue of fishes of Kochi Prefecture,
Japan. Reports of the USA Marine Biological Station 8: 1-99.

Kim, I.S., Choi, Y„ Lee, C.L., Lee, Y.J., Kim, B.J. and Kim, J.H. 2005.
Illustrated book of Korean fishes. Kyohak Publishing, Co., Ltd.:
Seoul, (in Korean).

Knapp, L.W. 1999. Platycephalidae. Pp. 2385-2421 in: Carpenter,
K.E., and Niem, V.H. (eds). FAO species identification guide for
fishery purposes. The living marine resources of the western
Central Pacific, vol. 4. Bony fish part 2 (Mugilidae to Carangidae).
FAO: Rome.

Knapp, L.W. 2000. Family Platycephalidae (flathead). Pp. 607-608 in:
Randall, J.E., and Lim, K.K.P. (eds ). A checklist of the fishes of the
South China Sea. The Raffles Bulletin of Zoology 2000
Supplement (8): Singapore. 607-608.

Knapp, L.W., Imamura, H. and Sakashita, M. 2000. Onigicia
bimaculata, a new species of flathead fish (Scorpaeniformes:
Platycephalidae) from the Indo-Pacific. Special Publication of the
J. L. B. Smith Institute of Ichthyology (64): 1-10.

Lee, C.L. and Joo, D.S. 1998. Taxonomic review of flathead fishes
(Platycephalidae, Scorpaeniformes) from Korea. Korean Journal
of Ichthyology 10: 216-230.

Matsubara, K. 1955. Fish morphology and hierarchy. Ishizaki Shoten:
Tokyo, v, viii, xi, 1605, 135 pis (in Japanese).

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of the family Platycephalidae (the flatheads). Memoirs of the
College of Agriculture, Kyoto University, (68): 1-109.

McCulloch, A.R. 1914. Report on some fishes obtained by the FIS
Endeavour on the coasts of Queensland, New South Wales,
Victoria, Tasmania, south and southwestern Australia. Part II.
Zoological Results of the Fishing Experiments carried on by FIS
Endeavour 2: 77-165, pis. 13-34.

McCulloch, A.R. 1929. A check-list of the fishes recorded from
Australia. Part 3. Memoirs of the Australian Museum 5: 329-436.

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University of Agriculture 1: 1-228, map.

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Nakabo, T. (ed.). Fishes of Japan with pictorial keys to the species,
English edition. To kai University Press: Tokyo.

Ochiai, A. 1984. Platycephalidae. Pp. 321-322, pis 288-289 in:
Masuda, H., Amaoka, K., Araga, C., Uyeno, T., and Yoshino, T.
(eds). The fishes of the Japanese Archipelago. English text and
plates. Tokai University Press: Tokyo.

Okada, Y. and Matsubara, K. 1938. Key to the fishes and fish-like
animals of Japan. Sanshodo: Tokyo, xl, 584, 113 pis. (n Japanese).

Paxton, J.R., Hoese, D.F., Allen, G.R. and Hanley, J.E. 1989. Zoological
catalogue of Australia. Vol. 7. Pisces. Petromyzonidae to
Carangidae. Australian Government Publishing Service:
Canberra, xii, 664.

Sadovy, Y. and Cornish, A.S. 2000. Reef fishes of Hong Kong. Hong
Kong University Press: Hong Kong, xi, 321.

Sainsbury, K.J., Kailola, PJ. and Leyland, G.G. 1985. Continental
shelf fishes of northern and north-western Australia. Clouston
and Hall, and Peter Pownall Fisheries Information Service:
Canberra, vii, 375.

Revision of Ratabulus

33

Shao, K.T. and Chen, J.P. 1987. Fishes of the family Platycephalidae
(Teleostei: Platycephaloidei) of Taiwan with descriptions of two
new species. Bulletin of the Institute of Zoology, Academia Sinica
26: 77-94.

Shao, K.T. and Chen, J.P. 1993. Platycephalidae. Pp. 255-260 in: Shen,
S.C. (ed.). Fishes of Taiwan. Department of Zoology, National
Taiwan University: Taipei.

Shinohara, G., Endo, H., Matsuura, K., Machida, Y. and Honda, H.
2001. Annotated checklist of the deepwater fishes from Tosa Bay,
Japan. National Science Museum Monographs (20): 283-343.

Tanaka, S. 1917. Eleven new species of fish from Japan. Zoological
Magazine 29: 7-12 (in Japanese).

Tanaka, S. 1931. On the distribution of fishes in Japanese water.
Journal of the Faculty of Science, Imperial University of Tokyo.
Sect. 4, Zoology 3: 1-90, 3 pis.

Yatou, T. 1985. 371 Ratabulus megacephalus (Tanaka). Pp. 598-599,
729 in: Okamura, O.,

Machida, Y., Yamakawa, T., Matsuura, K., and Yatou, T. (eds). Fishes
of the Okinawa Trough and the adjacent waters , vol. 2. The
intensive research of unexploited fishery resources on continental
slopes (in Japanese with English description). Japan Fisheries
Resource Conservation Association: Tokyo.

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list. Academy Book: Seoul, 747 (in Korean).

Memoirs of Museum Victoria 67: 35-44 (2010)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia

Timothy Holland 1 ’ 2

1 School of Geosciences, Monash University, Vic, 3800, Australia

2 Museum Victoria, G.P.O. Box 666, Melbourne, Vic, 3001, Australia (tholland@museum.vic.gov.au)

Abstract Holland, T. 2010. Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia. Memoirs of Museum

Victoria 67: 35-44.

The Genoa River Trackway site in the Upper Devonian Combyingbar Formation represents one of the earliest
records of tetrapods (land vertebrates) from Australia. However, the osteichthyan assemblage from the site is poorly
known compared to other Devonian tetrapod localities. New information from a tetrapodomorph fish lower jaw possibly
indicates the first record of a tristichopterid from the Genoa River Beds . The specimen shares a posteroventral embayment
in the profile of lower jaw with Eusthenopteronfoordi, Platycephalichthys bischojfi, and Moroccan tristichopterid material,
as well an isolated specimen of the ‘osteolepidid’ Gyroptychius . Polyplocodont and dendrodont teeth are also described
from the Genoa River, with the latter indicating the presence of a large, porolepiform taxon. The antiarch placoderm
Remigolepis represents the most abundant fossil fish taxon recorded from the Genoa River. The possible presence of
phyllolepid placoderm material may support previous suggestions of a pre-Famennian age for the Genoa River Beds.

Keywords tristichopterid, Tetrapodomorpha, Genoa River, Devonian, polyplocodont, dendrodont

Introduction

The fine-grained sandstone of the Genoa River Beds
(-Combyingbar Formation [Vandenberg et al., 1992]) along
the Genoa River, southeastern Victoria, is renowned for
preserving the first record of Devonian tetrapods from
Australia and Gondwana (Warren and Wakefield, 1972).
Discovered by Norman A. Wakefield in 1971, this material
was described as three distinct tetrapod trackways by Warren
and Wakefield (1972) and Clack (1997), with subsequent work
focusing mainly on the two better preserved footprint sets,
believed to be distinct in form (Clack, 2002; Young, 2006;
2007). Biostratigraphic and lithological indicators summarised
by Young (2006) suggested a Frasnian age for the locality, but
did not describe any fossils to support this. Dunn (1897), Hall
(1959) and Douglass (1974) recorded Upper Devonian plant
material from the Genoa River, with the strata broadly
corresponding to the Merrimbula Group of the New South
Wales south coast (Lewis et al., 1994), more specifically,
lithofacies 1-3 of the Early to Mid-Frasnian, Twofold Bay
Formation (Simpson et al., 1997; Young, 2007). The age of the
Genoa River trackways are notable for being ‘...probably
contemporary with Obruchevichthys and Elginerpeton ’
(Clack, 2002; p. 92), previously considered to be the oldest
known tetrapods (Ahlberg, 1995). However, the recent
discovery of a tetrapod trackway site from the Eifelian
Northern Holy Cross Mountains, Poland (Niedzwiedzki et al.,
2010) is significantly older than this aforementioned material.

In contrast to the importance of the Genoa site, none of the
vertebrate assemblage has been described. Such details have
been published regarding the fauna from Northern Hemisphere
tetrapod localities, such as those from East Greenland
(summarised by Blom etal., 2007). The record of the Devonian
fish fauna from the Genoa River includes the placoderm taxa
Bothriolepis, Remigolepis and Groenlandaspis (Young, 1988)
and a phyllolepid taxon (recorded in the field notes of Anne
Warren [Anne Warren, La Trobe University, pers. comm.,
2009]). Isolated ‘crossopterygian skull elements’ have also
been reported from the site (Young, 1988; p. 192), along with
bone elements and scales from the porolepiform fish
Holoptychius (Young, 1993), although this record has since
been questioned (Young, 2007). A large, poorly preserved
partial lower jaw of an ‘osteolepiform’ fish was identified by
Ahlberg and Clack (1998), based on a broad distribution of
denticles on the prearticular and a posterodorsally directed
glenoid surface of the articular. New descriptive work on this
specimen, NMV P198470, is presented here following further
preparation. Several similarities to tristichopterid specimens
from the Frasnian of Canada, Russia and the Famennian of
Tafilalt, Morocco, are suggested. Two distinct sarcopterygian
fish tooth morphologies from the Genoa River fauna are also
described in detail for the first time. This includes a small
polyplocodont tooth and a large dendrodont tusk. The former
morphology is known from tetrapodomorph fishes and lower
tetrapods, while the latter is unique to the Porolepiformes

36

T. Holland

(Schultze, 1970). A list is presented of the fossil fish specimens
collected from the Genoa River and stored in Museum
Victoria. The biostratigraphic implications of the Genoa River
fauna are discussed, including the possible presence of
phyllolepid material.

Material and methods

The aforementioned specimens were collected in 1973 by Ian
Stewart on a field trip led by James W. Warren along the bank
of the Genoa River, Victoria, Australia. Preliminary work on
the jaw (NMV P198470) was carried out by Alec L. Panchen
and Anne Warren, but was never published (Anne Warren, La
Trobe University pers. comm., 2009). Manual surface
preparation was undertaken by Per E. Ahlberg and Jennifer A.
Clack (Ahlberg and Clack, 1998), with bone material being
dissolved in hydrochloric acid by T.H in 2008. A black latex
peel of the specimen was then dusted with sublimate of
ammonium chloride and photographed with a Nikon D80
camera using a Nikon DX 18-135 mm lens. The polyplocodont

(NMV P229479) and dendrodont (NMV P229477) teeth were
sectioned by Ian Stewart, and photographed with a Leica
DFC500 camera using a Leica M205C microscope.
Anatomical terminology follows that of Jeffery (2003).

Institutional abbreviations

NMV, Museum Victoria, Melbourne; MNHN, Museum
national d’Histoire naturelle, Paris, France; NRM,
Naturhistoriska riksmuseet (Natural History Museum of
Sweden).

Systematic descriptions

Superclass Osteichthyes Huxley, 1880
Class Sarcopterygii Romer, 1955

Horizon. Combyingbar Formation (Vandenberg et al., 1992)
Order Tetrapodomorpha Ahlberg, 1991

Nungatta Creek

Yambulla Peak

Yambulla Creek

Genoa River

Keyhole Creek

Trackway site

Murmuring Creek

Mt Merragunegin

NMV PI 98470

Watervine Creek

Wangarabell Road

New South Wales

Victoria

Yambulla
Peak Trail

Wangarabell Creek

Figure 1. Geological map of Genoa River area, Genoa, Victoria, Australia (modified from Marsden, 1976; Simpson et al., 1997); grey = Late
Devonian, Combyingbar Formation; diagonal lines = Early Devonian, Maramingo Granite; white = Ordovician slate, sandstone.

Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia

37

tetrapodomorph gen. et sp. indet.

Large incomplete lower jaw (Ahlberg and Clack, 1998; p. 35)

Poorly preserved lower jaw; belonged to an osteolepiform fish
(Young, 2006; p. 413)

Poorly preserved osteolepiform sarcopterygian lower jaw (Young,
2007; p. 1003)

Referred specimen. NMV P198470, an incomplete lower jaw, including
infradentaries, prearticular, articular, and possible dentary and
coronoid elements.

Locality. Along the Genoa River, between the branches of
Murmuring Creek and Keyhole Creek (fig. 1).

Description. Only the lingual surface of NMV PI 98470 is
preserved (figs 2a-b), consisting of an elongate, relatively
narrow, posterior half of a lower jaw. It measures 18 cm long
from the preserved anterior to the posterior tip of the articular,
and 5 cm deep from the level of the infradentaries to the dorsal
margin of the dentary. Parts of the dorsal and ventral margins are
incomplete, most notably the dorsal area anterior to the articular.
Most of the surface of the jaw is covered by the prearticular
(fig. 2b), which anteriorly carries small patches of denticles (fig.
2b) near the dorsal margin. These patches are replaced posteriorly
by a roughened surface of bone, which is marked with a
conspicuous crescent-shaped contour (fig. 2b) towards the
posterior margin of the prearticular.

Posterior to the prearticular is the articular (fig. 2b), which
displays a posterodorsally directed, concave glenoid surface. It is
bordered anterodorsally and posteroventrally by two rounded
prongs, with the former being preceded by a prominent concave
dip in the preserved dorsal margin of the jaw. This could possibly
represent the inner margin of the adductor fossa (fig. 2b).

Anterior to this region, the dorsal margin of the jaw is
represented by two strips of bone: the dentary (fig. 2b), which is
preserved anteriorly, and a presumed coronoid (fig. 2b),
represented by a posteriorly positioned ledge that sits lingual to
the dentary. Both the dentary and the coronoid are incompletely
preserved.

Although the ventral margin of the lower jaw is incomplete
anteriorly, a small slither of bone positioned labially to the
prearticular may represent the third infradentary (fig. 2b). It is
followed by a small break, and then by a similar bone, possibly
the fourth infradentary (fig. 2b). This bone is bordered dorsally
by a narrow groove, which rises transversely to delineate the
margin of the prearticular from a region of exposed meckelian
bone (fig. 2b). A small depression is present in this area, ventral
to the glenoid surface of the articular. This may represent the
opening for the ramus mandibularis of n. facialis (fig. 2b). The
posteroventral outline of the jaw shows a distinct concave, step-
like margin towards the articular (fig. 2b).

Comparisons to other taxa. Affinities of NMV PI 98470 to
various osteichthyan clades can be eliminated quickly through
comparisons of lower jaw gross morphology. NMV P198470
differs from Devonian ‘palaeoniscid’ fishes such as ‘Mimia’ and
Moythomasia (Gardiner, 1984) (fig. 2c) in lacking a dorsally
directed glenoid fossa on the articular (and also the latter by the
absence of a double prearticular); Devonian actinistians, such as
Miguashaia (Forey et al., 2000) (fig. 2d) and Styloichthys

(Friedman, 2007) by lacking a ventral mandibular flange
protruding below the level of the infradentaries and the
‘sympletic’ articulation at the posterior of the jaw; Devonian
dipnoans, such as Chirodipterus and Rhinodipterus (Jarvik,
1967) by having a prearticular, coronoid series and lacking a
principle tooth plate; and porolepiforms, such as Holoptychius
(Jarvik, 1972) (fig. 2e) by lacking a rounded posteroventral
margin and large dorsoventral depth. Among tetrapodomorphs,
NMV PI 98470 differs from rhizodontids by lacking an unossified
articular and other meckelian elements (Jeffery, 2003) (fig. 2f);
Devonian tetrapods, such as Ventastega (Ahlberg and Clack,
1998) (fig. 2g) by lacking an articular situated posterodorsally to
the dentary tooth row; and most ‘osteolepidids’, such as
Ectosteorhachis (Thomson, 1964) (fig. 2h) in having a gentle,
transverse posteroventral outline towards the articular.

NMV PI 98470 is comparable to Platycephalichthys bischoffi
(Vorobyeva, 1962: plates XVI-XVII) and some specimens of
Eusthenopteron foordi (fig. 2i; Jarvik, 1980: fig. 125), in which
the posteroventral margin of the lower jaw displays a concave
profile. This embayment is also present in the unnamed
tristichopterid specimen MNHN n° MCD 42 from Tafilalt,
Morocco (Lelievre and Janvier, 1986) (fig. 2k), which also shares
the acute crescent shaped contour on the posterior region of the
prearticular (fig. 2k). Among the lower jaw morphology of other
tristichopterids, a posteroventral concave profile is absent from
Eusthenopteron save-soderberghi (Vorobyeva, 1962: plate I), E.
kurshi (Zupins, 2008), Jarvikina wenjukowi (Vorobyeva, 1962:
plate XXVIII), Eusthenodon wangsjoi (Jarvik, 1952) and
Tristichopterus (Traquair, 1875). A posteroventral embayment is
also present in the lower jaw of Gyroptychius specimen NRM PZ
P1409 (fig. 2j), although this element is preserved in labial view.

tetrapodomorph gen. et sp. indet.

Referred specimen. NMV P229479, small parabasally sectioned,
polyplocodont tooth.

Locality. Along the Genoa River, approximately 30-50 m
upstream from the Yambulla Peak Track (fig. 1).

Description. The specimen, represented by a single tooth (figs
3a-c), measures approximately 1.5 cm from the apex to the
preserved parabasal section, and is 8 mm in diameter. Several
loose plications (fig. 3a) are present at the base of the external
surface. These extend approximately one -third of the total
height of the tooth, and are replaced apically by a smooth layer
of enamel. The pulp cavity (fig. 3b) is visible in parabasal
section, and seems to be free from osteodentine. It is surrounded
by loose, simple folds of orthodentine (figs 3b-c), which appear
to be interrupted intermediately by bone (fig. 3c).

Comparisons with other taxa. Tooth morphology of NMV
P229479 closely matches that of polyplocodont teeth in having
a free pulp cavity, simple orthodentine folds and bone of
attachment extending between folds (Schultze, 1970: fig. la).
This morphology is present in rhizodontids, Megalichthys,
Eusthenopteron and Tristichopterus (Schultze, 1970). However,
the lingual surface of NMV P229479 differs from that of
rhizodontids, such as Barameda (Holland et al., 2007) in
lacking fine, raised striae on the enamel.

38

T. Holland

Adductor fossa . Crescent on particular

Denticles

Infradentary four

Dentary

Coronoid

Prearticular

Infradentary three

Meckalian groove

Concavity in the
posteroventral margin
of the lower jaw

Foramen for
the ramus
m and ibu laris
of n. facialis

Articular

Concavity in the posteroventral margin
of the lower jaw

Crescent on prearticular

Concavity in the posteroventral margin
of the lower jaw

Figure 2. Osteichthyan lower jaws; a, photograph of possible tristichopterid NMV P198470; b, interpretive drawing of NMV P198470; c,
Moythomasia (modified from Gardiner, 1984); d, Miguashaia (modified from Forey et al., 2000); e, Holoptychius (modified from Jarvik, 1972);
f, Rhizodus (modified from Jeffery, 2003); g, Ventastega (modified from Ahlberg and Clack, 1998); h, Ectosteorhachis (modified from Thomson,
1964); i, Eusthenopteron specimen NRM PZ P35; j, Gyroptychius specimen NRM PZ P1409; k, tristichopterid specimen MNHN n° MCD 42
(modified from Lelievre and Janvier, 1986). Scale = 1 cm; scale not included for c-h; all jaws excluding j are in lingual view; j is in labial view.

Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia

39

Bone of attachment

Orthodentine

Orthodentine

Orthodentine

Plication

Orthodentine

Osteodentine

Osteodentine

Figure 3 . Tooth morphologies of tetrapodomorph fishes; a, labial view of polyplocodont tooth, NMV P229479; b, stylised drawing showing complete
parabasal section of NMV P229479; c, orthodentine folding of NMV P229479 in parabasal section; d, vertical section of dendrodont tooth, NMV
P229477; e, orthodentine folding of NMV P229477; f, parabasal section of NMV P229477. Scale = 5 mm; scale not included for c, e.

Order Porolepiformes Jarvik, 1942

? porolepiform gen. et sp. indet.

Referred specimen. NMV P229477, a large, vertically and
parabasally sectioned dendrodont tooth element.

Locality. Along the Genoa River, approximately 30-50 m
upstream from the Yambulla Peak Track (fig. 1).

Description. Specimen NMV P229477 is represented by a
large fang (figs 3d-f), being approximately 6 cm in height,
although parts of the base may not have been collected. The
external surface is badly damaged, preserving no plications,
with the proximal end sectioned vertically, exposing the pulp
cavity (figs 3d-f). The width of the pulp cavity varies, being
approximately half the total width of the specimen (total
width - 2.2 cm) towards the proximal end, to be only one-fifth
of the total diameter approaching the apex. Inside the pulp
cavity is a granular column of bone, probably representing
osteodentine (figs 3d, f). Lateral to the pulp cavity are wide
expanses of orthodentine (figs 3d-f), which are marked with
fine, densely packed lines, representing folding. This intense
folding is more apparent in parabasal section, and appears
darker and tighter towards the pulp cavity (fig. 3e). Several
regions of orthodentine are interrupted by channels of
osteodentine emanating from the pulp cavity (fig. 3f). In some
instances, these channels appear to bifurcate and narrow as
they approach the outer surface of the tooth.

Comparisons with other taxa. The combination of complicated
folded orthodentine and a pulp cavity filled with osteodentine in
NMV P229477 closely matches the description of dendrodont
tooth morphology from porolepiform fishes, such as Porolepis
(Schultze, 1970) and Laccognathus (Schultze, 1969). There is
no intrusion of outside bone of attachment between the folds of
orthodentine, in contrast to eusthenodont teeth (Schultze, 1970).

Discussion

Of the osteichthyan remains from the Genoa River Beds, the
lower jaw of NMV P198470 may offer a link to tristichopterid
taxa from outside East Gondwana in having a posteroventral
embayment in the profile of the lower jaw. This particular
feature is present in the Eusthenopteron- like tristichopterid
material from Famennien of Morocco (Lelievre and Janvier,
1986), and some specimens of Eusthenopteron foordi from the
Frasnian of Miguasha, Quebec, Canada (e.g. NRM PZ P35
from the Swedish Museum of Natural History) . Eusthenopteron
is considered among the more primitive members of the
Tristichopteridae, along with the Scottish Givetian taxon,
Tristichopterus (Ahlberg and Johanson, 1997). A possible
relationship between these taxa and NMV P198470 is
surprising, because the record of tristichopterids in Australia
otherwise consists of mandageriids. This includes
Cabonnichthys Ahlberg and Johanson, 1997 and Mandageria
Johanson and Ahlberg, 1997 from the Fransian Mandagery
Formation of Canowindra, New South Wales; Eusthenodon

40

T. Holland

from the Famennian of the Hunter Formation of Grenfell, New
South Wales (Johanson, 2004); and the Worange Point
Formation, south of Eden, New South Wales (Ahlberg et al.,
2001). Eusthenodon was initially described from the
Famennian of East Greenland (Jarvik, 1952). However, the
assignment of the Eden tristichopterid within Eusthenodon has
been questioned by Young (2008), placing it with the
Canowindra tristichopterids in a possibly endemic Gondwanan
subfamily — the Madageriidae — based on the presence of
accessory vomers on the palate. The only other notable
Australian taxon previously linked with the Tristichopteridae
is Marsdenichthys Long, 1985 from the Givetian of Mt.
Howitt, Victoria. This taxon has also been associated with the
Rhizodopsidae (Long, 1999), although new material described
by Holland et al. (2010) shows no affinities to this clade or the
Tristichopteridae. In addition, the lower jaw of Marsdenichthys
(Holland et al., 2010: fig. 2) lacks the posteroventral
embay ment of NMV PI 98470 and some tristichopterid taxa.
However, evaluating the distribution of the posteroventral
lower jaw concavity seen in NMV PI 98470 throughout the
Tristichopteridae requires caution, because the area is either
not well preserved or described as the Madageriidae (e.g.
Young et al., 1992; Ahlberg and Johanson, 1997; Johanson and
Ahlberg, 1997) and Langlieria (Clement, 2002); and appears
absent from Eusthenodon wangsjdi (Jarvik, 1952), Jarvikina
(Vorobyeva, 1962), the European species of Eusthenopteron
(e.g. Vorobyeva, 1962; Zupins, 2008) and the basal
Tristichopterus (Traquair, 1875). A poorly developed
posteroventral lower jaw concavity also appears in E. foordi
specimen P.2197 (Jarvik, 1996: fig. 19), although the relevant
area in this specimen is possibly crushed. Outside the
Tristichopteridae, a posteroventral embayment is also present
in the lower jaw of Platycephalichthys from the Frasnian of
Russia (Vorobyeva, 1962). However, this taxon, which was
previously associated with tristichopterids, shares some
affinities with ‘elpistostegid’ fishes, such as Panderichthys
(Coates and Friedman, in press), and thus may occupy a more
crownward phylogenetic position. It is also worth noting that a
concave posteroventral profile marks the lower jaw of the
cosmine-covered ‘osteolepidid’ Gyroptychius specimen, NRM
PZ P1409. Although no cosmine was recorded on the scant
labial surface material of NMV P198470 (before preparation),
the presence of a posteroventral notch in the jaws of both
specimens is potentially significant, as Gyroptychius is the
sister taxon to the Tristichopteridae in Ahlberg and Johanson
(1998) and shares a number of other characters with the group,
including vomers with long posterior processes, an elongate
ethmosphenoid block, and a trifurcate tail. However, the lower
jaws of several specimens of G. agssizi (Jarvik, 1948: fig. 74),
G. milleri (Jarvik, 1948: fig. 80) and G. groenlandicus (Jarvik,
1950: fig. 21) have rounded posteroventral profiles. Thus,
posteroventral jaw morphology may vary within Gyroptychius.
Alternatively, this region may be damaged in NRM PZ P1409
and may not represent a shared character between Gyroptychius
and the Tristichopteridae. Furthermore, Spodichthys from the
Frasnian of East Greenland has been shown to be the immediate
sister taxon to the Tristichopteridae (Snitting, 2008),
subsequent to the analysis of Ahlberg and Johanson (1998).

This taxon displays a rounded posteroventral lower jaw profile
(Snitting, 2008: fig. 7). Thus, it appears unlikely that concave
posteroventral mandible morphology was present in
Gyroptychius, was lost in Spodichthys and Tristichopterus, and
then re-evolved at the node containing Eusthenopteron.

The phylogenetic relationships of the polyplocodont tooth
NMV P229479 are difficult to discern, based on the widespread
prevalence of similar tooth morphology throughout several
tetrapodomorph groups, including rhizodontids,
megalichthyids, tristichopterids, ‘elpistostegids’ and early
tetrapods (Schultze, 1970). Based on relative size and spatial
proximity to each other, it is possible that the lower jaw of
NMV PI 98470 and NMV P229479 are attributable to the same
form, although several other Palaeozoic fish sites are known to
contain multiple tetrapodomorph fish genera (e.g. Mt. Howitt,
Holland et al., 2010). Thus, isolated elements such as these
must be interpreted with caution, because they may belong to
different taxa.

The use of polyplocodont and eusthendont tooth
morphologies as phylogenetic indicators may help clarify the
relationships of advanced tristichopterids. In the cladistical
analysis of Clement et al. (2009), Eusthenodon and Langlieria
are grouped in an apical clade with Mandageria and
Cabonnichthys . However, the former two taxa are described as
having eusthenodont teeth (Schultze, 1970; Clement, 2002),
while polyplocodont teeth are recorded from the later (Ahlberg
and Johanson, 1997; Johanson and Ahlberg, 1997). It would be
of great interest to determine the tooth morphology of the Eden
tristichopterid to compare with the Northern Hemisphere
specimens of Eusthenodon and the tristichopterids from
Canowindra, to ascertain phylogenetic information. Curiously,
the use of eusthenodont tooth morphology as a character is not
included in the phylogenetic analysis of Ahlberg and Johanson
(1997) or Johanson and Ahlberg (1997), with Eusthenodon
coded with Mandageria and Cabonnichthys as having
polyplocodont teeth. It is not stated in the relative literature
whether the teeth of Mandageria (Johanson and Ahlberg,
1997) and Cabonnichthys (Ahlberg and Johanson, 1997) have
been sectioned or are only known from natural moulds, as in
other tetrapodomorph fishes from Canowindra, such as
Gooloogongia (Johanson and Ahlberg, 2001). Among other
tetrapodomorph fishes, eusthenodont teeth have also been
described from Platycephalichthys (Vorobyeva, 1959) and
Litoptychus (Schultze and Chorn, 1998). This latter form,
known from the Frasnian of Colorado, United States of
America (Schultze and Chorn, 1998), is placed as the sister
taxon to the Megalichthyidae in Coates and Friedman (in
press).

The identification of NMV P229477 as a dendrodont tooth
confirms the record of porolepiform taxa among the Genoa
River fauna. In overall proportions, NMV P229477 is slightly
larger than the vomerine fang of Barameda decipiens from the
Carboniferous of Mansfield, Victoria (Holland et al., 2007), a
taxon probably 3^4 m in total body length (pers . obs ., 20 1 0) . As
some specimens of the cosmopolitan porolepiform Holoptychius
exceed 2.5 m in length (Long, 1995), it is not unreasonable to
suggest a very large body size for NMV P229477, possibly
among the largest recorded for porolepiform fishes.

Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia

41

Of the faunal assemblage recorded from the Genoa River Beds
(table 1), the most common taxon represented in the collections of
Museum Victoria is the antiarch placoderm, Remigolepis, with up
to eight registered specimens. Several large incomplete antiarch
plates collected from the Genoa River (ANU 3269-3272) are also
stored in the Research School of Earth Sciences, Australian
National University (Gavin Young, ANU pers. comm., 2010).
High occurrences of Remigolepis have been noted from the Upper
Famennian tetrapod sites of Ningxia, China and East Greenland
(Lebedev, 2004) . Further similarities between the fossils localities
of the Genoa River and East Greenland also occur in the presence
of other taxa. As with the Genoa River Beds, the faunal assemblage
of the Aina Dal Formation of East Greenland includes Remigolepis
and Bothriolepis, as well as phyllolepid, tristichopterid,
porolepiform and tetrapod material (Blom et al., 2007). The
remains of Groenlandaspis, also reported from the Genoa River,
are possibly present in the younger Harder Bjerg Formation of
East Greenland (Blom et al., 2007). Plates attributed to
Groenlandaspis from the Genoa River (e.g. NMV PI 86587)
show evenly spaced, fine tubercles, similar to the condition in
regularly ornamented specimens of Groenlandaspis (e.g.
Daeschler et al., 2003: figs 2, 4). As stated by Young (2006), this
material contradicts spurious reports of a smooth, unomamented

form of Groenlandaspis from the Genoa River by Young (1993),
which had been used previously to equate the age of the Genoa
River Beds to that of the Famennian Worange Point Formation.

Of the sarcopterygian material collected from the Genoa
River, the rounded scales of specimen NMV P230291 are not
characteristic of the genus Holoptychius, reported from the
area by Young (1993). The abovementioned sarcopterygian
scales stored in Museum Victoria are marked with fine radiating
ridges on the exposed external surface, similar to those
preserved in some rhizodontids (e.g. Holland et al. 2007: fig. 4)
and non-madageriids tristichopterids (e.g. Jarvik, 1952: fig.
30c). This pattern is distinct from the lateral scales of
Holoptychius, in which the exposed external surface bares
much thicker, broadly separated, radiating lines (Cloutier and
Schultze, 1996: fig 11a). Aside from the examples described in
this paper, several other sarcopterygian teeth have been
collected from the Genoa River. Although varying in size and
shape, it is difficult to discern the affinities of these specimens
without obtaining detailed cross-sections. These specimens
include NMV P229478, a partial jaw with several teeth. In
contrast to the possible tristichopterid jaw NMV P198470, the
incomplete nature of NMV P229478 does not allow detailed
comparisons with known sarcopterygian forms.

Table 1. List of Museum Victoria specimens collected from the Genoa River.

Museum number

taxon

material

NMV P229529

Remigolepis

Plate

NMV P229545

Remigolepis

Plate

NMV P230039

Remigolepis

Plates and scales

NMV P230040

Remigolepis

Plate

NMV P230041

Remigolepis

Plate

NMV P229544

Remigolepis

Plates, shoulder joint

NMV P186587

Groenlandaspis

Plate

NMV P186582

Groenlandaspis and Remigolepis

Plate impressions

NMV P230038

Bothriolepis, Remigolepis and ?phyllolepid

Plate impressions

NMV P230683

Placoderm indet.

Plate

nMV P230682

Placoderm indet.

Plate

NMV P229476

Placoderm indet.

Plate

nMV P230042

Sarcopterygian indet.

Tooth

nMV P230043

Sarcopterygian indet.

Tooth

nMV P229478

Sarcopterygian indet.

Partial jaw and teeth

nMV P230291

Sarcopterygian indet.

Scales

nMV P229477

Porolepiform indet.

Tusk

nMV P229479

Tetrapodomorph indet.

Tooth

nMV PI 98470

Tetrapodomorph indet.

Jaw

nMV P41321

Tetrapod indet.

Trackways 1 and 2

nMV P41322

Tetrapod indet.

Trackway 3

42

T. Holland

In regards to the age of the Genoa River Beds, the potential
record of phyllolepid material is compelling as abiostratigraphic
indicator. This record consists of elements observed in the field
by Anne Warren (pers comm., La Trobe University, 2009), as
well as the impression of a plate on specimen NMV P230038
(table 1), a large block red sandstone. This material is of great
importance, as the record of phyllolepids from Australia
extends to Givetian and Frasnian fossil sites (Young, 2005), in
contrast to the Famennian range for the group from Euramerican
localities (e.g. Blom et al., 2007). This record could support an
earlier, possibly Frasnian, age for the Genoa River localities, as
suggested by Young (2006). It should be noted that although
five phyllolepid genera have been described from the pre-
Famennian sites in Australia (listed in Young, 2010),
indeterminate phyllolepid material has also been reported from
other Famennian sites in East Gondwana (Young, 2005: fig. 4).
Furthermore, the possible impression of a phyllolepid plate on
NMV P230038 consists of a small, incomplete region of
parallel ridged ornament. This region could alternatively be
interpreted as the parallel, laterally running ridges marking the
anterior ventrolateral plate of Remigolepis (e.g. Johanson, 1997:
fig 12e). Nevertheless, the report of phyllolepid plates in the
field (Anne Warren, pers comm., 2009) possibly provides
significant new information regarding the age of the Genoa
River Beds. Of the other placoderm taxa reported from the
Genoa River, Bothriolepis, Remigolepis, and Groenlandaspis
have been recorded from both Frasnian (e.g. Canowindra;
Young, 2008) and Famennian (e.g. Grenfell; Johanson, 1997)
fossils sites from Australia.

The possibly Frasnian age of the Genoa River Beds does
not contradict current ideas regarding the occurrence and
dispersal of tristichopterid and porolepiform fishes throughout
the early Late Devonian. Such hypotheses include the origin of
the Tristichopteridae in Euramerica, based on the presence of
the stem taxon Spodichthys and the earliest member of the
group Tristichopterus in the Northern Hemisphere (Snitting,
2008). The discovery of NMV P198470 from the Genoa River
may support the dispersal of the group to Gondwana as early
or before the Early-Mid Frasnian, rather than at the Frasnian-
Famennian boundary. This scenario is possibly contradicted
by the presence of tristichopterid material from the Givetian
Bunga Beds, on the south coast of New South Wales (Young,
2007; 2008). However, this material, including scales and an
isolated cleithrum, does not include any known tristichopterid
synapomorphies, and should be regarded with caution.

The age of porolepiform material from other East
Gondwana sites far exceeds that of NMV P229477. This
includes elements from the Pragian-Eifelian Dulcie Sandstone
and Cravens Peak Beds of Central Australia (Young and
Goujet, 2003). This East Gondwanan record appears as old as
the earliest porolepiform taxa from the Northern Hemisphere,
such as Porolepis (Jarvik, 1972).

Acknowledgments

I would like to thank Lucinda Gibson of Museum Victoria for
her help with photography, and also Anne Warren of La Trobe
University, Bundoora, John Long of Museum Victoria and

Gavin Young of the Research School of Earth Sciences, The
Australian National University, Canberra, for their shared
discussion regarding this material. Appreciation is also given
to Thomas Mors and Jonas Hagstrom of the Swedish Museum
of Natural History, Stockholm, for aiding access to material,
and to Patricia Vickers-Rich of the School of Geosciences,
Monash University, Clayton, for her supervision. Finally, I
would like to thank Dave Pickering of Museum Victoria for
attempting to contact Ian Stewart. T.H. is supported by a
Monash University Dean’s Postgraduate Scholarship.

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Young, G.C. 1993. Middle Palaeozoic macrovertebrate biostratigraphy
of eastern Gondwana. Pp. 208-251 in: Long, J.A. (ed). Palaeozoic
Vertebrate Biostratigraphy and Biogeography. Belhaven Press,
London.

Young, G.A. 2005. An articulated phyllolepid fish (Placodermi) from
the Devonian of central Australia: implications for non-marine
connections with the Old Red Sandstone continent. Geological
Magazine 142: 173-186.

Young, G.C. 2006. Biostratigraphic and biogeographic context for
tetrapod origins during the Devonian: Australian evidence.
Alcheringa Special Issue 1: 409-428.

Young, G.C. 2007. Devonian formations, vertebrate faunas and age
control on the far south coast of New South Wales and adjacent
Victoria. Australian Journal of Earth Sciences 54: 991-1008.

Young, G.C. 2008. Relationships of tristichopterids (osteolepiform
lobe-finned fishes) from the Middle-Late Devonian of East
Gondwana. Alcheringa 32: 321-336.

Young, G.C. 2010. Placoderms (armored fish): dominant vertebrates of
the Devonian period. Annual Review of Earth and Planetary
Sciences 38: 523-550.

Young, G.C. and Goujet, D. 2003. Devonian Fish Remains from the
Dulcie Sandstone and Cravens Peak Beds, Georgina Basin,
Central Australia. Records of the Western Australian Museum 65:
1-85.

Young, G.C., Long, J.A. and Ritchie, A. 1992. Crossopterygian fishes
from the Devonian of Antarctica: systematic, relationships and
biogeographic significance. Records of the Australian Museum
14: 1-77.

Zupins, I. 2008. A new tristichopterid (Pisces, Sarcopterygii) from the
Devonian of Latvia. Proceedings of the Latvian Academy of
Sciences, Section B 62: 40-46.

Memoirs of Museum Victoria 67: 45-59 (2010)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea:
Decapoda: Caridea) from Australia

Tomoyuki Komai 1 and Joanne Taylor 2

1 Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba, 260-8682 Japan (komai@chiba-muse.
or.jp)

2 Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia (jtaylor@museum.vic.gov.au)

Abstract Komai, T. & Taylor, J. 2010. Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda:

Caridea) from Australia. Memoirs of Museum Victoria 67: 45-59.

Examination of collections from waters off southern Australia resulted in significant findings of three new species
of the crangonid genus Metacrangon Zarenkov, 1965: M. australis sp. nov. from the Southern Ocean off Tasmania, west
of Maquarie Island; M.poorei sp. nov. from the Tasman Sea; and M. spinidorsalis sp. nov. from waters off southwestern
Western Australia and the Tasman Sea. The two former species are morphologically similar to M. variabilis (Rathbun,
1902) from the north Pacific and M. proxima Kim, 2005 from Japan. The third is referred to the informal M.jacqueti (A.
Milne-Edwards, 1881) species group. Differentiating characters of these three new species are discussed and a key to their
identification is provided.

Keywords Crustacea, Decapoda, Caridea, Crangonidae, Metacrangon , new species, key, Australia, Victoria, Western Australia,

Tasmania, Macquarie Island

Introduction

The crangonid shrimp genus Metacrangon Zarenkov, 1965 is
rather diverse, with 26 named species and one subspecies (De
Grave et al., 2009; Komai and Komatsu, 2009; Komai, in
press). It is characterised by the shallowly depressed gastric
region of the carapace, the usual presence of a pair of
submedian teeth on the carapace, the laterally flared pleuron
of the sixth abdominal somite, and the second pleopod with an
appendix masculina being much shorter than the endopod
(Zarenkov, 1965; Butler, 1980; Kim and Hayashi, 2003; Kim,
2005). Christoffersen (1988) supported the monophyly of the
genus. Many of these 27 taxa are rare, reported from limited
geographic locations that are often confined to or near their
type localities. Although species of Metacrangon are well
represented in the north Pacific Ocean, no species of the genus
have been described from Australia. Poore (2004) reported
that unidentified species were known to occur on the
southeastern shelf off eastern Tasmania at around 500-600 m
depth, but to date, these have remained undescribed. A
nominal new species from southwestern Australia was also
recently reported by Poore et al. (2008) from sampling cruises
off the continental margin of Western Australia onboard the
FRV Southern Surveyor in 2005, mounted by CSIRO Marine
and Atmospheric Research (CMAR) and Museum Victoria
(project entitled ‘Mapping benthic ecosystems on the deep

continental shelf and slope in Australia’s southwest region’).

This present study was initiated to describe the new
species from southwestern Australia reported by Poore et al.
(2008), but our examination of the Metacrangon specimens
lodged in Museum Victoria, referred to by Poore (2004),
confirmed two further species, both new to science. In this
paper, these three new species are described and illustrated:
M. australis sp. nov. from waters southeast of Tasmania off
Macquarie Island; M. spinidorsalis sp. nov. (- Metacrangon
sp. Poore, 2004 and Metacrangon sp. MoV 5423 Poore et al.,
2008) from southwestern Australia and the Tasman Sea; and
M. poorei sp. nov. from the Tasman Sea. These three species
occur in rather high latitudinal areas.

The examined material remains in Museum Victoria,
Melbourne (MV) and the Western Australian Museum, Perth
(WAM). The abbreviation ‘sp. MoV’ refers to the unique
Museum Victoria number allocated to new or undetermined
taxa (and is not the same as sp. nov., which refers to a new
species). The measurement provided is of the postorbital
carapace length (cl) measured from the level of the posterior
margin of the orbit to the midpoint of the posterodorsal
margin. In order to avoid unnecessary repetition, only M.
spinidorsalis is fully described and differential descriptions
are given for the other two new species.

46

T. Komai &J. Taylor

Taxonomic account

Metacrangon australis sp. nov.

Figures 1-3

Material examined. Holotype: Australia, Tasmania, southwestern
Pacific west of Macquarie Island (54°42.42'S, 158°45.12'E- 54°41.36'S,
158°43.12'E), 700-900 m, 22-23 Jan 1999, FRV Southern Surveyor ,
epibenthic sled (stn SS01/99/65), NMV J60424 (1 female, cl 13.1 mm).

Paratypes: same data as holotype, NMV J61200 (2 females, cl 9.9,
12.1 mm).

Description. Body (fig. 1) moderately robust. Rostrum (figs
2a-b) narrowly triangular with acute apex in dorsal view,
directed forward or slightly ascending, about 0.20 times as long
as carapace; dorsal surface with middorsal carina in proximal
half; lateral margin slightly convex in lateral view, merging
into orbital margin; midventral carina distinct, ventral margin
nearly straight in lateral view. Carapace (figs 1, 2a) very slightly
widened anteriorly, longer than wide postorbitally; surface
covered with very short setae; dorsal midline with two
moderately small, subequal teeth; anterior (epigastric) tooth
arising at 0.10 of carapace length, posterior (cardiac) tooth
broken off, arising at 0.75-0.80 of carapace length; submedian
and hepatic teeth moderately small; antennal tooth moderately
strong, directed forward in dorsal view, weakly ascending
(same degree as rostrum) in lateral view, acuminate, falling
short of rostral tip; orbital cleft absent; anterolateral margin
between antennal and branchiostegal teeth sinuous with obtuse
lobule (holotype) or with tiny denticle (paratypes) inferior to
base of antennal tooth; branchiostegal tooth moderately strong,
very slightly diverging anteriorly in dorsal view and strongly
ascending in lateral view, distinctly overreaching dorsodistal
margin of antennal basicerite; pterygostomial tooth small,
visible in lateral view; postorbital carina clearly delimited,
accompanied by longitudinal suture; weak epibranchial carina
present.

Thoracic sternites depressed; fifth sternite with small,
forwardly directed median tooth, otherwise unarmed.

Abdomen (figs 1, 2c) moderately sculptured; first to fifth
somites with sharply delimited, crested middorsal carina,
anterior end of middorsal carina on second somite produced
anteriorly. Pleuron of anterior four somites rounded marginally.
Fifth somite with posterodorsal margin slightly produced
medially; posterolateral margin unarmed; pleuron with
posteroventral angle rounded, ventral margin gently convex.
Sixth somite 1.6 times longer than wide, with distinct, slightly
curved submedian carinae, not reaching posterodorsal margin;
dorsolateral carina distinct, reaching to posterodorsal margin;
posterodorsal margin produced, distinctly bilobed; pleuron
flared laterally, with small posteroventral tooth; posterolateral
process moderately strong, directed posteriorly, terminating in
sharp tooth. Telson (figs 2c-d) tapering distally to acute apex,
with three pairs of minute dorsolateral spines, anteriormost
pair located at posterior 0.35; three pairs of spiniform setae
posterior to third pair of dorsolateral spines.

Eye (figs 2a-b) as long as wide; cornea as wide as eyestalk,
darkly pigmented, corneal width 0.13-0.15 of carapace length;
eyestalk with small, papilla-like dorsal tubercle.

Figure 1. Metacrangon australis sp. nov., holotype, female (cl 13.1
mm), NMV J60424, entire animal in lateral view (left fifth pereopod
lost). Scale bar = 5 mm.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

47

Figure 2. Metacrangon australis sp. nov., holotype, female (cl 13.1 mm), NMV J60424: a, carapace and cephalic appendages, dorsal view (setae
partially omitted); b, anterior part of carapace (left side) and left cephalic appendages, dorsal view; c, abdomen, dorsal view; d, posterior part of
telson, dorsal view. Scale bars = 5 mm for a and c; 2 mm for b; 1 mm for d.

48

T. Komai &J. Taylor

Figure 3. Metacrangon australis sp. nov., holotype, female (cl 13.1 mm), NMV J60424: a, distal two segments of left third maxilliped, dorsal
(extensor) view (setae omitted); b, distal part of antepenultimate segment of left third maxilliped, ventral view; c, left first pereopod, lateral view;
d, same, subchela and carpus, dorsal (extensor) view; e, left second pereopod, lateral view; f, right third pereopod, lateral view; g, left fourth
pereopod, lateral view; h, same, dactylus, flexor view; i, right fifth pereopod, lateral view; j, same, dactylus, flexor view; k, posterolateral tooth of
left uropodal exopod, dorsal view (setae omitted). Scale bars = 2 mm for a-g and i; 1 mm for h and j; 0.5 mm for k.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

49

Antennular peduncle (figs 2a-b) moderately stout, reaching
midlength of antennal scale. First segment with prominent,
blunt distolateral process directed dorsally; distomesial
margin unarmed; stylocerite falling far short of distolateral
process of first segment, terminating in slender, sharp tooth,
lateral margin gently convex. Second segment widened
distally, slightly longer than wide, with prominent, blunt
distolateral process. Third segment much wider than long.
Outer flagellum consisting of 13 articles.

Antennal basicerite (figs 2a-b) stout, with acutely pointed
dorsodistal lateral angle and long ventrolateral tooth distinctly
overreaching dorsodistal lateral angle. Antennal scale (figs
2a-b) 0.50-0.55 times as long as carapace and about 2.3 times
longer than wide; lateral margin weakly concave; distolateral
tooth moderately broad, slightly overreaching rounded lamella.

Third maxilliped (fig. 3a) relatively stout, overreaching
antennal scale by full length of ultimate segment; ultimate
segment about 4.5 times longer than wide; penultimate segment
about 2.3 times longer than wide; antepenultimate segment
with two subequal spiniform setae subdistally (fig. 3b).

First pereopod (fig. 3 c) moderately stout, slightly
overreaching antennal scale; palm (fig. 3d) 2.9 times longer
than wide, not widened proximally or distally; lateral and
mesial margins faintly sinuous; thumb relatively long; carpus
with small ventrolateral tooth, otherwise unarmed on
distolateral margin; merus with small dorsodistal tooth,
ventral margin sinuous, crested. Second pereopod with
dactylus about 0.3 times as long as palm (fig. 3e); length ratio
of chela to ischium 1:23:2.2:1.8. Third pereopod (fig. 3f)
slender; length ratio of dactylus to ischium 1:1.83.8:2.8:2.8.
Fourth pereopod (fig. 3g) relatively stout, reaching nearly
distal margin of antennal scale; dactylus (fig. 3H) elongate
subovate, spatulate, about 0.5 times as long as propodus,
margins naked; dactylus-propodus articulation about 60°;
propodus about 4.0 times longer than wide; propodus-carpus
combined distinctly shorter than merus-ischium combined.
Fifth pereopod (fig. 3i) shorter than fourth pereopod; dactylus
(fig. 3j) spatulate, subequal in length to dactylus of fourth
pereopod, about 0.7 times as long as propodus.

Uropodal exopod with blunt posterolateral tooth and with
three spiniform setae (fig. 3k).

Colouration. Not known.

Distribution. Known only from the type locality in the
Southern Ocean southeast of Tasmania, west of Macquarie
Island, at depths of 700-900 m.

Remarks. Metacrangon australis is somewhat similar to M.
proxima, M. variabilis and M. poorei sp. nov. in the general
disposition of carapacial teeth and the carination of the
abdomen, but it is quite unique within the genus in having the
combination of the following characters: the rostrum is
narrowly triangular and distinctly overreaches the distal
corneal margins; the carapace has two middorsal teeth, of them
the anterior tooth is distinctly postrostral and the posterior
(cardiac) tooth arises at 0.75-0.80 of the carapace length; the
lateral margin of the rostrum merges into the orbital margin,
thus no cleft is defined; the first to fifth abdominal somites bear

sharp, crested middorsal carina; and the anterior three pleura
are rounded marginally. Of particular note is the lack of an
orbital cleft, an uncommon trait previously known only in M.
knoxi (Yaldwyn, 1960) (see Komai, 1997). M. knoxi is referred
to the M. jacqueti (A. Milne -Edwards, 1881) species group
(Komai, 1997) and is readily distinguished from M. australis
by the anterior middorsal tooth on the carapace that arises at
the midlength of the rostrum and the presence of a ventral tooth
on each first to third abdominal pleuron.

Etymology. Named ‘ australis ’, Latin meaning ‘southern’,
alluding to the type locality of this new species, representing
the southernmost locality of the genus.

Metacrangon poorei sp. nov.

Figures 4, 5

Material examined. Holotype: Australia, off southeastern Victoria,
(39°53.76'S, 149°03.39'E), 1608 m, 28 Apr 2000, FRV Southern
Surveyor, epibenthic sled, (stn SS01/00/246), NMV J52069 (1 female,
cl 8.1 mm).

Description. Based on holotype female. Body (figs 4a-e)
moderately robust. Rostrum (figs 4a-b, 5a) triangular with
acute apex in dorsal view, strongly ascending (angle against
horizontal plane of carapace about 45°), 0.20 times as long as
carapace; dorsal surface with low, but clearly delimited
middorsal carina; lateral margin faintly sinuous in lateral view,
merging into postorbital region of carapace; midventral carina
distinct, ventral margin sinuous in lateral view. Carapace (figs
4a-b) not widened posteriorly, longer than wide postorbitally;
dorsal midline with two moderately small teeth; anterior
(epigastric) tooth arising at 0.18 of carapace length, posterior
(cardiac) tooth broken off, arising at 0.68 of carapace length;
submedian and hepatic teeth moderately small; antennal tooth
moderately strong, directed forward in dorsal view, strongly
ascending (same degree as rostrum) in lateral view, acuminate,
falling slightly short of rostral apex; orbital cleft distinct;
anterolateral margin between antennal and branchiostegal
teeth concave, unarmed; branchiostegal tooth moderately
strong, directed forward in dorsal view, strongly ascending in
lateral view, reaching dorsodistal margin of antennal basicerite;
pterygostomial tooth small, clearly visible in lateral view;
postorbital carina clearly delimited, accompanied by
longitudinal suture; epibranchial carina weakly absent.

Fifth to eighth thoracic sternites each with distinct median
keel: on fifth, spiniform, directed forward; on sixth,

terminating anteriorly in tiny tooth; on seventh, angulated
anteriorly; and on eighth, rounded.

Abdomen (figs 4c-e) slightly sculptured, surface sparsely
punctate; first somite with trace of middorsal carina, second to
fourth somites with rather broad, clearly delimited middorsal
carina. Pleuron of anterior four somites rounded marginally.
Fifth somite with low, rather broad, clearly delimited middorsal
carina; posterodorsal margin faintly produced medially;
posterolateral margin unarmed; pleuron with posteroventral
angle rounded, ventral margin gently convex. Sixth somite 1.7
times longer than wide, with distinct, slightly curved
submedian carinae, not reaching posterodorsal margin;

50

T. Komai &J. Taylor

Figure 4. Metacrangon poorei sp. nov., holotype, female (cl 8.1 mm), NMV J52069: a, carapace and cephalic appendages, dorsal view; b, same,
lateral view; c, first to third abdominal somites, dorsal view; d, fourth abdominal somite to telson, dorsal view; e, entire abdomen, lateral view.
Scale bar = 2 mm.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

51

Figure 5. Metacrangon poorei sp. nov., holotype, female (cl 8.1 mm), NMV J52069: a, anterior part of carapace (left side) and left cephalic
appendages, dorsal view; b, distal two segments and distal part of antepenultimate segment of left third maxilliped, ventral (flexor) view; c, left
first pereopod, lateral view; d, same, subchela, flexor view; e, left second pereopod, lateral view; f, left third pereopod, lateral view; g, left fourth
pereopod, lateral view; h, same, dactylus, flexor view; i, right fifth pereopod, lateral view; j, posterolateral tooth of left uropodal exopod, dorsal
view. Scale bars = 1 mm for a-g and i; 0.5 mm for h and j.

52

T. Komai &J. Taylor

dorsolateral carina distinct, reaching to posterodorsal margin;
posterodorsal margin produced, weakly bilobed; pleuron
flared laterally, posteroventral tooth small; posterolateral
process moderately strong, directed slightly laterally,
terminating in sharp tooth. Telson (fig. 4d) damaged, but one
pair of dorsolateral spines still preserved.

Eye (figs 4a, 5a) slightly longer than wide; cornea as wide
as proximal part of eyestalk, darkly pigmented, corneal width
0.17 of carapace length; eyestalk with small, papilladike
dorsal tubercle.

Antennular peduncle (figs 4a, 5a) moderately stout. First
segment with prominent, blunt distolateral process directed
dorsally; distomesial margin unarmed; stylocerite just
reaching tip of distolateral process, subacutely pointed, lateral
margin gently convex. Second segment slightly widened
distally, distinctly longer than wide, with prominent, blunt
distolateral process. Third segment wider than long. Outer
flagellum consisting of 10-11 articles.

Antennal basicerite (fig. 5a) stout, with rounded dorsodistal
lateral angle and long ventrolateral tooth distinctly
overreaching dorsodistal lateral angle. Antennal scale
damaged (fig. 5a).

Third maxilliped (fig. 5b) relatively slender, overreaching
antennal scale by 0.6 length of ultimate segment; ultimate segment
gradually tapering distally, 6.0 times longerthan wide; penultimate
segment about 2.7 times longer than wide; antepenultimate
segment with two unequal spiniform setae subdistally.

First pereopod (figs 5c-d) moderately stout; palm 3.5
times longer than wide, not widened proximally or distally;
lateral and mesial margins faintly sinuous; thumb moderately
broad; carpus with small ventrolateral tooth, otherwise
unarmed on lateral margin; merus with small dorsodistal
tooth, ventral margin sinuous, crested. Second pereopod (fig.
5e) with dactylus about 0.5 times as long as palm; length ratio
of chela to ischium 1:1. 7: 1.6: 1.5. Third pereopod (fig. 5f)
slender; length ratio of dactylus to ischium 1:1. 9:3. 4:2. 5:2. 6.
Fourth pereopod (fig. 5g) moderately stout; dactylus (fig. 5h)
narrowly spatulate, about 0.6 times as long as propodus,
margins naked; dactylus-propodus articulation about 30°;
propodus about 5.0 times longer than wide; carpus shorter
than propodus, with numerous long setae on dorsal margin;
row of long setae on dorsal and ventral margins of merus and
ischium (dorsal setae longer than ventral setae). Fifth pereopod
(fig. 5i) slightly shorter than fourth pereopod; dactylus slender,
subspatulate, longer than dactylus of fourth pereopod, about
0.8 times as long as propodus.

Uropodal exopod with minute posterolateral tooth and minute
spinule just located mesial to posterolateral tooth (Fig. 5j).

Colouration. Not known.

Distribution. Known only from the type locality off
southeastern Victoria, at a depth of 1608 m.

Remarks. Metacrangon poorei sp. nov. is morphologically very
similar to M. variabilis (Rathbun, 1902) from the northeastern
Pacific and M. proximo Kim, 2005 from Japan in the disposition
of teeth on the carapace and the development of the middorsal
carina of the abdomen. Nevertheless, the new species is easily

distinguished from the latter two species by the following
characters: the rostrum is acutely pointed and reaches to the
distal corneal margins in M. poorei , whereas it is blunt or
subacute at the tip and falls far short of the distal corneal
margins in the latter two species; the anterolateral angle of the
postorbital carina is rounded in M. poorei, rather than bearing
a small triangular tooth or denticle in the latter two species; the
lateral carina on the fifth abdominal somite is obsolete in M.
poorei sp. nov, but it is distinct in the latter two species; and the
stylocerite of the antennule reaches the distolateral process on
the first peduncular segment in the new species, rather than
falling far short of it in the latter two species.

Etymology. It is our pleasure to dedicate this new species to our
esteemed colleague, Dr. Gary C. B. Poore.

Metacrangon spinidorsalis sp. nov.

Metacrangon sp.(Poore, 2004, 139, fig. 36f)

Metacrangon sp. MoV 5423. (Poore et al., 2008, 81)

Figures 6-10

Material examined. Holotype: Australia, Western Australia, off Point
Hillier (35°22.54'S, 117°12.25'E-35°22.54'S, 117°12.25'E), 539 m, 22
Nov 2005, FRV Southern Surveyor, beam trawl (stn SS10/2005/019),
WAM C45115 (1 female, cl 10.4 mm).

Paratypes: Tasmania, Tasman Sea off Maria Island (42°42.8'S,
148°22.2'E), 450 m, 25 Jun 1984, RV Soela, demersal beam trawl (stn
S03/84/77), NMV J40886 (1 female, cl 7.6 mm); Southern Ocean, 48
km west of Richardson Point (4P15.0'S, 144°08.0'E), 520 m, 20 Oct
1984, Frank and Bryce demersal trawl (stn S05/84/51), NMV J40954
(1 female, cl 9.0 mm). Western Australia, off Point Hillier (35°22.54'S,
117°12.25'E - 35°22.54'S, 117°12.25'E), 539 m, 22 Nov 2005, FRV
Southern Surveyor, beam trawl (stn SS10/2005/019), NMV J54497 (3
females, cl 5.4-10.4 mm, 6 males, cl 6. 1-7.0 mm); off Bald Island
(35°04.01'S, 118°39.50'E - 35°13.40'S, 118°40.30'E), 728-710 m, 23
Nov 2005 (stn SS10/2005/032), NMV J19215 (1 female, cl 7.7 mm);
(35°12.49'S, 118°39.04'E - 35°12.14'S, 118°40.08'E), 431-408 m, 24
Nov 2005 (stn SS10/2005/034), WAM C45116 (1 female, cl 6.9 mm);
off Perth Canyon (31°59.33'N, 115 o 10.59'E-32°00.07'S, 115°10.4rE),
508-478 m, 29 Nov 2005 (stn SS10/2005/068), NMV J54512 (4
females cl 67-9.5 mm, 1 male cl 6.5 mm).

Description. Female. Body (fig. 6) moderately robust. Rostrum
(figs 7a-b) narrowly triangular with acute apex in dorsal view,
directed forward, 0.20-0.25 times as long as carapace; dorsal
surface nearly flat; lateral margin slightly arched in lateral
view, merging into postorbital region of carapace; midventral
carina low, ventral margin slightly sinuous in lateral view.
Carapace (figs 6, 7a) slightly widened posteriorly, slightly
longer than wide postorbitally; surface covered with very short
setae; dorsal midline with two moderately small teeth; anterior
tooth arising at rostral base, not overlapping rostrum, slightly
larger than posterior (cardiac) tooth; posterior (cardiac) tooth
arising at 0.55-0.60 of carapace length; submedian teeth
moderately small; hepatic tooth relatively small; antennal tooth
moderately strong, directed forward in dorsal view, somewhat
ascending in lateral view (angle about 30° against horizontal
plane of carapace), acuminate, far falling short of rostral apex;
orbital cleft present, but only weakly delimited; anterolateral
margin between antennal and branchiostegal tooth concave,

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

53

with tiny spinule inferior to base of antennal tooth;
branchiostegal tooth moderately strong, directed forward in
dorsal view and somewhat dorsally in lateral view, reaching
dorsodistal margin of antennal basicerite; pterygostomial tooth
small, not visible in lateral view; postorbital carina clearly
delimited, accompanied by longitudinal suture; epibranchial
carina weakly delimited.

In spawning molts, thoracic sternum concave, armature
absent; only fifth sternite with small tubercles medially. In
nonspawning molts, fifth sternite with sharp, procurved tooth;
sixth to seventh somites each with rounded, strongly
compressed prominence, becoming higher posteriorly.

Abdomen (figs 6, 7c) slightly sculptured; anterior two
somites without trace of middorsal carina anteriorly, but second
tergum with distinct spiniform middorsal tooth located at
anterior end of posterior section; third somite with trace of
middorsal carina, and fourth somite with broad, clearly delimited
middorsal carina. First and second pleura each with blunt tooth
on ventral margin; third pleuron with blunt tooth at anteroventral
angle; fourth pleuron unarmed. Fifth somite with low, but
distinct middorsal carina; posterodorsal margin faintly produced
medially; posterolateral margin unarmed; pleuron with
posteroventral angle subacutely pointed, ventral margin gently
convex. Sixth somite with distinct, straight submedian carinae,
not reaching posterodorsal margin; dorsolateral carina distinct,
reaching to posterodorsal margin; posterodorsal margin
produced, faintly bilobed; pleuron flared laterally, posteroventral
tooth small, acute or subacute; posterolateral process strong,
directed slightly laterally, terminating in sharp tooth. Telson (fig.
7c) longer than sixth somite, tapering to acute tip; dorsal surface
deeply grooved mesially, with three pairs of lateral spines,
anterior-most pair located at about midlength; two plumose
setae posterior to third pair of lateral spines (fig. 7d).

In spawning molt, first to fourth abdominal sternites
unarmed; fifth sternite with low median tubercle. Sixth
abdominal sternite shallow depressed medially.

Eye (figs 7a-b) slightly longer than wide; cornea slightly
wider than eyestalk, light brown or opaque in preservative,
corneal width 0.14-0.15 of carapace length; eyestalk with
small, papilla-like dorsal tubercle.

Antennular peduncle (figs 7a-b) moderately stout,
overreaching midlength of antennal scale. First segment with
prominent, blunt distolateral process directed dorsally;
distomesial margin unarmed; stylocerite falling slightly short
of distolateral process, acutely or subacutely pointed, lateral
margin gently convex. Second segment slightly widened
distally, distinctly longer than wide, with prominent, blunt
distolateral process. Third segment wider than long. Outer
flagellum overreaching distal margin of lamella of antennal
scale by about 0.4 length, consisting of 10-11 articles.

Antennal basicerite (fig. 7e) stout, with sharply pointed
dorsodistal lateral angle and short ventrolateral tooth slightly
overreaching dorsodistal lateral angle; carpocerite
subcylindrical, reaching distal 0.20 of antennal scale. Antennal
scale (figs 7a, 7e) about 0.50 times as long as carapace and 2.3
times longer than wide; lateral margin faintly sinuous;
distolateral tooth relatively wide, slightly falling short of
rounded distal margin of lamella.

Figure 6. Metacrangon spinidorsalis sp. nov., holotype, female (cl
1 0 .4 mm), WAM C45 115, entire animal in lateral view. Scale bar = 5 mm.

54

T. Komai &J. Taylor

Figure 7. Metacrangon spinidorsalis sp. nov., holotype, female (cl 10.4 mm), WAM C45115: a, carapace and cephalic appendages, dorsal view;
b, anterior part of carapace (left side), left eye and left antennule, dorsal view (setae omitted); c, abdomen, dorsal view; d, distal part of telson,
dorsal view; e, left antennal scale, dorsal view. Scale bars = 5 mm for a and c; 2 mm for b and e; 1 mm for d.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia 55

Figure 8. Metacrangon spinidorsalis sp. nov., holotype, female (cl 10.4 mm), WAM C45115; left thoracic appendages: a, third maxilliped, dorsal
view (setae omitted); b, distal part of antepenultimate segment of third maxilliped, ventral view; c, first pereopod, lateral view; d, same, subchela,
dorsal (extensor) view; e, second pereopod, lateral view (coxa damaged); f, same, chela, extensor view; g, third pereopod, lateral view; h, fourth
pereopod, lateral view; i, same, dactylus, flexor view; j, fifth pereopod, lateral view; k, same, dactylus, flexor view; 1, posterolateral tooth of right
uropodal exopod, dorsal view (setae omitted). Scale bars = 2 mm for a-h and j; 1 mm for i and k; 0.5 mm for 1.

56

T. Komai &J. Taylor

Figure 9. Metacrangon spinidorsalis sp. nov., paratype, male (cl 6.5 mm), NMV J54512: a, carapace and cephalic appendages, dorsal view; b,
same, lateral view; c, thoracic teeth, ventrolateral view; d, abdomen, lateral view; e, left fourth pereopod, lateral view; f, same, dactylus, flexor
view; g, endopod of left first pleopod, ventral view; h, endopod and appendix masculina of left second pleopod, mesial view. Scale bars = 2 mm
for a, b and d; 1 mm for c and e; 0.5 mm for f-h.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

57

Figure 10. Metacrangon spinidorsalis sp. nov., paratype, male (cl 6.2 mm), NMV J54497, lateral view taken of live specimen on board FRV
Southern Surveyor. Scale bar = 10 mm.

Third maxilliped (fig. 8a) relatively slender, overreaching
antennal scale by 0.6 length of ultimate segment; margins and
dorsal surface of distal two segments with numerous short to
long setae. Ultimate segment gradually tapering distally, 6. 5-7.0
times longer than wide. Penultimate segment about 2.6 times
longer than wide. Antepenultimate segment sinuously curved in
dorsal view, with thick tuft of long setae dorsodistally; ventral
surface with two minute spiniform setae subdistally (fig. 8b).
Coxa with rounded lateral process (not figured). Exopod falling
far short of distal margin of antepenultimate segment, with well-
developed flagellum.

First pereopod (figs 8c-d) moderately stout, reaching distal
margin of antennal scale; palm 3 .0-3 .5 times longer than wide,
not widened proximally or distally, cutting edge oblique; lateral
and mesial margins nearly straight; thumb moderately broad;
carpus with small ventrolateral tooth, otherwise unarmed on
lateral margin; merus with small dorsodistal tooth, ventral margin
sinuous, crested. Second pereopod (fig. 8e) reaching nearly to
midlength of antennal scale; dactylus about 0.5 times as long as
palm; cutting edges of fingers with row of minute spiniform setae
(fig. 8f); length ratio of chela to ischium 1: 1 .9: 1 .6: 1 .5; coxa with
prominent flap-like process (not figured). Third pereopod (fig.
8g) slender, nearly reaching distal margin of antennal scale by tip
of dactylus; length ratio of dactylus to ischium 1:2.2:4.6:2.9:3.2.
Fourth pereopod (fig. 8h) moderately stout, slightly overreaching
midlength of antennal scale by dactylus; dactylus (fig. 8i)

spatulate, about 0.8 times as long as propodus, upper margin with
row of dense stiff setae, lower margin with few sparse setae; tip
of dactylus terminating in two unequal projections flanking
unguis, upper projection longer, with minute setae; dactylus-
propodus articulation about 45°; propodus about 3 .6 times longer
than wide, bearing row of dense stiff setae on dorsal and ventral
margins; carpus shorter than propodus, with numerous long setae
on dorsal margin; row of long setae on dorsal and ventral margins
of merus and ischium (dorsal setae longer than ventral setae).
Fifth pereopod (fig. 8j) distinctly shorter than fourth pereopod,
falling far short of base of branchiostegal tooth; dactylus (fig. 8k)
subspatulate, shorter than dactylus of fourth pereopod, about 0.6
times as long as propodus; setation much less than in fourth
pereopod.

Uropod (fig. 6) not reaching tip of telson; exopod with nearly
straight lateral margin, posterolateral angle terminating in
truncate tooth (fig . 81) ; no movable spinule mesial to posterolateral
tooth; endopod longer and narrower than exopod.

Male. Rostrum nearly spiniform (fig. 9A), 0.25-0.35 times as
long as carapace, slightly overreaching distal comeal margins.
Carapace (figs 9a-b) with two prominent middorsal teeth,
anterior tooth elongate, arising slightly anterior to or just at level
of posterior margin of orbit, overlapping rostrum; posterior tooth
strong, hooked; branchiostegal tooth strong, curved slightly
laterally, distinctly overreaching distolateral angle of antennal
basicerite. Fifth to eighth thoracic stemites (fig. 9c) with

58

T. Komai &J. Taylor

prominent, acute teeth becoming larger posteriorly.

Middorsal tooth on second abdominal somite larger than in
female; pleural ventral teeth on anterior three somites more
pronounced in females (fig. 9d).

Comeal width 0.18-0.20 of carapace length (fig. 9a). Outer
antennular flagellum (fig. 9a-b) overreaching antennal scale by

0.8 length, consisting of 15-18 articles. Antennal scale (fig. 9a)
0.55-0.60 times as long as carapace; distolateral tooth distinctly
overreaching distal lamella.

Fourth pereopod (fig. 9e) more slender than in females
(propodi about 4.5 times longer than wide); dactylus (fig. 9f)
narrowly spatulate.

Endopod of first pleopod (fig. 9g) tapering distally to rounded
tip, bearing four stiff setae terminally. Second pleopod with
appendix masculina reaching about distal 0.6 of endopod, bearing
about 10 long spiniform setae (fig. 9h).

Colouration. Carapace, abdominal somites, pereopods, telson
and uropods relatively uniform browny red colour. Body, legs
and first pereopods covered with whitish pigment spots; similar
spots also present, but less pronounced on generally paler second
to fifth pereopods.

Distribution. The present material contains specimens from two
rather distant localities, namely southwestern Australia and
Tasmania, suggesting that this species is widely distributed in
southern Australia, at depths of 408-728 m.

Remarks. As is apparent from the above description, Metacrangon
spinidorsalis sp. nov. shows considerable degree of sexual
dimorphism in the development of the middorsal teeth on the
carapace and the shape of the antennal scale.

This new species is referable to the Metacrangon jacqueti
species group because of the disposition of teeth on the carapace,
the presence of ventral tooth on each first to third abdominal
pleuron, and the setose margins of the dactyli of the fourth and
fifth pereopods (Komai, 1997). The following nine species are
referred to this informal species group (Komai, 1997; Retamal
and Gomy, 2003): M. agassizi (Smith, 1882) from the
northwestern Atlantic; M. bahamondei Retamal and Gomy, 2003
from southern part of Chile; M. bellmarleyi (Stebbing, 1914)
from western to southern Africa; M. crosnieri Komai, 1997 from
Madagascar; M. jacqueti from the northeastern Atlantic; M.
knoxi from the Chatham Rise, New Zealand; M. ochotensis
(Kobjakova, 1955) from the South Kuril Islands in the
northwestern Pacific; M.procax (Faxon, 1893) from California
to Peru in the eastern Pacific; and M. similis Komai, 1997 from
Japan. However, M. spinidorsalis is unique even within the
genus, as it possesses a distinct middorsal tooth on the second
abdominal tergite. Furthermore, in this new species, the orbital
cleft is only weakly delimited. In this regard, this new species is
intermediate between M. knoxi (where the orbital cleft is absent)
and other species in the M. jacqueti species group (where the
orbital cleft is distinct). The small anterior middorsal tooth on the
carapace in females, which does not overlap the rostrum, also
distinguishes M. spinidorsalis from other species in the M.
jacqueti species group.

Etymology. Named in reference to the characteristic spiniform
tooth on the second abdominal tergite.

Concluding remarks

This study reports the significant findings of three new species of
Metacrangon from rather high latitudinal areas in the southern
oceans of Australia, increasing the number of species known
from the southern hemisphere from five to eight. The previously
described species known from the southern hemisphere are M.
bahamondei, M. bellmarleyi, M. crosnieri, M. knoxi and M.
richardsoni (Yaldwyn, 1960) (cf. Yaldwyn, 1960; Komai, 1997;
Retamal & Gomy, 2003). With the exception of M. richardsoni,
all of them are referred to the M. jacqueti species group. Although
Metacrangon has been well represented by the north Pacific
species, it is suggested that the diversity of the genus in the
southern hemisphere is much higher than we expected. Other
genera of the family have also been shown to be species rich in
southern Australian waters with new species and range extensions
recently reported for Lissosabinea Christoff ersen, 1988 and
Philocheras Stebbing, 1900 (Komai, 2008; Taylor and Collins,
2009; Taylor, 2010).

Key to Australian species of Metacrangon

1. Carapace with anterior middorsal tooth arising at level of
rostral base; second abdominal tergite with middorsal

tooth; dactylus of fourth pereopod with marginal setae

M. spinidorsalis sp. nov.

- Carapace with anterior middorsal tooth arising distinctly
posterior to rostral base; second abdominal tergite
unarmed; dactylus of fourth pereopod naked marginally ..
2

2. Rostrum overreaching distal corneal margins; orbital
cleft absent; middorsal carina on first to fifth abdominal
somites crested; antennular stylocerite falling short of

distolateral process of first peduncular segment

M. australis sp. nov.

- Rostrum just reaching distal corneal margins; orbital cleft
present; middorsal carina on first to fifth abdominal
somites not crested; antennular stylocerite reaching

distolateral process of first peduncular segment

M.poorei sp. nov.

Acknowledgments

The preliminary identifications of the material lodged at
Museum Victoria were made by Gary Poore.

Thanks to Alan Williams and Rudy Kloser from CSIRO
Marine and Atmospheric Research (CMAR) who were largely
responsible for the sampling design of the ‘Voyages of
Discovery’ research program, which generated the Western
Australian proportion of the material listed in this report.
Collection of that material has been funded through the
Commonwealth Environment Research Facilities (CERF)
program, an Australian Government initiative supporting
world-class, public research. Special thanks to Karen Gowlett-
Holmes for permission to use the photograph of Metacrangon
spinidorsalis sp. nov. that is published here.

Three new species of the crangonid genus Metacrangon Zarenkov (Crustacea: Decapoda: Caridea) from Australia

59

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Memoirs of Museum Victoria 67: 61-95 (2010)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers
(Echinodermata: Holothuroidea: Apodida)

P. Mark O’Loughlin 1 and Didier VandenSpiegel 2

1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (pmo@bigpond.net.au)

2 Musee royal de l’Afrique centrale. Section invertebres non-insects, B-3080, Tervuren, Belgium (dvdspiegel@
africamuseum.be)

Abstract O’Loughlin, P.M. and VandenSpiegel, D. 2010. A revision of Antarctic and some Indo-Pacific apodid sea cucumbers

(Echinodermata: Holothuroidea: Apodida). Memoirs of Museum Victoria 67: 61-95.

Eight new apodid species from Antarctica are described: myriotrochids Achiridota smirnovi sp. nov., Myriotrochus
nikiae sp. nov., Prototrochus linseae sp. nov., Prototrochus barnesi sp. nov., and chiridotids Kolostoneura griffithsi sp.
nov., Scoliorhapis bipearli sp. nov., Scoliorhapis massini sp. nov., Taeniogyrus prydzi sp. nov. Genera Scoliorhapis H. L.
Clark, Taeniogyrus Semper and Trochodota Ludwig are reviewed. Scoliodotella Oguro is ajunior synonym of Scoliorhapis
H. L. Clark. Trochodota Ludwig type species is fixed as Holothuria (Fistularia) purpurea Lesson. Trochodota Ludwig is
a junior synonym of Taeniogyrus Semper. Sigmodota Studer type species is fixed as Chiridota contorta Ludwig, and
Sigmodota Studer is raised out of synonymy with Taeniogyrus Semper. Species assigned to Sigmodota are Chiridota
contorta Ludwig, Taeniogyrus dubius H. L. Clark (as Sigmodota dubia) and Taeniogyrus magnibaculus Massin and
Heterier (as Sigmodota magnibacula). Non-Antarctic new genus Rowedota gen. nov. is erected with type species
Taeniogyrus allani Joshua, and other assigned species Trochodota epiphylca O’Loughlin, Trochodota mira Cherbonnier,
Trochodota shepherdi Rowe and Trochodota vivipara Cherbonnier. Trochodota species not assigned to Rowedota gen.
nov. and Sigmodota Studer are assigned to Taeniogyrus Semper. Other Antarctic apodid species discussed are Myriotrochus
antarcticus Smirnov and Bardsley, Myriotrochus hesperides O’Loughlin and Manjon-Cabeza and Taeniogyrus antarcticus
Heding. Non-Antarctic apodid species discussed are Chiridota pisanii Ludwig, Chiridota australiana Stimpson and
Trochodota maculata H. L. Clark. The spelling of the species name Myriotrochus macquariensis Belyaev and Mironov is
corrected. A table with Antarctic Apodida species and their distributions is provided. A table with specimen and ossicle
sizes for some Taeniogyrinae species is provided. A key to genera of Taeniogyrinae is provided. Species names are
standardized to: macquariensis ; studerv, theeli.

Keywords Achiridota, Chiridota, Kolostoneura, Myriotrochus, Paradota, Prototrochus, Scoliorhapis, Scoliodotella, Sigmodota,

Taeniogyrus, Trochodota, emended diagnoses, new genus, new species, Antarctic, Bellingshausen Sea, Indo-Pacific,
Prydz Bay, Ross Sea, Scotia Sea, Weddell Sea.

Introduction

This review of Antarctic apodid genera and species has become
possible with the opportunities to study numbers of Antarctic
holothuroid collections (Table 1).

ANARE 1993 collected with a Van Veen grab and epibenthic
sled, and recent BIOPEARL expeditions used an epibenthic sled,
and both yielded many very small specimens including the
apodids that are reported in this study. Tissue samples from
recent NIWA and US AMLR and BAS collections are currently
being processed for molecular sequences, but this systematic
study is based exclusively on morphological characters and the
species recognized as morpho-species .

Numerous systematic problems have arisen during the study
of species of Taeniogyrinae Smirnov, 1998. There has been a

significant history of misidentifications of Antarctic and
Magellanic species. A succession of authors has been dissatisfied
with the systematic status of one or both of the genera Taeniogyrus
Semper, 1867 and Trochodota Ludwig, 1891, including Dendy
(1909), Joshua (1914), H. L. Clark (1921), Rowe (1976), Rowe
(in Rowe and Gates 1995), Smirnov (1997), Massin and Heterier
(2004) and O’Loughlin and VandenSpiegel (2007). The presence
or absence of clusters of wheels in the body wall is the generic
diagnostic distinction between Taeniogyrus and Trochodota
species, and has proved to be subjective and unsatisfactory. And
a useful generic diagnostic character for taeniogyrinid species,
namely the arrangement of teeth on the inner rim of the wheel
ossicles, became potentially lost because of a misunderstanding
in a revision of a type species . We attempt to improve systematic
clarity around these issues.

62

P.M. O’Loughlin & D. VandenSpiegel

Table 1. Antarctic collections studied.

Names of expeditions

Localities

Specimens lodged

Terra Nova 1910-1913

Ross Sea

Natural History Museum (London)

Discovery Expedition

South Atlantic, Scotia Sea

Natural History Museum (London)

BANZARE

Eastern Antarctica,
Kerguelen Islands

South Australian Museum

US Antarctic Research Program

Antarctic Ocean

US National Museum of Natural
History (Smithsonian Institution)

ANARE

Prydz Bay, Heard Island

Museum Victoria, Tasmanian
Museum, South Australian Museum

Tangaroa Ross Sea

Ross Sea

New Zealand Institute of Water and
Atmospheric Research

Hesperides BENTART-2003,
BENTART-2006

Amundsen Sea,
Bellingshausen Sea

University of Malaga

British Antarctic Survey BIOPEARL 2006,
BIOPEARL 2008

Bellingshausen Sea,
Scotia Sea

Natural History Museum (London),
Museum Victoria

US Antarctic Marine Living Resources
2004, 2005, 2009

South Atlantic, Scotia Sea

Museum Victoria

US Antarctic Marine Living Resources
2006

Antarctic Peninsula

US National Museum of Natural
History (Smithsonian Institution)

Methods

Photographs of preserved specimens (Figure 1) were taken
using a Pentax K-7 camera with Olympus 38 mm macro lens
on bellows. Photographs were taken at f 1 1 — f 1 6 using twin
flashes. The photograph of a preserved specimen of
Myriotrochus antarcticus was taken using a Leica MZ12.5
compound microscope, Q imaging camera, and Auto-Montage
software. The photograph of a preserved specimen of
Taeniogyrus australianus was taken using a SLR Canon
EOS5D digital camera with 65 mm lens. For scanning electron
microscope (SEM) observations ossicles were cleared of
associated soft tissues in commercial bleach, air-dried, mounted
on aluminium stubs, and coated with gold. Observations were
made using a JEOL JSM-6480LV SEM. Measurements were
made with Smile view software. Montage photographs of
ossicles were taken using a Leica CTR5000 compound
microscope, Leica DC500 digital camera, and Auto-Montage
software. Drawings were done by Mark O’Loughlin.

Corrected taxa spellings

Myriotrochus macquariensis Belyaev and Mironov, 1981 was
named for material collected near Macquarie Island, and the
original spelling of the species name as macquoriensis was a
lapsus calami. We correct the “incorrect original spelling” to
macquariensis in accord with Article 32.5 of the ICZN (1999).

Ludwig (1875) and some subsequent authors used the
generic name Chirodota instead of Chiridota Eschscholtz, 1829.

The correct spelling Chiridota is used throughout this paper.

Theel (1886a) erected the species Chiridota studerii. The
species name has various spellings in the literature and we use
the appropriate spelling studeri throughout this work.

Heding (1928) erected the species Scoliodota theelii. We
use the appropriate spelling theeli throughout this work.

Abbreviations

AM

AMLR

ANARE

BANZARE

BAS

BIOROSS-NIWA

ICZN

MNA

NHM

NIWA

NMNH

Australian Museum (echinoderm registration
numbers with prefix J) .

Antarctic Marine Living Resources .
Australian National Antarctic Research
Expedition.

British, Australian, New Zealand Antarctic
Research Expedition.

British Antarctic Survey.

Tangaroa 2004 expedition to the Ross Sea.
International Code of Zoological
Nomenclature (1999).

University of Genoa registration number
prefix for BIOROSS Tangaroa 2004 and
other Ross Sea holothuroid specimens.
British Museum of Natural History.

New Zealand Institute of Water and
Atmospheric Research.

National Museum of Natural History,
Smithsonian Institution.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

63

NMV Museum Victoria (echinoderm registration

numbers with prefix F) .

NZIPY-CAML New Zealand International Polar Year-
Census of Antarctic Marine Life Project
cruise TAN0802.

RBINS Royal Belgian Institute of Natural Sciences.

All SEM material observed during this study
is deposited in RBINS.

SAM South Australia Museum.

TMAG Tasmania Museum and Art Gallery.

USARP United States Antarctic Research Progam.

USNM United States National Museum. Historically

three types of registration have been used for
USNM specimens: Echinoderm catalogue
numbers prior to 1920 did not have a prefix,
subsequently had the prefix E, and since 2001
the EMU on-line system has been used and
registrations reported as USNM without an
E prefix.

Numbers in brackets after registrations refer to numbers of
specimens in lots.

In this work Antarctic refers to the region south of the
Polar Front/Antarctic Convergence.

Relevant history of species misidentification

Three chiridotid species occur in the Magellanic/Falklands region
(north of the Polar Front/Antarctic Convergence): Chiridota
pisanii Ludwig, 1886 (12 tentacles, wheels in papillae in body
wall, lacking hooks); Chiridota contorta Ludwig, 1875 (12
tentacles, wheels in discrete clusters / papillae in the body wall,
and hooks); Holothuria (Fistularia) purpurea Lesson, 1830 (10
tentacles, wheels not in clusters in body wall, and hooks).
Taeniogyrus antarcticus Heding, 1931 (10 tentacles, wheels in
some groups, and hooks in body wall) has been found only south
of the Polar Front at South Georgia, Shag Rocks and the S Orkney
Is (see below), but not in the Falklands/Magellanic region.

Lampert (1885, 1886) discussed Chiridota purpurea

(Lesson), and provided complete synonymies. Subsequently
Lampert ( 1 889) recognized that the material that he had described
was Chiridota contorta Ludwig, and retained only Bell (1881) in
his synonymy for Chiridota purpurea (Lesson).

Studer (1876) erected a new genus Sigmodota because of the
presence of sigmoid hooks in an apodid species from both the
Kerguelen Is and Magellanic region. He reported 12 tentacles,
ignored the original description of 10 tentacles by Lesson (1830),
and referred Holothuria ( Fistularia ) purpurea Lesson, 1830 to
his Sigmodota. Theel (1886a, page 16) thought that it was “very
peculiar” that no Challenger specimen from the Kerguelen Is and
Strait of Magellan had 12 tentacles and hooks but no wheels. All
Challenger specimens had aggregations of wheels, and Theel
wondered if “the very scattered aggregations of wheels had
escaped the attention of Studer”. After more than another century
of collecting in these two regions still no specimen with 12
tentacles and hooks but lacking wheels has been found. We agree
with Theel’s concern (1886a, page 16) and judge that Studer did
not notice the presence of wheels in his material since the only
Chiridotidae species that has been found in the Kerguelen Is (see

O’Loughlin 2009) and also in the Magellanic region (this work)
is Chiridota contorta Ludwig (with 12 tentacles, hooks and
wheels). We agree with Ludwig’s (1898) judgment that Studer’s
Sigmodota purpurea (Lesson, 1830) is a junior synonym of
Chiridota contorta Ludwig.

Theel (1886a) judged that the material referred by Studer
(1876) to Sigmodota purpurea (Lesson) was not Holothuria
( Fistularia ) purpurea Lesson, presumably because of an absence
of wheels. He erected a new species Chiridota studeri Theel,
1886, understanding the species to have 12 tentacles, hooks, and
no wheels. Lampert (1889) retained Chiridota studeri Theel, but
described material with 10 tentacles, hooks and wheels not in
papillae. That material was Holothuria ( Fistularia ) purpurea
Lesson. For the reasons given in the paragraph above we again
agree with Ludwig’s (1898) judgment that Chiridota studeri
Theel, 1886 is a junior synonym of Chiridota contorta Ludwig.

In the same report Theel (1886a) described specimens from
the Falkland Is as having 12 tentacles, scattered aggregations of
wheels in the body wall, an absence of hooks, and minute rods in
the muscle bands. He judged that this material from the Falkland
Is was the true Holothuria ( Fistularia ) purpurea Lesson since it
came from the type locality. We again agree with Ludwig’s
(1898) judgment that Theel’s Holothuria ( Fistularia ) purpurea
Lesson is Chiridota pisanii Ludwig (described in the same year
1886).

Ludwig (1891) included the two species Chiridota studeri
Theel, 1886 and Chiridota venusta Semon, 1887 in his new genus
Trochodota Ludwig, 1891 . Ludwig (1898) subsequently changed
the identification of his included species Chiridota studeri Theel,
1886 to Trochodota purpurea (Lesson, 1830).

Ossicle clusters in generic diagnosis

Dendy and Hindle (1907) remarked that in regard to wheels
being grouped into papillae or scattered it was “undesirable to
recognize any generic distinction between these two forms”.
In establishing the new species Chiridota benhami Dendy,
1909 (a junior synonym of Chiridota dunedinensis Parker,
1881) Dendy remarked that it was “clearly impossible to base
generic distinctions merely upon the arrangement or even
upon the presence or absence of the wheels”. Joshua (1914)
found the degree to which wheels were aggregated in
Trochodota allani Joshua, 1912 varied greatly, and that
“grouped” and “scattered” applied to wheels in this one
species. H. L. Clark (1921) used the generic distinction in his
key of “wheels gathered into sharply defined papillae” or
“wheels scattered, often numerous enough to be crowded into
ill-defined heaps”. Rowe (1976) noted that for his species
Trochodota shepherdi wheels are grouped into more than “ill-
defined heaps” but not into papillae. He judged that the degree
to which wheels were grouped was not a useful generic
diagnostic character. Rowe also noted the anomaly that H. L.
Clark (1921) used grouping of hooks for species distinction
and grouping of wheels for generic distinction.

We have observed a range of wheel and hook arrangements
in the body wall (see O’Loughlin and VandenSpiegel 2007).
Wheels may be: clustered into discrete papillae, macroscopically
noticeable as white spots, as in Chiridota australiana Stimpson,

64

P.M. O’Loughlin & D. VandenSpiegel

1855, Chiridota contorta Ludwig, Taeniogyrus heterosigmus
Heding, 1931 and Taeniogyrus magnibaculus Massin and
Heterier, 2004; or clustered into longitudinal interradial bands
as in Trochodota shepherdi Rowe, 1976; or aligned in irregular
bands adjacent to the longitudinal muscles as in Trochodota
roebucki Joshua, 1914 and Taeniogyrus tantulus O’Loughlin,
2007 (in O’Loughlin and VandenSpiegel); or loosely grouped
into small clusters in larger specimens as in Taeniogyrus
antarcticus Heding.

Hooks may be: grouped closely into small papillae as in
Chiridota australiana Stimpson; aligned over and adjacent to
the longitudinal muscles as in Taeniogyrus papillis O’Loughlin,
2007 (in O’Loughlin and VandenSpiegel); aligned transversely
in paired series over the edges of the longitudinal muscles as in
Trochodota roebucki Joshua; scattered in all interradii as in
Taeniogyrus heterosigmus Heding.

We judge that the arrangement of wheels and hooks in the
body wall of chiridotid species may be useful as a species
diagnostic character but not useful at generic level. A
consequence is our synonymy below of Trochodota Ludwig,
1891 with Taeniogyrus Semper, 1867.

Wheel form in generic diagnosis

H. L. Clark (1921) thought that the teeth on the inner rim of the
wheels of Trochodota purpurea were in groups. This was
erroneous according to Smirnov (1997), and our observations.
A figure of a wheel of Chiridota purpurea (as Chiridota studeri
Theel) in Lampert 1889 (fig. 12a) does show a discontinuous
series of teeth that are only over the spokes. This does not occur
in any chiridotid wheels, and must be an error in illustration.

Rowe (1976) based his emended diagnoses of Taeniogyrus
and Trochodota on the diagnostic character of wheels with
continuous and those with discontinuous grouped teeth on the
inner rim. Rowe (1976) considered Trochodota purpurea to be
the type species for Trochodota, and followed H. L. Clark
(1921) in thinking that the teeth on the inner rim of the wheels
were discontinuous. Trochodota species were those with
discontinuous series of teeth. As noted by Smirnov (1997) the
Rowe review was unacceptable because the teeth are continuous
around the rim in the type species.

O’Loughlin and VandenSpiegel (2007) followed the
emended diagnosis of Trochodota by Rowe (1976). We now
reject that position. But we agree with Rowe (1976) that the
form of the wheels is useful at a generic diagnostic level, and
erect a new genus to accommodate taeniogyrinid species with
discontinuous series of teeth on the wheels.

Tentacle number in generic diagnosis

Tentacle number is not an inconsequential variable in apodid
genera and species since it is interdependent with the structure
of the calcareous ring. This is recognized in the diagnoses of
genera of Myriotrochidae where species with 12 tentacles and
two ring plates each with pairs of anterior projections are
assigned to genus Myriotrochus Steenstrup, 1851, and those
with 10 tentacles and single anterior projections on all 10 ring
plates are assigned to genus Prototrochus Belyaev and
Mironov, 1982. The Taeniogyrinae genera type species of

Taeniogyrus Semper and Trochodota Ludwig have 10 plates in
the calcareous ring and 10 tentacles, and the historical
diagnostic character distinguishing the species of these two
genera is the unsatisfactory degree to which wheels are
grouped in the body wall. We judge that these two genera are
synonyms, and raise the genus Sigmodota Studer, 1876, for
which the type has 12 calcareous ring plates and 12 tentacles,
out of synonymy to accommodate taeniogyrinid species with
wheels, hooks and 12 tentacles.

Key to genera of subfamily Taeniogyrinae

1 . Irregular thick spinous plates with wheel-spoked perforations

present in body wall

ArchedotaO’Loughlin (in O’Loughlin and VandenSpiegel)

— Lacking thick spinous plates in body wall 2

2. Lacking sigmoid hooks and chiridotid wheels in body

wall Kolostoneura Becher

— Sigmoid hooks with or without chiridotid wheels in body

wall 3

3. Sigmoid hooks only in body wall

Scoliorhapis H. L. Clark

— Sigmoid hooks and chiridotid wheels in body wall 4

4. Chiridotid wheels with discontinuous series of teeth

around inner rim of wheels Rowedota gen. nov.

— Chiridotid wheels with continuous series of teeth around

inner rim of wheels 5

5. Tentacles 12, and 12 plates in calcareous ring

Sigmodota Studer

— Tentacles 10, and 10 plates in calcareous ring . .

Taeniogyrus Semper

Order Apodida Brandt, 1835 (sensu Ostergren 1907)

Suborder Myriotrochina Smirnov, 1998

Diagnosis (Smirnov 1998). Ten or 12 digitate or peltato -digitate
tentacles. Plates of calcareous ring with large anterior
projections; excavations for tentacular ampullae are on anterior
side of calcareous ring. Madreporite placed close to water ring.
No ciliated funnels. One polian vesicle. Body wall ossicles
represented by wheels with large numbers of spokes (8-25) and
without a complex hub (single family Myriotrochidae).

Myriotrochidae Theel, 1877
Diagnosis. As for suborder.

Acanthotrochus antarcticus Belyaev and Mironov, 1981
Table 2

Acanthotrochus antarcticus Belyaev and Mironov, 1981a: 526-528, pi.
1(4-7), figs 3a-d, tables 3, 4.— Belyaev and Mironov, 1982: 108, fig. 18.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

65

Table 2. Antarctic species of Apodida, and their distributions. Chiridota pisanii Ludwig* and Taeniogyrus purpureus (Lesson)* are listed but to
date have not been recorded south of the Polar Front and are not Antarctic. Except for undescribed species, sources of data are given in the text.

Taxon

Distribution

Myriotrochidae Theel, 1877

Acantliotrochus antarcticus Belyaev and Mironov, 1981

Eastern Antarctica, 65°S 155°E, 2800 m.

Acanthotrochus species (by Bohn in Gebruk et al. 2003, p. 119)

South Orkney Is, 2914 m.

Achiridota smirnovi sp. nov.

Prydz Bay, Amery Depression, Fram Bank, 518-788 m.

Myriotrochus antarcticus Smirnov and Bardsley, 1997

Eastern Antarctica, MacRobertson Shelf, 113 m;

Western Antarctica, S Orkney Is, 216 m; Weddell Sea, 193 m.

Myriotrochus hesperides O’Loughlin and Manjon-Cabeza, 2009

Antarctic Peninsula, 65.47°S 69.03°W, 350 m.

Myriotrochus macquariensis Belyaev and Mironov, 1981

SW Pacific Ocean, Hjort Trench, 59°S 158°E, 3010-4640 m.

Myriotrochus nikiae sp. nov.

Eastern Antarctica, Ross Sea, 71°S 175°E, 2283 m.

Myriotrochus species (in Belyaev and Mironov 1982, p. 104)

Drake Passage, South Sandwich Trench.

Myriotrochus species (by Bohn in Gebruk et al. 2003, p. 119)

South Orkney Is, NW Weddell Sea, 2084-5190 m.

Neolepidotrochus variodentatus (Belyaev and Mironov, 1978)

South Sandwich Trench, 6766-7934 m.

Prototrochus barnesi sp. nov.

Scotia Sea, Shag Rocks, 206 m.

Prototrochus bipartitodentatus (Belyaev and Mironov, 1978)

South Sandwich Trench, 7700-8100 m.

Prototrochus linseae sp. nov.

Scotia Sea, South Shetland Is, 192-1544 m.

Prototrochus species (in Belyaev and Mironov 1982, pp. 92, 93)

South Sandwich Trench, 6050-6150 m.

Prototrochus species (by Bohn in Gebruk et al. 2003, p. 119)

South Orkney Is, 2375-5190 m.

Chiridotidae Ostergren, 1898

*Chiridota pisanii Ludwig, 1886

South America S of 42°S, Falkland Is, Burdwood Bank,
0-102 m.

Kolostoneura grijfithsi sp. nov.

Scotia Sea, South Orkney Is, 506 m.

Scoliorhapis biopearli sp. nov.

South Shetland Is, 1544 m.

Scoliorhapis massini sp. nov.

Scotia Sea, Shag Rocks, 206 m (? Falkland Is, 118 m).

Sigmodota contorta (Ludwig, 1875)

Antarctic Ocean, Bouvet I., South Georgia, S Shetland Is, S
Orkney Is, 46-503 m.

South America, south of 42° in the west, S of 47°S in the
east. Indian Ocean, Heard, Kerguelen, Marion Is, 2-228 m.
Indonesia, Java Sea, 82 m.

Sigmodota magnibacula (Massin and Heterier, 2004)

Western Antarctica, Weddell Sea, S Orkney Is, 172-240 m.
Eastern Antarctica, Ross Sea, Terre Adelie, Wilkes Land,
Prydz Bay, MacRobertson Shelf, 8-525 m.

Taeniogyrus antarcticus Heding, 1931

Scotia Sea, S Orkney Is, South Georgia, Shag Rocks,
206-216 m.

Taeniogyrus prydzi sp. nov.

Eastern Antarctica, MacRobertson Shelf, 109-121 m;
Prydz Bay Channel, Outfall slope, 795-830 m.

*Taenio gyrus purpureus (Lesson, 1830)

Falkland Is; Magellanic region, 0-64 m.

Paradota weddellensis Gutt, 1990

Antarctic Ocean, Antarctic Peninsula, 126-265 m;
Bellingshausen Sea, 97-1191 m; Ross Sea, 85-658 m; Prydz
Bay, 505-578 m; Scotia Sea, 59-759 m; Weddell Sea,
225-655 m; Heard I., 120-215 m.

Synaptidae Burmeister, 1837

Labidoplax species (by Bohn in Gebruk et al. 2003, p. 119)

South Orkney Is, 2893-3683 m.

66

P.M. O’Loughlin & D. VandenSpiegel

Figure 1. Colour photos of preserved specimens of apodid species, a, Prototrochus linseae sp. nov. (7 mm long; S Shetland Is; holotype, NMV
F168631); b, Kolostoneura griffithsi sp. nov. (12 mm long; insert drawings of tentacle rods, 96-112 pm long; S Orkney Is; holotype, NMV
F168634); c, Scoliorhapis biopearli sp. nov. (6 mm long; insert drawing of tentacle rod, 115 pm long; S Shetland Is; holotype, NMV F168633); d,
Scoliorhapis massini sp. nov. (20 mm long; Shag Rocks; holotype, NMV F168635); e, Sigmodota magnibacula (Massin and Heterier, 2004) (15
mm long; S Orkney Is; NMV F168629); f, Taeniogyrus antarcticus Heding, 1931 (15 mm long; S Orkney Is; NMV F168630).

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

67

Figure 2. a, drawing of tentacle and part of non-calcareous ring of Achiridota smirnovi sp. nov. (Prydz Bay; holotype, NMV F68687); b, drawing
of tentacle and part of calcareous ring of Kolostoneura griffithsi sp. nov. (S Orkney Is; holotype, NMV F168634).

Distribution. Eastern Antarctica, off Oates Land, Mawson
Peninsula, 65°S 155° E, 2800 m.

Remarks. The distinguishing characters of species of
Acanthotrochus Danielssen and Koren, 1879 are: wheel ossicles
of more than one type, some with outward-pointing teeth on
rim; wheels with outward pointing teeth lack inward-pointing
teeth; anterior projections of the calcareous ring plates longer
than the basal height of the plate; radial canal pore situated
lower than the base of the anterior plate projection (see Gage
and Billet 1986).

Achiridota Clark, 1908

Achiridota H. L. Clark, 1908: 126.— Heding, 1935: 16.— Smirnov,
1998: 521.

Diagnosis (of type species, following Fisher 1907, and Smirnov
1998). Tentacles 12, trunk stout, digits small; 6-8 pairs of
digits per tentacle, increasing in size distally, distal digits
paired not single; calcareous ring well developed, radial and
interradial plates with anterior projection/tooth, straight
posterior margin; lacking ossicles; single polian vesicle;
madreporite at anterior edge of dorsal mesentery, close to ring
canal; gonad tufts with central trunk and simple or dichotomous
branches.

Type species. Anapta inermis Fisher, 1907 (Hawaiian Is, 466-
772 m).

Other species. Achiridota profunda Heding, 1935 (N Atlantic,
2700 m); Achiridota smirnovi sp. nov. (E Antarctica, 518-788 m).

Remarks. Achiridota Clark, 1908 was initially assigned to
Chiridotidae, but was subsequently assigned to Myriotrochidae
by Smirnov (1998) on the basis of having: large anterior teeth on
the plates of the calcareous ring; single polian vesicle; madreporite
close to ring canal. Although Heding (1935) referred two new

species to Achiridota he argued convincingly that one of his
species ( Achiridota ingolfi Heding, 1935) was more Chiridota-
like because of the form of the calcareous ring and presence of 1 1
polian vesicles. We judge that Achiridota ingolfi belongs more
appropriately in Chiridota Eschscholtz, 1829.

Achiridota smirnovi sp. nov.

Figure 2a; table 2

chiridotid sp. MoV 2019 O’Loughlin et al., 1994: 553, 554.

Material examined. Holotype. Eastern Antarctica, Prydz Bay, Amery
Depression, 68°06'S 72°15'E, 788 m, stn ANARE AA93-60, M.
O’Loughlin, 28 Jan 1993, NMV F68687.

Paratype. Prydz Bay, edge of Fram Bank, 66°55’S 69°12’E, 518 m,
stn ANARE AA93-75, 31 Jan 1993, NMV F68686.

Diagnosis. Achiridotid species up to 15 mm long (posterior end
of body missing on 11 mm long holotype); 12 peltato-digitate
tentacles, 4 pairs of digits per tentacle, pair distally, increasing
in size distally; tentacle ampullae cup-like, on anterior edge of
non-calcareous ring; lacking ossicles in body wall, tentacles;
lacking calcareous ring; single polian vesicle; madreporite on
long straight canal; gonad tubules with multiple branching;
lacking ciliated funnels.

Colour (preserved). Holotype with reddish-brown flecking on
semi-translucent off-white body, red-brown tentacles; other
specimen semi-translucent off-white, tentacles pale reddish-
yellow.

Distribution. Eastern Antarctica, Prydz Bay, 518-788 m.

Etymology. Named for Alexei Smirnov (Zoological Institute of
the Russian Academy of Sciences, St. Petersburg) in recognition
of his description of the first myriotrochid species from the
Antarctic shelf, and with appreciation of his significant
contribution to the systematics of Apodida.

P.M. O’Loughlin & D. VandenSpiegel

Remarks. Apodid specimens collected from Prydz Bay during the
same cruise as these specimens, and preserved in the same way
(directly in 70% ethanol), have retained their ossicles in good or
slightly eroded condition. It is most unlikely that the complete
absence of calcareous parts in the two specimens on which
Achiridota smirnovi sp. nov. is based is a result of preservation.

Generic assignment of this new species is problematic
because of the absence of a calcareous ring and body wall
ossicles. But the tentacles do arise from anterior cup-like
depressions around the non-calcareous ring, as in Achiridota H.
L. Clark. The species has the Achiridota diagnostic characters of:
12 peltato-digitate tentacles; single polian vesicle; lacking
ossicles in tentacles, body wall; lacking ciliated funnels. But the
madreporite is situated at the end of a long straight canal, and is
not close to the water canal. We assign the new species to
Achiridota Clark with reservations because of the absence of a
calcareous ring and position of the madreporite distant from the
ring canal. These two characters distinguish Achiridota smirnovi
sp. nov. from Achiridota inermis (Fisher) and Achiridota profunda
Heding. Additional diagnostic distinctions are the presence of
6-8 pairs of tentacle digits in Achiridota inermis (4 pairs in A.
smirnovi), and 7-8 pairs of tentacle digits and unbranched gonad
tubules in Achiridota profunda (branched in A. smirnovi).

Myriotrochus Steenstrup, 1851

Diagnosis ( following Gage and Billet 1986). Myriotrochid
with 12 tentacles; calcareous ring bilaterally symmetrical;
dorsolateral radial plates with two anterior projections; wheel
ossicles of one type with rim teeth pointing only towards centre
of hub; wheel hub lacking holes, and if holes are present they
are distributed regularly in a circle around the centre of the
hub; rod ossicles absent.

Myriotrochus antarcticus Smirnov and Bardsley, 1997
Figures 3, 4; table 2

Myriotrochus sp. MoV 2039 O’Loughlin et ah, 1994: 553, table 2.

Myriotrochus antarcticus Smirnov and Bardsley, 1997: 109-111,
fig. 1, table 1.— O’Loughlin et al. 2009: 9.

Material examined. Holotype. Eastern Antarctica, 66°55'S 62°32'E,
113 m, M. O’Loughlin, 11 Feb 1993, NMV F69125.

Other material. Western Antarctica, South Orkney Is, 60.82°S
46.49°W, 216 m, BAS stn PB-EBS-4, 18 Mar 2006, NMV F168638

(1) ; F168643 (1 whole, for molecular sequence, tissue code MOL AF
805); RBINS IG 31 459 (2 whole, for SEM figures); NHM 2010. 48-49

(2) ; Weddell Sea, 71.25° S 13.00° W, 193 m, RBINS 628686 (1).

Diagnosis ( following Smirnov and Bardsley 1997 for Prydz
Bay holotype). Myriotrochid species up to 8 mm long; 12
peltato-digitate tentacles, up to 8-12 digits per tentacle,
distalmost pair longest; lacking tentacle ossicles; calcareous
ring comprising 10 plates, ventral plates not significantly longer
than dorsal plates; two dorsolateral radial plates each with 2
prominent tapering anterior projections, remaining plates with
single prominent tapering anterior projection; posterior margin
of calcareous ring slightly undulating, not concave; ossicles
myriotrochid wheels only, scattered sparsely and uniformly
throughout body wall; wheels of one type, all teeth pointing to

centre of hub; wheel ossicle diameters 140-150 pm (for S
Orkneys specimens 80-144 pm\ for Weddell Sea specimen
108-133 pm), spokes 15-16 (for S Orkneys 13-16; for Weddell
Sea 13-15), teeth 22-24 (for S Orkneys 19-25; for Weddell Sea
22-24), spokes/teeth % 66.7-68.2 (for South Orkneys 64-68;
for Weddell Sea 60.8-65.2), hub diameter/wheel diameter %
18.0-18.6 (for S Orkneys 16.0-18.1), teeth length/wheel
diameter % 18.0-18.6 (for S Orkneys 14.4-16.0).

Colour (preserved). Body grey, translucent; tentacles white.

Distribution. Antarctic shelf species; Eastern Antarctica,
western MacRobertson Shelf, 113 m; Western Antarctica, South
Orkney Is, 216 m; Weddell Sea, 193 m (range 113-216 m).

Remarks. For Myriotrochus antarcticus Smirnov and Bardsley,
1997 there are only minor morphological differences for the
limited number of measurements for specimens from the Scotia
Sea, Weddell Sea and Prydz Bay, suggesting a morpho-species
with an eastern and western Antarctic distribution.

Myriotrochus hesperides O’Loughlin and Manjon-Cabeza, 2009
Figure 5a; table 2

Myriotrochus hesperides O’Loughlin and Manjon-Cabeza (in
O’Loughlin et al., 2009): 9, fig. 2e, f, table 1.

Diagnosis ( following O’Loughlin et al. 2009). Myriotrochid
species up to 13 mm long; 12 peltato-digitate tentacles, about 7
small rounded digits per tentacle; lacking tentacle ossicles;
plates of calcareous ring asymmetrical with pointed anterior
projections / teeth, 2 radial plates each with 2 anterior
projections, remaining plates with single anterior projection,
wide rounded tongue -like posterior projections of variable
length; ossicles myriotrochid wheels only, few only in posterior
dorsal body wall; wheels of one type, all teeth pointing towards
centre of hub; spokes irregular, about half branching proximally,
some branches not reaching rim, some spokes with cross-
connections; teeth variably sub-equal or different in size; hubs
small, irregular, not disc-like, lacking perforations, formed by
junction of spokes; largest wheel with diameter 248 pm, hub
diameter 40 pm, 13 spokes at hub, 23 spokes at rim, 30 equal
teeth; smallest wheel with diameter 200 pm, hub diameter 24
pm, 12 spokes at hub, 16 spokes at rim, 28 unequal teeth.

Distribution. Antarctic Peninsula, 65.47°S 69.03°W, 350 m.

Remarks. Myriotrochus hesperides O’Loughlin and Manjon-
Cabeza, 2009 was the second Antarctic shelf Myriotrochus
species to be described. Amongst Myriotrochus species it is
closest to Myriotrochus clarki Gage and Billett, 1986 from the
Rockall Trough in the N Atlantic at 1605-2515 m. The species
are distinguished in O’Loughlin et al. 2009.

Myriotrochus macquariensis Belyaev and Mironov, 1981
Table 2

Myriotrochus macquoriensis Belyaev and Mironov, 1981b: 169-170,
fig. 4, tables 3, 4, pi. figs 4, 5.— Belyaev and Mironov, 1982: 105, fig. 15.

Distribution. SW Pacific/Antarctic Ocean, Hjort Trench, 59°S
158°E, 3010-4640 m.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

69

Figure 3. Photo of Myriotrochus antarcticus Smirnov and Bardsley, 1997, showing 12 peltato-digitate tentacles, calcareous ring with high
anterior pointed projections, and wheels distributed throughout body wall (7 mm long; S Orkney Is, 216 m; RBINS IG 31 459).

Remarks. The spelling of the species name is corrected as
reported at the beginning of this paper.

Myriotrochus nikiae sp. nov.

Figure 5b; table 2

Material examined. Holotype. Antarctica, Ross Sea, 71.23° S 174.44°
E, 2281-2283 m, NZ IPY-CAML stn TAN0802/171, N. Davey, 26 Feb
2008, NIWA 37812 (in two parts).

Diagnosis. Myriotrochid species up to 33 mm long (2 parts
combined); body wall thick, soft; 12 tentacles, withdrawn;
lacking tentacle ossicles; calcareous ring comprising 10 plates;
two dorsolateral radial plates each with 2 prominent tapering
anterior projections, remaining plates with single prominent
tapering anterior projection; posterior margin of calcareous
ring slightly undulating, not concave; sac -like calcareous
madreporite, dorsal, close to ring; single ventral polian vesicle;
gonad comprises thick digitiform unbranched tubules arising
in a series along gonoduct, 9 on one side of mesentery; ossicles
in body wall myriotrochid wheels only, found throughout body
wall; wheels of one type, all teeth pointing to centre of hub; hub
large and disc-like, perforated, with regular circle of
perforations separated by sometimes irregular hub spokes
aligned with longer outer wheel spokes, some hub spokes
branched distally; teeth blunt and rounded, of irregular length,
generally longer over spokes than between spokes.

Measurements for 9 wheels: wheel ossicle diameters 320-
400 pm, hub disc diameters 168-216 pm, inner hub diameter
24-32 pm, teeth length 48 pm, spokes 11-15, teeth 22-36,
spokes/teeth % 36-50, hub disc diameter/wheel diameter %
51-52, teeth length/wheel diameter % 12-15.

Colour (preserved). Body grey, not translucent; few small dark
brown spots on tentacles.

Distribution. Eastern Antarctica, Ross Sea, 2283 m.

Etymology. Named for Niki Davey (NIWA), with appreciation
of her generous and skilled collaborative assistance in
determining Antarctic and New Zealand holothuroids.

Remarks. The specimen size, grey colour, form and size of the
wheels, and form of the calcareous ring of Myriotrochus nikiae
sp. nov. are similar to those of Myriotrochus bathybius H. L.
Clark, 1920 from the eastern tropical Pacific Ocean (4°33'S
87°43'W, 3669 m), and according to Gage and Billett (1986)
from the northeast Atlantic Ocean (Rockall Trough, 1800-
2925 m, and Porcupine Seabight, 3680-4310 m). Gage and
Billett (1986) further judged Myriotrochus bathybius to be
cosmopolitan at abyssal depths. Differences are such that we
judge that the Antarctic specimen represents a related but
different species.

For Myriotrochus bathybius , Clark (1920) gave three
wheel diameters of 240, 300 and 340 pm (size range smaller
than for M. nikiae ), and two teeth counts of 37 and 38 (more
numerous than for M. nikiae). More significant for us is a
wheel of M. bathybius that was illustrated in Clark 1920, and
the teeth are uniform in length and distinctly pointed. Smirnov
(1999, fig. 4) illustrated similar teeth for Myriotrochus
( Oligotrochus ) bathybius. The form of these teeth is in contrast
with the irregular lengths of the quite rounded teeth of M.
nikiae. Clark (1920) noted seven dark spots between tentacle
bases. These are not present in the Antarctic specimen, but
some dark spots are present on the tentacles.

70

P.M. O’Loughlin & D. VandenSpiegel

Figure 4. Myriotrochus antarcticus Smirnov and Bardsley, 1997 (S Orkney Is, 216 m; RBINS IG 31 459). a, SEM of plates of the calcareous ring
with prominent anterior projections and undulating posterior margin and canals, dorso-lateral radial plate (lower left) with two anterior
projections; b, SEM of wheels from the body wall.

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71

Figure 5. a, montage photo of wheels from body wall of holotype of Myriotrochus hesperides O’Loughlin and Manjon-Cabeza, 2009 (Antarctic
Peninsula; slide NMV F161516); b, montage photos of wheels from body wall of holotype of Myriotrochus nikiae sp. nov., with insert drawing of
dorso-lateral radial (2 anterior projections) and adjacent inter-radial (single anterior projection) plates from calcareous ring (Ross Sea; NIWA 37812).

For Myriotrochus bathybius, Gage and Billett (1986)
reported a possible rough texture of the body wall in larger
specimens (soft, thick body wall for M. nikiae), and the
calcareous ring clearly visible through the body wall (not
visible through the thick body wall of M. nikiae ). Gage and
Billett (1986) illustrated tapered teeth of uniform length with
narrowly rounded points (in contrast with the bluntly rounded
teeth of variable length for M. nikiae).

As noted in the diagnosis the tentacles of the single
specimen of Myriotrochus nikiae are withdrawn, and it was
not possible to confidently describe them as “conical with
lateral digits” or “peltato-digitate” as required for assignment
to subgenus Oligotrochus M. Sars or subgenus Myriotrochus
Steenstrup respectively (see Smirnov 1999).

Neolepidotrochus variodentatus (Belyaev and Mironov, 1978)
Table 2

Myriotrochus variodentatus Belyaev and Mironov, 1978: 202-
204, fig. 4, tables 1, 4, pi. 1 figs 4-6, 9.

Lepidotrochus variodentatus Belyaev and Mironov, 1980:
1812, 1818, tables 1, 5.— Belyaev and Mironov, 1982 : 109, fig. 18.

Neolepidotrochus variodentatus Bohn, 2005: 234.

Distribution. South Sandwich Trench, 6766-7934 m.

Remarks. Bohn (2005) recognized that Lepidotrochus Belyaev
and Mironov, 1980 is a junior homonym of Lepidotrochus
Koken, 1894, and erected the replacement name
Neolepidotrochus. The distinguishing characters of species of
Neolepidotrochus Bohn, 2005 are: wheel ossicles with
outward-pointing teeth also have inward-pointing teeth;
anterior projections of the calcareous ring plates are lower than
the basal height of the plates; radial canal pore is situated
higher than the base of the anterior plate projection (see Gage
and Billet 1986).

Prototrochus Belyaev and Mironov, 1982

Diagnosis ( after Gage and Billett 1986). Myriotrochid with 10
tentacles; calcareous ring symmetrical, with dorsal and ventral
plates subequal in size; dorsolateral radial plates with single
anterior projection; wheels with teeth distributed regularly,
pointing towards centre of hub; rods absent from body wall,
sometime occurring in and around tentacles.

Remarks. Belyaev and Mironov (1982) noted in erecting their
new genus Prototrochus that the wheels of their included
species Prototrochus bipartitodentatus (Belyaev and Mironov,
1978) were exceptional within their diagnosis as there are small
external teeth at the base of the internal teeth.

Prototrochus barnesi sp. nov.

Figure 6; table 2

Material examined. Holotype. Antarctica, Scotia Sea, Shag Rocks,
53.63°S 40.91° W, 206 m, BAS stn SR-EBS-4, 11 Apr 2006, NMV
F168637.

Paratypes. Type locality and date, NHM 2010.54 (1); RBINS IG
31 459 (1, SEM).

Diagnosis. Myriotrochid species up to 3 mm long; 10 peltato-
digitate tentacles, 7 digits per tentacle, including a distal
terminal one; tentacle rods present, straight and curved, some
with central swelling, some with swollen end, 40-170 pm long;
sparse myriotrochid wheels in body wall, slightly scalloped
margin at each tooth; wheel ossicle diameters 72-104 pm,
spokes 13-15, teeth 22-27, spokes/teeth % 50-59, hub
diameters 19-26 pm, hub diameter/wheel diameter % 25-38,
teeth length/wheel diameter % 12-20.

Colour (preserved). Off-white.

Distribution. Western Antarctica, Scotia Sea, Shag Rocks, 206 m.

72

P.M. O’Loughlin & D. VandenSpiegel

Figure 6. Prototrochus barnesi sp. nov. (Shag Rocks; RBINS IG 31 459). a, SEM of plates of the calcareous ring with prominent single anterior
projections; b, SEM of wheels from the body wall; c, SEM of rods from tentacles.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

73

Etymology. Named for David Barnes (British Antarctic
Survey), in appreciation of his role in the BAS BIOPEARL
expeditions and the collection of specimens studied here.

Remarks. Belyaev and Mironov (1982) referred 12 species to
their new genus Prototrochus . O’Loughlin and VandenSpiegel
(2007) added three new Prototrochus species from the
continental slope of Australia, all lacking tentacle ossicles. The
only Prototrochus species recorded from Antarctica is
Prototrochus bipartitodentatus (Belyaev and Mironov, 1978)
from the South Sandwich Trench at 7700-8100 m. This species
lacks tentacle rods and has external teeth around the rim of the
wheels. Prototrochus barnesi sp. nov. has wheels with internal
teeth only, and has tentacle rods, a rare character for
Prototrochus species. The only other Prototrochus species with
tentacle rods is the similarly small Prototrochus minutus
(Ostergren, 1905), described from the coast of Korea at 60-65
m depth. Three diagnostic characters distinguish Prototrochus
minutus from Prototrochus barnesi : sometimes distally and
centrally branched tentacle rods; longer tentacle rods (mostly
140-200 pm long); significantly larger wheels (mostly 100-
150 pm diameter).

Prototrochus bipartitodentatus (Belyaev and Mironov, 1978)
Table 2

Myriotrochus bipartitodentatus Belyaev and Mironov, 1978: 201-
202, fig. 3, tables 1,3, pi. 1 fig. 1.

Prototrochus bipartitodentatus.— Belyaev and Mironov, 1982:
86, 92, fig. 6.

Distribution. South Sandwich Trench, 7700-8100 m.

Remarks. Prototrochus bipartitodentatus (Belyaev and
Mironov) is distinguished from all other Prototrochus species
by the presence of small external teeth at the base of the internal
teeth of the wheels.

Prototrochus linseae sp. nov.

Figures la, 7; table 2

Material examined. Holotype. Antarctica, Scotia Sea, South Shetland
Is, 62.53°S 61.83°W, 192 m, BAS stn LI-EBS-4, 4 Mar 2006, NMV
F168631.

Paratypes. Type locality and date, NHM 2010.50 (1); RBINS IG
31 459 (2, SEM).

Other material. South Shetland Is, 61.61°S 55.22°W, 1544 m, BAS
stn EI-EBS-1, 12 Mar 2006, NHM 2010.53 (1).

Diagnosis. Myriotrochid species up to 7 mm long; 10 tentacles;
lacking tentacle rods; scattered myriotrochid wheels only in
body wall; wheel ossicle diameters 125-137 pm, spokes 10-11,
teeth 22-24, spokes/teeth % 45, hub diameters 30-34 pm, hub
diameter/wheel diameter % 24.0-24.7, teeth length/wheel
diameter % 12.9-13.0.

Colour (preserved). Body pale yellow-brown to off-white,
translucent; tentacles yellow.

Distribution. Antarctica, Scotia Sea, South Shetland Is, 192-
1544 m.

Etymology. Named for Katrin Linse (British Antarctic Survey),
in appreciation of her role in the BAS BIOPEARL expeditions
and the collection of specimens studied here, and with gratitude
for her gracious collaboration in making BAS specimens
available for this study and providing relevant data.

Remarks. Most of the 12 species referred to their new genus
Prototrochus by Belyaev and Mironov (1982) lack tentacle
ossicles, as does Prototrochus linseae sp. nov. Most of the
species lacking tentacle ossicles are from the deep trenches
(6450-10700 m in Belyaev and Mironov 1882). Only
Prototrochus bipartitodentatus (Belyaev and Mironov, 1978)
has been recorded from the Antarctic (South Sandwich Trench,
7700-8100 m). It has external teeth around the rim of the wheels
that Prototrochus linseae does not. Belyaev and Mironov (1982)
list three shallower species (540-3000 m), two from the
Mediterranean Sea and Prototrochus australis (Belyaev and
Mironov, 1981) from the Tasman Sea. O’Loughlin and
VandenSpiegel (2007) list three additional species from the
Tasman Sea: Prototrochus burni O’Loughlin, 2007*,

Prototrochus staplesi O’Loughlin, 2007* and Prototrochus
taniae O’Loughlin, 2007* (*all in O’Loughlin and
VandenSpiegel 2007). As demonstrated by reference to Table 1
and Figure 10 in O’Loughlin and VandenSpiegel (2007),
Prototrochus linseae is distinguished from the four Tasman Sea
Prototrochus species by the form of the wheels: smallest wheels;
largest teeth; widest uniformly broad spokes, with sharp
constriction at the hub; largest hub.

Suborder Synaptina Smirnov, 1998

Diagnosis ( Smirnov , 1998). Plates of calcareous ring without
prominent anterior projections; excavations for tentacular
ampullae lie on outer side of calcareous ring. Madreporite
situated far from water ring at end of long stone canal. Ciliated
funnels present. One to many polian vesicles. Body wall
ossicles may be wheels of chiridotid type with 6 spokes and a
complex hub and/or sigmoid hooks, or anchors and anchor
plates. Wheels of larvae and juveniles with more spokes and
small denticles on inner side of rim.

Families. Chiridotidae Ostergren, 1898; Synaptidae Burmeister,
1837.

Chiridotidae Ostergren, 1898

Diagnosis ( Smirnov 1998). Synaptina with 10, 12 or 18 peltato-
digitate, pinnate or bifurcate tentacles. Juveniles with bifurcate
tentacles. Body wall ossicles wheels of chiridotid type and/or
sigmoid hooks. Chiridotid type wheels with 6 spokes, numerous
small denticles on inner side of rim and a complex hub; on
lower side of each spoke a branch leans against the lower end of
the hub forming a star structure. Ossicles in tentacles usually
rods with branched ends.

Subfamilies. Taeniogyrinae Smirnov, 1998; Chiridotinae
Ostergren, 1898 (sensu Smirnov 1998).

Subfamily Taeniogyrinae Smirnov, 1998

74

P.M. O’Loughlin & D. VandenSpiegel

Figure 7. Prototrochus linseae sp. nov. (S Shetland Is; RBINS IG 31 459). a, SEM of plates of the calcareous ring with prominent single anterior
projections; b, SEM of wheels from the body wall.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

75

Diagnosis (Smirnov 1998). Chiridotidae with 10 or 12 tentacles.
Body wall ossicles wheels of chiridotid type and sigmoid hooks
or sigmoid hooks only. Radial plates of calcareous ring not
perforated but sometimes slightly notched in anterior (upper)
face for passage of nerves.

Included genera. Archedota O’Loughlin, 2007 (in O’Loughlin
and VandenSpiegel 2007); Kolostoneura Becher, 1909;
Rowedota gen. nov.; Scoliorhapis H. L. Clark, 1946; Sigmodota
Studer, 1876; Taeniogyrus Semper, 1867.

Remarks. Achiridota Clark, 1908 was initially placed in
Chiridotidae, but was assigned to Myriotrochidae by Smirnov
(1998) on the basis of having: large anterior teeth on the plates
of the calcareous ring; single polian vesicle; madreporite close
to calcareous canal .Sigmodota Studer is raised out of synonymy
with Taeniogyrus Semper (below). Trochodota Ludwig, 1891 is
made a junior synonym of Taeniogyrus Semper (below).
Scoliodotella Oguro, 1961 is made a junior synonym of
Scoliorhapis H. L. Clark (below).

Kolostoneura Becher, 1909

Kolostoneura Becher, 1909: 42.— H. L. Clark, 1921: 164.—
Mortensen, 1925: 384-386.— Heding, 1928: 277, 278.— Pawson, 1970:
44.— Smirnov, 1998: 519.

Diagnosis (of type species, following Dendy and Hindle 1907
and Mortensen 1925). Taeniogyrinid genus with 10 peltato-
digitate tentacles, each with 12 digits increasing in size distally;
tentacle rods present; lacking ossicles in body wall; calcareous
ring present, 5 subrectangular, transversely elongate radial and
5 subequal, interradial plates, lacking anterior projections /
teeth; single polian vesicle; madreporite canal long, straight,
madreporite distant from water ring; branched gonad tubules;
ciliated funnels present.

Type species. Rliabdomolgus novae -zealandiae Dendy and
Hindle, 1907 (New Zealand, Chatham Is, coastal shallows).

Other species. Kolostoneura griffithsi sp. nov. (Antarctica,
Scotia Sea, 506 m).

Remarks. Smirnov (1998) noted that although Kolostoneura
lacks ossicles in the body wall its morphological characters
place it near Taeniogyrus and Trochodota and thus in family
Taeniogyrinae. Mortensen (1925) found a very small, damaged
specimen that he judged to be Kolostoneura novae -zealandiae
(Dendy and Hindle) that was infested with parasitic snails and
had hooks and wheels in the body wall. He postulated that the
ossicles of the ancestral species were present through the
influence of the parasite. Perhaps ossicles are typically present
in small specimens of this species, and then lost as size increases
as happens with many holothuroid species (such as with the
apodid Taeniogyrus magnibaculus Massin and Heterier, 2004
below). If this is the case then genus Kolostoneura would be a
junior synonym of Taeniogyrus Semper, 1867 (see below).

Kolostoneura griffithsi sp. nov.

Figures lb, 2b; table 2

Material examined. Holotype. South Orkney Is, 60.99°S 46.83°W,
506 m, BAS stn PB-EBS-3, 18 Mar 2006, NMV F168634.

Diagnosis. Taeniogyrinid species 12 mm long; 10 peltato-
digitate tentacles, 3-4 pairs of digits per tentacle; no ossicles in
body wall; tentacle rods present, some with central hub, some
with bifurcate ends, 96-112 pm long; solid calcareous ring,
plates lack anterior projections/teeth, posterior margin slightly
concave; single polian vesicle; madreporite with short curved
canal, close to water ring; gonad tubules not present; lacking
ciliated funnels.

Colour (preserved). Dark purple-red flecking on grey to off-
white semi -translucent body wall.

Distribution. Antarctica, Scotia Sea, South Orkney Is, 506 m.

Etymology. Named for Huw Griffiths (British Antarctic
Survey), in appreciation of his role in the BAS BIOPEARL
expeditions and the collection of specimens studied here.

Remarks. Kolostoneura griffithsi sp. nov. exhibits the diagnostic
characters of the type species of Kolostoneura Becher, 1909,
except that it has fewer tentacle digits, has calcareous ring
plates that are not transversely elongate, and lacks ciliated
funnels. However, Mortensen (1925) noted that ciliated funnels
were sometimes scarce or absent along the mesentery in the
material that he examined.

Rowedota gen. nov.

Figure 8

Diagnosis. Taeniogyrinid genus with 10 tentacles, each with
1-4 pairs of digits; chiridotid-type wheels and sigmoid hooks
in body wall, rods in tentacles; wheels with teeth on inner rim
in 6 discrete groups, discontinuous between spokes, not
continuous series around rim; wheels scattered or aligned in
the body wall, not in distinct papillae; sigmoid hooks scattered;
single polian vesicle; ciliated funnels present.

Type species. Taeniogyrus allani Joshua, 1912 (see O’Loughlin
and VandenSpiegel 2007).

Other species. Trochodota epiphyka O’Loughlin, 2007 (in
O’Loughlin and VandenSpiegel); Trochodota shepherdi Rowe,
1976; Trochodota vivipara Cherbonnier, 1988; Trochodota
mira Cherbonnier, 1988.

Etymology. Named for Dr Frank W. E. Rowe, Senior Fellow of
the Australian Museum, with appreciation of his contribution
to echinoderm systematics and of his role as a valued mentor
and colleague.

Remarks. Rowe (1976) based his emended diagnoses of
Taeniogyrus and Trochodota on an important diagnostic
character that distinguished two types of wheels amongst
species of Taeniogyrinae, namely those with a continuous
series and those with discontinuous grouped teeth on the inner
rim. We judge that this is a significant generic character and on
this diagnostic character erect the new genus. The genus has
further morphological coherence with all included species
having 10 tentacles, up to only 4 pairs of tentacle digits, a single

76

P.M. O’Loughlin & D. VandenSpiegel

Figure 8. a, b, SEM of wheels from body wall of Rowedota allani (Joshua, 1912) illustrating the Rowedota gen. nov. generic character of
discontinuous series of teeth around the inner rim (SE Australia; from lot NMV F82715).

polian vesicle, and the wheels never grouped into discrete
papillae or bands. The new genus has geographical coherence
with all included species occurring on both sides of the Indian
Ocean off Australia and Madagascar.

We have included the presence of a single polian vesicle
and ciliated funnels in the generic diagnosis of Rowedota gen.
nov., but the presence of ciliated funnels in the Cherbonnier
(1988) species and presence of a single polian vesicle in
Trochodota mira have yet to be confirmed.

Frank Rowe (pers. comm.) considers the rods in the body wall
illustrated by Cherbonnier (1988) for his Trochodota mira to be
contaminants. Species to date referred to Trochodota, other than
those referred here to the new genus Rowedota, are referred to
Taenio gyrus (below). The wheels of Trochodota maculata H. L.
Clark, 1921 have continuous series of teeth in the inner rim (see
below). Rowe (1976) followed Clark (1921) who thought that the
wheels in his new species Trochodota maculata had discontinuous
series of teeth. The 1921 illustration of a wheel suggests that Clark
was viewing the wheel from the side on which the continuous
series is not evident as the series is partly obscured by the spokes.

No species of Rowedota gen. nov. occurs in Antarctica.
The new genus is erected within this revision of genera of
Taeniogyrinae.

Scoliorhapis Clark, 1946
Table 3

Scoliodota Heding, 1928: 277, 278, 319 (junior homonym of
Scoliodota H. L. Clark, 1908).

Scoliorhapis H. L. Clark, 1946: 461.— Rowe (in Rowe and Gates),
1995: 267.— Smirnov, 1998: 519,-Kerr, 2001: 57.-0’Loughlin and
VandenSpiegel, 2007: 53.

Scoliodotella Oguro, 1961: 2-3.

Diagnosis ( emended from H. L. Clark 1946). Taeniogyrinae
with 10 or 12 peltato-digitate tentacles, each with up to 8 pairs

of digits; single polian vesicle; ciliated funnels present; body
wall ossicles sigmoid hooks only, hooks scattered or some
clustering or alignment; lacking wheels in body wall; tentacle
ossicles bracket- shaped or rods.

Type species. Scoliodota theeli Heding, 1928.

Type locality. Australia, New South Wales, Port Jackson.

Other species. Scoliorhapis biopearli sp. nov. (Scotia Sea, S
Shetland Is, 1544 m); Scoliodota lindbergi Djakonov (in
Djakonov et al.), 1958 (Sea of Okhotsk, South Sakhalin and
South Kurile Is, 8-22 m); Scoliorhapis massini sp. nov. (Scotia
Sea, Shag Rocks, 206 m); Scoliodotella uchidai Oguro, 1961
(Cape Aikappu, Japan, shallows).

Remarks. The significant emendation to the diagnosis by H. L.
Clark (1946) is the inclusion of species that have 12 tentacles.
This is in response to Scoliorhapis massini sp nov. (below)
having 12 tentacles.

Oguro (1961) erected his new genus Scoliodotella for
chiridotid specimens with sigmoid hooks but no wheels in the
body wall. He considered referring his new species to
Scoliodota H. L. Clark, 1908, unaware that H. L. Clark had
rejected Scoliodota and erected the genus Scoliorhapis. Oguro
did not refer his new species to Scoliodota because the hooks
were not grouped into papillae. We reject clustering of ossicles
as a sound generic diagnostic character, and consider
Scoliodotella to be a junior synonym of Scoliorhapis.

Chiridota japonica Marenzeller, 1881 was erected for
material from Japan. The species was poorly described. Theel
(1886a) referred damaged specimens from Australia (New
South Wales) to Chiridota japonica Marenzeller. Scoliodota
H. L. Clark, 1908 was erected as a new monotypic genus for
Chiridota japonica Marenzeller from Japan and the Theel
material from New South Wales on the basis of hook ossicles
only in the body wall. But Clark (1908) commented that the

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77

Table 3. Specimen and ossicle sizes for some species of Taeniogyrinae.

Selected species of Scoliorhapis,
Sigmodota and Taeniogyrus

Locality (this work)
or authors

Maximum

preserved

length

Wheel

diameter

pm

Hook

length

pm

Tentacle rod

length

pm

Tentacle

number

Scoliorhapis species

Soliorhapis theeli (type species)

Heding 1928

no data

absent

100-140

106

10

Scoliorhapis biopearli sp. nov.

S Shetland Is (this work)

6 mm

absent

168-184

112-120

10

Scoliorhapis massini sp. nov.

Shag Rocks (this work)

20 mm

absent

88-104

up to 80

12

Scoliorhapis ? massini sp. nov.

Falkland Is (this work)

50 mm

absent

120-176

120-160

12

Sigmodota species

Sigmodota contorta
(type species for Sigmodota )

Ludwig 1875

45 mm

94

140-160

no data

12

Ludwig 1898

45 mm

44—130

170-200

156-182

12

H.L. Clark 1921

no data

42-130

170-210

170

12

Ekman 1925

44 mm

45-120

140-170

no data

no data

Pawson 1964

50 mm

42-130

140-200

170

12

Massin & Heterier 2004

45 mm

35-100

160-250

140-170

12

Bouvet I. (this work)

10 mm

40-88

224-240

120-128

12

Falkland Is (this work)

40 mm

40-128

136-200

80-184

12

S Georgia (this work)

45 mm

48-96

192-248

80-144

12

S Georgia (this work)

8 mm

56-72

no data

up to 128

12

S Orkney Is (this work)

9 mm

40-88

152-184

96-112

12

S Shetland Is (this work)

15 mm

56-104

168-176

96-112

12

Magellanic (this work)

40 mm

up to 136

up to 208

up to 184

12

Heard I. (this work)

no data

88-104

192-208

104-168

12

Summary

50 mm

35-136

136-250

80-184

12

Sigmodota dubia

Fisher 1907

60 mm

90-175

185-230

no data

12

Sigmodota magnibacula

Massin & Heterier 2004

72 mm

100-200

160-180

170-280

12

S Orkney Is (this work)

70 mm

112-160

192-216

256-296

12

S Orkney Is (this work)

15 mm

56-120

128-176

176-216

12

Ross Sea (this work)

25 mm

96-136

152-200

192-240

12

Ross Sea (this work)

73 mm

110-140

170-190

200-270

12

Wilkes Land (this work)

60 mm

up to 104

up to 192

up to 272

12

Prydz Bay (this work)

105 mm

80-186

176-192

184-320

12

Summary

105 mm

56-200

128-216

170-320

12

Taeniogyrus species

Taeniogyrus antarcticus

Heding 1931

no data

no data

172-200

83-103

10

S Orkney Is (this work)

15 mm

64-80

200-208

128

10

S Orkney Is (this work)

7 mm

40-64

120-152

80-104

10

S Orkney Is (this work)

3 mm

48-72

112-120

128

10

S Orkney Is (this work)

2 mm

40-56

80-88

80

10

Shag Rocks (this work)

8 mm

48-64

144-168

no data

10

Shag Rocks (this work)

1 mm

48-64

64-80

64-72

10

Summary

15 mm

40-80

64-208

64-128

10

Taeniogyrus australianus
(type species for Taeniogyrus )

Australia (this work)

95 mm

48-88

112-136

72-104

10

Heding 1928

60 mm

70-100

up to 110

110

10

Summary

95 mm

48-100

110-136

72-110

10

Taeniogyrus maculatus

H.L. Clark 1921

26 mm

50-100

66-77

about 45

10

Newcastle (this work)

21 mm

48-104

80-96

56-72

10

Summary

26 mm

48-104

66-96

45-72

10

Taeniogyrus prydzi sp. nov.

Prydz Bay (this work)

7 mm

up to 90

232-240

136-144

10

Prydz Bay (this work)

50 mm

up to 90

256-272

136-152

10

Summary

50 mm

up to 90

232-272

136-152

10

Taeniogyrus purpureus

Ludwig 1898

33 mm

165-182

135-156

78-87

10

Ekman 1925

33 mm

70-180

80-150

no data

no data

H.L. Clark 1921

no data

154-182

125-150

76-87

no data

Pawson 1964

100 mm

130-180

120-130

average 78

10

Magellanic (this work)

15 mm

80-144

120-160

56-120

10

Summary

100 mm

70-182

80-160

56-120

10

78

P.M. O’Loughlin & D. VandenSpiegel

Marenzeller and Theel materials might represent different
species. Ohshima (1913) reported that Chiridota japonica von
Marenzeller had both hook and wheel ossicles, and belonged
to Trochodota Ludwig, 1891.

Clark (1921) rejected his own genus Scoliodota Clark, 1908
on the grounds that his type species Chiridota japonica von
Marenzeller belonged to Trochodota Ludwig. Heding (1928)
stated that Clark (1921) abandoned Scoliodota Clark without
considering the Theel specimens from New South Wales.
Heding (1928) erected a new species for the New South Wales
specimens that lacked wheels, and referred them to Scoliodota
Clark. Heding (1928) nominated his Scoliodota theeli Heding,
1928 as the new type for Scoliodota Clark, 1908.

Clark (1946) insisted that he selected Japanese material as
type for his Scoliodota, although that is not explicit in the text
(Clark 1908). Clark (1946) rejected Heding’s resurrection of
Scoliodota Clark, and erected anew monotypic genus Scoliorhapis
Clark, 1946 with type species Scoliodota theeli Heding, 1928.

Clark (1921) noted that Chiridota geminifera Dendy and
Hindle, 1907 had hook ossicles but lacked wheels, but because
of the single small and damaged type specimen considered the
material to be unreliable for referral to Scoliodota Clark. He
considered Chiridota geminifera Dendy and Hindle to be a
junior synonym of Trochodota dunedinensis (Parker, 1881).

Scoliorhapis biopearli sp. nov.

Figure lc; tables 2, 3

Material examined. Holotype. South Shetland Is, 61.61°S 51.22°W,
1544 m, BAS stn EI-EBS-1, NMV F168633.

Diagnosis. Conical form, widest orally, tapered anally, dorsal
projecting over ventral orally, 6 mm long; 10 tentacles, number
of digits not evident; tentacle ossicles curved bracket-shaped
rods with bluntly spinous distal outer edges, 112-120 pm long;
body wall ossicles sigmoid hooks only, in close transverse
alignment, not clustered, hooks 168-184 pm long.

Colour (preserved). White to translucent.

Distribution. South Shetland Is, 1544 m.

Etymology. Named for the British Antarctic Survey BIOPEARL
expedition that collected and documented this specimen.

Remarks. This species is erected for a single, small BAS
BIOPEARL specimen with tentacle and body wall ossicles that
are eroded but retain distinguishable form and size. The presence
of hooks only, and not wheels, in the body wall distinguishes
this apodid specimen as a species of Scoliorhapis Clark. The
distinctive presence of 10 tentacles distinguishes this species
from the second new species of Soliorhapis Clark from
Antarctica described below. Scoliorhapis biopearli sp. nov. is
distinguished from the type species Scoliorhapis theeli (Heding)
from eastern Australia by the larger size of the hooks (see Table
3) and transverse arrangement of the hooks in the body wall, and
the spinous ends of the bracket- shaped tentacle ossicles.

Scoliorhapis massini sp. nov.

Figures Id, 9a, b; tables 2, 3

Material examined. Holotype. Scotia Sea, Shag Rocks, 53.63°S
40.91°W, 206 m, BAS stn SR-EBS-4, NMV F168635.

Other material. Falkland Is, 50°55'S 59°58'W, 118 m. Discovery
Expedition, William Scoresby stn 756, 10 Oct 1931, NHM 2010.105-
109 (5).

Diagnosis. Elongate, thin, 20 mm long; 12 tentacles, 3 pairs of
digits; tentacle ossicles slightly curved rods with central swelling
and bifurcate ends, up to 80 pm long; body wall ossicles sigmoid
hooks only, not clustered, hooks 88-104 pm long.

Colour (preserved). Reddish brown.

Distribution. Scotia Sea, Shag Rocks, 206 m.

Etymology. Named for Dr Claude Massin (Royal Belgian
Institute of Natural Sciences), with appreciation of a lifetime of
magnificent contribution to holothuroid systematics and in
particular here to Antarctic apodid studies.

Remarks. Scoliorhapis massini sp. nov. is erected for a single
BAS BIOPEARL specimen that is in good condition. It is
distinguished from the other two species of Scoliorhapis by the
presence of 12 tentacles. In addition it has tentacle ossicles that
are rods, not bracket-shaped, and the sigmoid hooks and tentacle
rods are smaller than in the other two species (see Table 3).

Herouard (1906) provided the very limited description of
12 tentacles and hook ossicles for specimens from the
Bellingshausen Sea that he determined to be Sigmodota
studeri (Theel). The few diagnostic details would fit
Scoliorhapis massini sp. nov.

The Discovery (1931) Falkland Is specimens occur at a
similar depth to the Shag Rocks specimen and are provisionally
referred to Scoliorhapis massini sp. nov. They are: brown; lack
wheels; longer (up to 50 mm long); have 12 tentacles, each
with more pairs of digits (up to 6); the ossicles are eroded; the
tentacle rods are of similar form but longer (120-160 um long);
the hooks longer (120-176 um long). The significantly larger
ossicles may reflect the larger specimen sizes or another new
species of Scoliorhapis.

Sigmodota Studer, 1876
Table 3

Sigmodota Studer, 1876: 454— Theel, 1886a: 16, 33— Ludwig,
1898: 73, 81-82 .-Ostergren, 1898: 117-118.-H. L. Clark, 1908:
121.— Dendy and Hindle, 1907: 113.— Heding, 1928: 277.— Pawson,
1964: 466.— Smirnov, 1998: 519.

Diagnosis. Taeniogyrinid genus with 12 plates in calcareous
ring, 12 tentacles, peltato-digitate, 4—7 pairs of digits per
tentacle, terminal pair longest; rods in tentacles; chiridotid
wheels and sigmoid hooks in body wall; teeth on the inner rim
of wheels in continuous series; wheels grouped into papillae,
hooks scattered in body wall; no miliary granules in longitudinal
muscles; 3-10 polian vesicles; ciliated funnels present.

Type species. Type species fixed here (under Article 70.3.2 of
the 1999 edition of the ICZN Code) as Chiridota contorta
Ludwig, 1875, misidentified as Holothuria (Fistularia)
purpurea Lesson, 1830 in the original monotypy designation
by Studer (1876).

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79

Figure 9. a, b, ossicles from the holotype of Scoliorhapis massini sp. nov. (Shag Rocks; NMV F168635). a, SEM of hook from the body wall; b,
SEM of rods from a tentacle, c, enhanced colour photo of preserved specimen of Taeniogyrus australianus (Stimpson, 1855), showing wheel
cluster papillae (large) and hook clusters (small) (27 mm long; Australia, Sydney Harbour, 1968; AM J16377).

80

P.M. O’Loughlin & D. VandenSpiegel

Other species. Taeniogyrus dubius H. L. Clark, 1921 (as S.
dubia)\ Taeniogyrus magnibaculus Massin and Heterier, 2004
(as S. magnibacula ).

Remarks. We follow Ludwig (1898) and H. L. Clark (1908) and
agree that Studer’s Sigmodota purpurea (Lesson, 1830) is a
junior synonym of Taeniogyrus contortus (Ludwig, 1875).
Studer (1876) erected his new genus Sigmodota for the species
Holothuria ( Fistularia ) purpurea Lesson, 1830 because of the
presence of sigmoid hooks in apodid specimens from the
Kerguelen Is and Magellanic region. He reported 12 tentacles,
and ignored the 10 tentacles in the species Holothuria
( Fistularia ) purpurea Lesson. As understood by Ludwig 1898
and H. L. Clark 1908 and as argued above by us in “Relevant
history of species misidentification” the material that Studer
examined had 12 tentacles, hooks and wheels.

H. L. Clark (1908) judged Sigmodota Studer to be a junior
synonym of Taeniogyrus Semper, 1867, which he diagnosed as
having: 10 or 12 peltato-digitate tentacles; 1 or several polian
vesicles; ciliated funnels not in stalked clusters; wheels in
papillae; large sigmoid hooks scattered in body wall; lacking
miliary granules. He included two species: Taeniogyrus
australianus (Stimpson, 1855), with 10 tentacles, hooks in
papillae, single polian vesicle; and Taeniogyrus contortus
(Ludwig, 1875), with 12 tentacles, hooks scattered, 6 or 7
polian vesicles.

As discussed above in “Ossicle clusters in generic
diagnosis” we reject ossicle aggregation in the body wall as a
generic diagnostic character, and judge that tentacle number is
a good generic character. Thus we raise Sigmodota Studer out
of synonymy with Taeniogyrus Semper, 1867 on the basis of
the type species Chiridota contorta Ludwig having 12
tentacles and the type species of Taeniogyrus, Chiridota
australiana Stimpson, having 10.

We examined specimens of Sigmodota contorta (Ludwig)
and Sigmodota magnibacula (Massin and Heterier) and found
that there were consistently 12 subequal plates in the calcareous
ring. Plates were fused, and joined beneath the tentacle bases
attached to the outside face of the ring. Each plate had a low
anterior projection and a shallow concave posterior indentation.

Sigmodota contorta (Ludwig, 1875)

Figure 10; tables 2, 3

Chiridota contorta Ludwig, 1875: 80-81, pi. 6, figs 6a-c.—
Lampert, 1885: 234.—’ Theel, 1886a: 16, 33, pi. 2 fig. 2.— Theel, 1886b:
20.— Lampert, 1889: 853-854.— Ludwig, 1891: 359.— Ostergren,
1897: 154.— Ludwig, 1897: 217-219.-Ludwig, 1898: 73-83, pi. 3 figs
37-42. -Perrier, 1905: 77-78.

Sigmodota purpurea .— Studer, 1876: 454.— Studer, 1879: 123.

Chiridota purpurea.— Bell, 1881: 101.— Lampert, 1885: 236.—
Lampert, 1886: 18-21, figs 17-20.-Ludwig, 1886: 29, 30.

Chiridota studeri Theel, 1886a: 33.— Lampert in Studer, 1889:
163, 283, 285, 308.

Sigmodota contorta— Ostergren, 1898: 118.— Sluiter, 1901: 134.

Sigmodota studeri.— Herouard, 1906: 15.

Taeniogyrus contortus.— H. L. Clark, 1908: 121-123, pi. 7 figs
8-13.— H. L. Clark, 1921: 165,-Ekman, 1925: 147-148.-Ekman,
1927: 416-417 (part Sigmodota magnibacula (Massin and Heterier,
2004)).— Heding, 1928: 311, fig. 66(l-9).-Deichman, 1947: 348-

349.— Pawson, 1964: 466, 467.— Pawson, 1969a: 126, 141.— Pawson,
1969b: 38, map 6.— Pawson, 1971: 289, figs 1, 2.— Arnaud, 1974:
585.— Hernandez, 1981: 164, figs lk, 1, 4d, e.-Gutt, 1988: 24 (part if
not all Sigmodota magnibacula (Massin and Heterier, 2004)).— Gutt,
1991: 324 (part if not all Sigmodota magnibacula (Massin and
Heterier, 2004)).-Massin, 1992: 311,-Branch et al„ 1993: 40, 56, 61,
65.— O’Loughlin, 2002: 298, 300, 301, tables 1, 2 (part Sigmodota
magnibacula (Massin and Heterier, 2004); see material examined).—
Massin and Heterier, 2004: 442, 443, table 1.— O’Loughlin, 2009: 2,
table 1.

Taeniogyrus contortus antarcticus .— Panning, 1936: 17, 18, fig.
8.— Pawson, 1969a: 141. (non Taeniogyrus antarcticus Heding, 1931)

Chiridota pisanii.— O’Loughlin, 2002: 298, tables 1, 4 (=
Sigmodota contorta ; non Chiridota pisanii Ludwig, 1886).

Material examined. South America, Argentina, Santa Cruz, east of
Grande Bay, Albatross Stn 2771, 51°34'S 68°00'W, 91 m, 1888, USNM
19826 (3); Chile, Inutil Bay, 53°35’S 69°45'W, 37-46 m, 1969, USNM
E33679 (13); 53°34'S 69°59'W, 82-91 m, 1970, USNM E33715 (9).

Falkland Is, 52°10’S 64°56’W, 150 m, Discovery Expedition,
William Scoresby stn 816, 14 Jan 1932, NHM 2010.55-62 (8); 54°00'S
64°58'W, 118 m, Discovery Expedition, William Scoresby stn 88, 6
Apr 1927, NHM 2010.63-68 (6); E Falkland Is, 75 m, Discovery
Expedition, William Scoresby stn 84, 24 Mar 1927, NHM 2010.69-70
(2); 296 m, Discovery Expedition, William Scoresby stn 773, 31 Oct
1931, NHM 2010.71-74 (4); Saldanka Bay, Discovery Expedition,
William Scoresby Marine Station 82, 6 Sep 1926, NHM 2010.75-84
(many).

Antarctic Ocean, Bouvet I., ICEFISH 2004 stn 80-BT42, 54.40°S
03.48°W, 159 m, NMV F104990 (1).

South Georgia, ICEFISH 2004 stn 38-BT18, 54.00°S 37.66°W, 46
m, NMV F104799 (1); 18-25 m, Discovery Expedition, William
Scoresby stn 25, 17 Dec 1926, NHM 2010.85-94 (11); RBINS IG 31
459 (1, SEM); 2 m, Discovery Expedition, William Scoresby stn 56, 14
Jan 1927, NHM 2010.95-96 (2); 179-235 m, Discovery Expedition,
stn 39, 25 Mar 1926, NHM 2010.97-98 (2); 0-100 m, Discovery
Expedition, stn 126, 19 Dec 1926, NHM 2010.99 (1); 26-83 m,
Discovery Expedition, William Scoresby stn 62, 19 Jan 1927, NHM
2010.100-103 (4); Leith Harbour, 22-55 m, Discovery Expedition, stn
1941, 29 Dec 1936, NHM 2010.104 (1).

S Shetland Is, 61.34°S 55.20° W, 204 m, 3 Dec 2006, BAS stn EI-
EBS-4, NMV F168628 (1).

S Orkney Is, 53.60°S 37.90° W, 503 m, BAS stn SG-EBS-3, NMV
F168632 (1); NMV F168642 (2 specimens as tissues); NHM 2010.35-
47 (13); RBINS IG 31 459 (2, SEM figures).

Indian Ocean, Heard I., ANARE, 52°41'-53T3 , S 72°56'-73°41'E,
120-228 m, NMV F84977 (1), F84978 (1), F84979 (1); Kerguelen Is,
49°33'S 69°49'E, 20 m, SAM K2384 (1); BANZARE stn 12, 49°28'S
70°04'E, 4-5 m, 1929, SAM K1839 (1); BANZARE stn 49, 49°30'S
69°48'E, 2-20 m, 1930, SAM K1840 (1).

Diagnosis. Sigmodotid species up to 50 mm long (preserved);
tentacles 12, 4-7 pairs of digits, longest distally; wheels in
distinct papillae, scattered over interradii, more numerous
dorsally and anteriorly, wheels 35-136 pm diameter; hooks
scattered, frequently about twice the size of the wheels, 136-
250 pm long; tentacle rods 80-184 pm long, significantly
shorter than hooks; up to 7 unequal polian vesicles; gonad
tubules dichotomously branched.

Type locality. Unknown (3 specimens).

Colour ( preserved ). Predominantly dark reddish-brown, rarely
grey and translucent.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

81

Figure 10. SEM images for specimens of Sigmodota contorta (Ludwig, 1875). a, plates from calcareous ring (specimen from Discovery
Expedition , William Scoresby stn 25, South Georgia, 18-27 m, RBINS IG 31 459). b, c, ossicles from S Shetland Is specimen (NMV F168628);
b, wheels from body wall; c, hook and wheel from body wall (left), rods from tentacle (right).

82

P.M. O’Loughlin & D. VandenSpiegel

Distribution. Western Antarctica, Bouvet I., 159 m; South
Georgia, 46 m; S Shetland Is, 204 m; S Orkney Is, 503 m.

South America, south of 42° in the west, S of 47°S in the
east (Pawson 1969a).

Indian Ocean, Heard I., 120-228 m; Kerguelen Is, 2-20 m,
116 m (Lampert 1889); Marion I., 125-132 m (Branch et al.
1993).

Java Sea, 82 m (Sluiter 1901).

Remarks. Specimens of the species Sigmodota contorta
(Ludwig) have been subjected to many misidentifications, as
discussed at the beginning of this work and as is evident in the
synonymy above.

Sluiter (1901) determined 3 Siboga specimens (stn 319) up
to 10 mm long from the Java Sea at 82 m as Sigmodota
contorta (Ludwig).

Herouard (1906) provided the very limited description of
12 tentacles and hook ossicles for specimens from the
Bellingshausen Sea that he determined to be Sigmodota
studeri (Theel). The few diagnostic details would fit
Scoliorhapis massini sp. nov. (above).

Panning (1936) examined specimens from South Georgia
that he judged to be Taeniogyrus antarcticus Heding, 1931. He
found the specimens close to Taeniogyrus contortus (Ludwig),
and made the Heding species a subspecies ( Taeniogyrus
contortus antarcticus Heding). Based on tentacle number (12)
and ossicle sizes, we judge that Panning (1936) was in fact
examining specimens of Sigmodota contorta (Ludwig).

Massin and Heterier (2004) erected their species
Taeniogyrus magnibaculus for specimens from the Weddell
Sea subsequent to the work of Cherbonnier (1974), Gutt (1988,
1991), and O’Loughlin (2002) who reported Taeniogyrus
contortus (Ludwig) from the regions of Terre Adelie, the
Weddell Sea and Prydz Bay respectively. The Prydz Bay
material has been redetermined as Taeniogyrus magnibaculus,
which has also been found by us in the Ross Sea and off
Wilkes Land. Taeniogyrus contortus (Ludwig) has not been
found on coastal eastern Antarctica, and it is anticipated that
the Cherbonnier reference was to Taeniogyrus magnibaculus
and this synonymy is given below. Likewise Gutt (1988, 1991)
reported Taeniogyrus contortus from the Weddell Sea. We
anticipate that some or possibly all of his material was
Taeniogyrus magnibaculus. Taeniogyrus contortus and
Taeniogyrus magnibaculus are generally allopatric, except in
the South Orkney Is.

The mistaken identification by O’Loughlin (2002) of
Heard I. specimens as Chiridota pisanii Ludwig, 1886 was
corrected by Bohn in Altnoder et al. (2007), and discussed by
O’Loughlin (2009). The specimens are listed here as
Taeniogyrus contortus (Ludwig).

Sigmodota dubia (H. L. Clark, 1921)

Table 3

Taeniogyrus species Fisher, 1907: 735, pi. 82 fig. 2.

Taeniogyrus dubius H. L. Clark, 1921: 166 (key and note).—
Heding, 1928: 311 — Massin and Heterier, 2004: 442, 443, table 1.

Distribution. Hawaiian Is, Oahu I., 403-470 m.

Remarks. Fisher (1907) reported 12 tentacles, 10 polian vesicles
of unequal size, and aggregations of wheels. His specimen and
ossicle meaurements are given in Table 3. H. L. Clark (1921)
named the species. Fisher (1907) noted that this species was
close to Taeniogyrus contortus (Ludwig), and distinguished it
by the larger number of polian vesicles (10, unequal) and size
and form of the sigmoid ossicles. We found the wheels of
Sigmodota dubia as reported by Fisher (1907) to be significantly
larger than our measurements for Sigmodota contorta, but the
hooks not significantly smaller, as was judged by Fisher (1907)
(see Table 3).

Sigmodota magnibacula (Massin and Heterier, 2004)

Figure le, 11; tables 2, 3

Taeniogyrus contortus— Ekman, 1927: 416-417.— Cherbonnier,
1974: 610.— Gutt, 1988: 24.— Gutt, 1991: 324. (all or part probably non
Taeniogyrus contortus (Ludwig, 1875))

Taeniogyrus cf. contortus.— O’Loughlin et al., 1994: 553, 554.

Taeniogyrus magnibaculus Massin and Heterier, 2004: 441-444,
figs 1, 2, table 1.

Material examined. Scotia Sea, South Orkney Is, 60.82°S 46.49°W,
216 m, BAS stn PB-EBS-4, 2006, NMV F168629 (1); NMV F168639
(1 specimen as tissue sample MOL AF 798); NHM 2010.1-4 (4);
RBINS IG 31 459 (1, SEM figures); 60°50'S 44°30'W, 172 m, US
AMLR 2009 stn 16-31, NMV F168840 (13); 61°13'S 45°56'W, 240 m,
US AMLR stn 41-46, NMV F168841 (8).

Eastern Antarctica, Ross Sea, McMurdo Sound, 366 m. Terra
Nova stn 348, 13 Feb 1912, NHM 1932.8.11.216-217 (2); NHM
1932.8.11.218 (1); Cape Adare, 71°53'S 170°11'E, 220 m, BIOROSS stn
TAN0402/94, NIWA 61060 (2); Ross Sea, 74°43’S 164°08'E, 140 m,
MNA 2440 (1); 74°43'S 164°07'E, 120 m, MNA 2446 (10); 74°45’S
164°15'E, 219 m, MNA 2462 (1); IPY-CAML stn TAN0802/26,
74°58'S 170°27'E, 285 m, NIWA 35724 (4); stn TAN0802/31, 74°59'S
170°27'E, 283 m, NIWA 35776 (1).

Wilkes Land, 66°18'S 110°32'E, 101 m, USARP 1961, USNM
E33725 (25).

Prydz Bay, ANARE 1987, stn 51, 525 m, NMV F76847 (1); stn 22,
165 m, NMV F168862 (1); stn 38, 312 m, NMV F168863 (3); N end of
Fram Bank, 67°05'S 68°59'E, 216 m, ANARE AA93-131, NMV
F69100 (2).

MacRobertson Shelf, 67°16'S 65°25'E, 121 m, ANARE AA93-
127, NMV F68691 (20); 66°55'S 62°32'E, 113 m, AA93-124, NMV
F69099 (2); Mawson Base, 8 m, ANARE 1972, NMV F168839 (1).

Diagnosis. Sigmodotid species up to 105 mm long (preserved);
12 peltato-digitate tentacles, 6 pairs of digits per tentacle, apical
pair largest, decreasing in size towards base of tentacle trunk;
rows of interradial off-white papillae comprising clusters of
chiridotid wheels; wheels 56-200 pm diameter; hooks scattered
in body wall, 128-216 pm long, blunt spines on outer surface of
hook variably present and of variable size; tentacle rods
irregularly straight, slightly swollen mid-rod, tapered, short
branches distally, 170-320 pm long; calcareous ring and ossicles
sometimes absent in largest specimens; 3 polian vesicles.

Colour (preserved). Body wall and tentacles dark red-brown to
violet-brown epidermally, dark grey underneath.

Distribution. Antarctic, Weddell Sea, 72°29'S 26°57'W, 226 m
(Massin and Heterier 2004); South Orkney Is, 172-240 m;

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83

Figure 11. SEM images for specimens of Sigmodota magnibacula (Massin and Heterier, 2004). a, b, plates from calcareous ring (Prydz Bay;
RBINS IG 31 459 ex NMV F68691); c, wheels from body wall (S Orkney Is; RBINS IG 31 459 ex NMV F168842); d, hooks from body wall (S
Orkney Is; RBINS IG 31 459); e, rods from tentacles (S Orkney Is; RBINS IG 31 459 ex NMV F168842).

84

P.M. O’Loughlin & D. VandenSpiegel

Ross Sea, 120-366 m; Terre Adelie, 170-250 m (Cherbonnier
1974); Wilkes Land, 101 m; Prydz Bay, 165-525 m;
MacRobertson Shelf, 8-121 m.

Remarks. As can be seen in Table 3 the maximum size of
specimens, and length and form of the tentacle rods, are good
diagnostic characters for Sigmodota magnibacula (Massin and
Heterier) . Diameter of wheels and length of hooks are comparable,
but tentacle rods much larger. As discussed above in the Remarks
under Sigmodota contorta (Ludwig), the eastern Antarctic
material of Cherbonnier (1974) is judged to be Sigmodota
magnibacula, as is some or all of the Weddell Sea material of
Gutt (1988, 1991). In discussing Taenio gyrus contortus (Ludwig)
from the “Wmterstation”, Ekman (1927) made mention of what
is probably this species. Massin and Heterier (2004) noted that
this species was “living on the spines of cidarid echinoids”. We
have not seen evidence of such an association. The US AMLR
specimens were found inside Demospongiae.

Taeniogyrus Semper, 1867
Table 3

Taeniogyrus Semper, 1867: 23.— Smirnov, 1998: 519.

(For synonymies see Ludwig 1898 and Pawson 1964).

Trochodota Ludwig, 1891: 358.— Pawson, 1968: 24.— Pawson,
1970: 46.— Smirnov, 1998: 519.

(For synonymies see Ludwig 1898 and Pawson 1964).

Type species of Taeniogyrus Semper, 1867. Chiridota
australiana Stimpson, 1855 (monotypy).

Type species of Trochodota Ludwig, 1891. Rowe (in Rowe and
Gates 1995) proposed that “a case needs to be put to the ICZN
to establish Holothuria (Fistularia) purpurea Lesson, 1830 as
type species of Trochodota Ludwig, 1891. Type species:
Chiridota studeri Theel, 1886, sensu Ludwig, 1891 -Holothuria
(Fistularia) purpurea Lesson, 1830 (not Theel,
1886 -Taeniogyrus contortus (Ludwig, 1875)) by subsequent
designation, Rowe, F. W. E., this work.” The type species
proposed by Rowe is fixed here (under Article 69.2.4 of the
1999 edition of the ICZN Code) as Holothuria ( Fistularia )
purpurea Lesson, 1830, misidentified as Chiridota studeri
Theel, 1886a in the original inclusion of two species in his new
genus by Ludwig (1891).

Other included species. Taeniogyrus antarcticus Heding, 1931;
Chiridota benhami Dendy, 1909; Taeniogyrus cidaridis
Ohshima, 1915; Taeniogyrus clavus Heding, 1928; Taeniogyrus
dayi Cherbonnier, 1952; Taeniogyrus dendyi Mortensen, 1925;
Trochodota diasema H. L. Clark, 1921 (as T. diasemus)-,
Chiridota dunedinensis Parker, 1881; Taeniogyrus
heterosigmus Heding, 1931; Trochodota inexpectata Smirnov,
1989 (as T. inexpectatus)-, Chiridota japonica von Mareneller,
1881 (as T. japonicus)-, Taeniogyrus keiensis Heding, 1928;
Trochodota maculata H. L. Clark, 1921 (as T. maculatus)-,
Trochodota neocaladonica Smirnov, 1997 (as T.
neocaladonicus)\ Taeniogyrus papillis O’Loughlin, 2007 (in
O’Loughlin and VandenSpiegel); Taeniogyrus prydzi sp. nov.;
Holothuria (Fistularia) purpurea Lesson, 1830 (as T.
purpureus)-, Trochodota roebucki Joshua, 1914; Trochodota

rosea Ohshima, 1914 (as T. roseus)\ Taeniogyrus tantulus
O’Loughlin, 2007 (in O’Loughlin and VandenSpiegel);
Chiridota venusta Semon, 1887 (as T. venustus).

Diagnosis. Taeniogyrinid genus with 10 peltato- digitate
tentacles; 4-8 pairs of digits, terminal pair longest; rods in
tentacles; chiridotid wheels and sigmoid hooks in body wall;
teeth on the inner rim of wheels in continuous series; wheels
and hooks variably grouped or scattered in body wall; no
miliary granules in longitudinal muscles; 1-12 polian vesicles;
ciliated funnels present.

Remarks. The succession of authors who have expressed
dissatisfaction with the genera Taeniogyrus Semper and
Trochodota Ludwig is mentioned in the Introduction. As
discussed at the beginning of this paper and stated in the
Remarks under Sigmodota we reject ossicle aggregation in the
body wall as a generic diagnostic character, and judge that
tentacle number is a good generic character. We thus raise
Sigmodota Studer (taeniogyrinid species with 12 tentacles) out
of synonymy with Taeniogyrus Semper (10 tentacles), and
synonymise Trochodota Ludwig with Taeniogyrus Semper
(taeniogyrinids with 10 tentacles).

Taeniogyrus antarcticus Heding, 1931
Figures If, 12; tables 2, 3

Taeniogyrus antarcticus Heding, 1931: 685-691, fig. 15(1—12). —
Pawson, 1969a: 141.— Massin and Heterier, 2004: 442-443, table 1.

Material examined. Western Antarctica, Scotia Sea, S Orkney Is,
60.82°S 46.49°W, 216 m, BAS stn PB-EBS-4, 18 Mar 2006, NMV
F168630 (1); NMV F168640 (2 as tissue samples, codes MOL AF799,
800); NMV F168641 (2 as tissue samples, codes MOL AF801, 802);
NHM 2010.5-14 (10); NHM 2010.15-34 (20); RBINS IG 31 459 (2
whole, SEM figures).

Shag Rocks, 53.63°S 40.91° W, 206 m, BAS stn SR-EBS-4, 11
Apr 2006, NMV F168636 (1); RBINS IG 31 459 (1 whole, SEM
figures); NHM 2010.51-52 (2).

Diagnosis. Up to 15 mm long; 10 tentacles; 5 pairs of digits,
terminal/distal pair longest; 3-9 polian vesicles; few long, thin
ciliated funnels with long membranaceous collar, short funnel,
and distinct peduncle; chiridotid wheels 40-80 pm diameter (no
data in Heding 1931; 40-80 pm, S Orkney Is; 48-64 pm, Shag
Rocks), wheels in a few small clusters dorsally; hooks scattered
in body wall, 64—208 pm long (172-200 pm, Heding 1931 fig.
15; 80-208 pm, S Orkney Is; 64—168 pm. Shag Rocks); tentacle
rods straight to slightly bent to bracket-shaped, dichotomously
branching ends, rare blunt spines along rods, rare rounded hub
in mid-rod, 64—128 pm long (83-103 pm, Heding 1931 fig. 15;
80-128 pm, S Orkney Is; 64—72 pm, Shag Rocks).

Colour (preserved). Pale yellow-brown.

Distribution. Western Antarctica, Scotia Sea, South Orkney Is,
South Georgia, Shag Rocks, 206-216 m (0-15 m for South
Georgia in Massin and Heterier 2004).

Remarks. Heding (1931) erected his new species for many
specimens from South Georgia, by distinguishing them within
a discussion of Taeniogyrus contortus (Ludwig). Massin and

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Figure 12. SEM images for specimen of Taeniogyrus antarcticus Heding, 1931 (S Orkney Is; RBINS IG 31 459). a, plates from the calcareous
ring; b, hooks from the body wall; c, wheels from the body wall; d, rods from the tentacles.

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P.M. O’Loughlin & D. VandenSpiegel

Heterier (2004 table) listed 12 tentacles for Taeniogyrus
antarcticus Heding, but in the last sentence of his discussion
Heding (1931) noted that Taeniogyrus contortus specimens all
had 12 tentacles “apart from this example” (referring to his new
species Taeniogyrus antarcticus ).

Panning (1936) considered Taeniogyrus antarcticus
Heding to be at best a subspecies of Taeniogyrus contortus
Ludwig. Pawson (1969a) followed Panning (1936). On the
basis of tentacle number we disagree. Based on tentacle
number and ossicle sizes we judge that the specimens from
South Georgia that were examined by Panning (1936) were
Sigmodota contorta (Ludwig).

There are significant differences in the measurement sizes
available for ossicles from different localities in the Scotia
Sea, as can be seen in Table 3. The variations may be the result
of size of specimen sampled or location of the sample taken
from the body wall. Or the variations may indicate cryptic
speciation. In this study we found that for this species the
limited data available indicate that maximum ossicle size
increases with specimen size. The type locality for Taeniogyrus
antarcticus Heding is South Georgia.

Taeniogyrus antarcticus Heding, 1931 is the sole western
Antarctic species of Taeniogyrus. A new species of
Taeniogyrus from eastern Antarctica is described below, with
significantly longer sigmoid hooks and a single polian vesicle.

Taeniogyrus australianus (Stimpson, 1855)

Figures 9c, 13; table 3

Chiridota australiana Stimpson, 1855: 386.— Lampert, 1885:
230.— Ludwig, 1898: 82-83.

Taeniogyrus australianus— Semper, 1867: 23.— H. L. Clark,
1908: 122.-H.L. Clark, 1921: 166,-Heding, 1928: 310,311,315-316,
fig. 67(9-16).— H. L. Clark, 1946: 459.— Rowe (in Rowe and Gates),
1995: 267.

Sigmodota australiana Ostergren, 1898: 118.

Material examined. Australia, Sydney, Collaroy, Long Reef, in sand,
AM J20086 (2); same lot, RBINS IG 31 459 (1 whole for SEM images);
AM J12542 (1); Middle Harbour, AM J16377 (1); Lord Howe I., AM
J16373 (1); Heron I., AM J19570 (11).

Diagnosis. Taeniogyrid species up to 95 mm long (preserved);
10 peltato-digitate tentacles, 6-7 pairs of digits per tentacle,
longest distally; single madreporite at oral end of dorsal
mesentery, mushroom-like head; single ventral elongate sac-like
polian vesicle; 2 series of crowded ciliated funnels, on left edge
of left ventral radial muscle, and right edge of mid-ventral radial
muscle; chiridotid wheels in large to small discrete papillae,
wheels 48-88 pm diameter; sigmoid hooks in small papillae, not
scattered, hooks 120-136 p m long; tentacle rods thickened
centrally, slightly curved, bifurcate ends, 72-104 pm long.

Distribution. Eastern Australia (Mooloolaba in Queensland to
Ulladulla in New South Wales; Great Barrier Reef; Heron I.;
Lord Howe I.; 0-15 m (Rowe and Gates 1995).

Remarks. Taeniogyrus australianus (Stimpson) is not an
Antarctic apodid species but has not been fully illustrated in
previous works and is included here as it is the type species of
the revised genus Taeniogyrus, and its identity was confused

by Theel (1886a). He described two specimens of Chiridota
australiana Stimpson from Port William (with uncertainty
whether New Zealand or Falkland Is) that were 35 mm long,
with 10 tentacles, each with 4 pairs of digits, a single polian
vesicle, wheels 140 pm diameter with no evidence of clustering,
hooks 140 pm long. We agree with Ludwig (1898) that this
evidence points to Taeniogyrus purpureus (Lesson) and the
Falkland Is.

Taeniogyrus maculatus (H. L. Clark, 1921)

Figure 14; table 3

Trochodota maculata H. L. Clark, 1921: 163, pi. 36 figs 14—21. —
H. L. Clark, 1946: 460,-Rowe, 1976: 203-205, table l.-Rowe (in
Rowe and Gates) 1995: 268.— Smirnov, 1997: 16.

Material examined. E Australia, New South Wales, Newcastle,
Swansea Channel, 3 m, 30 Aug 1988, AM J21895 (9); same lot, RBINS
IG 31 459 (1 whole for SEM images).

Diagnosis. Up to 21 mm long (preserved; 26 mm long in Clark
1921); body wall papillate, semi -translucent; tentacles 10, 3
pairs of digits (4—5 in Clark 1921); curved to slightly bracket-
shaped tentacle rods; wheels with 6 spokes, spokes broad at rim,
narrow at hub, teeth on inner rim of wheels in continuous series;
sigmoid hooks with fine spinelets on outer surface of projecting
pointed hook, hooks scattered in body wall (hooks in small
groups and scattered in Clark 1921); ciliated funnels along base
of mid-dorsal mesentery; single sac-like polian vesicle; gonad
comprises 2 elongate sacs (ossicle measurements in Table 3).

Colour (preserved). Pale to dark reddish-brown. Colour live is
pink with numerous minute dark spots (Clark 1921).

Type locality. N Australia, Torres Strait, Murray Is, Mer, reef flat.

Distribution. Tropical Australia, 0-20 m (Rowe in Rowe and
Gates 1995).

Remarks. Taeniogyrus maculatus (H. L. Clark) is not an
Antarctic apodid species but is included here to clarify its
generic assignment. The figure of a wheel in H. L. Clark (1921)
indicates that the wheel was viewed from the outside and the
continuity of teeth around the inner rim obscured by the spokes.
This led H. L. Clark to wrongly conclude that the distribution of
the teeth was discontinuous. Rowe (1976) followed H. L. Clark
(1921). In this study all wheels have continuous series of teeth.

Taeniogyrus prydzi sp. nov.

Figure 15; tables 2, 3

Taeniogyrus sp. MoV 2007 O’Loughlin et al., 1994: 553, 554.
Taeniogyrus sp. MoV 2010 O’Loughlin et al., 1994: 553, 554.

Material examined. Holotype. Eastern Antarctica, MacRobertson
Shelf, stn ANARE AA 93-124, 66°55'S 62°32'E, 113 m, M.
O’Loughlin, NMV F68690.

Paratypes. MacRobertson Shelf, Edge of Nielsen Basin, stn
ANARE AA 93-127, 67°16'S 65°25'E, 109-121 m, NMV F69120 (1);
NMV F68689 (1, 2 pieces); Prydz Bay, N of Fram Bank, stn ANARE
AA 93-142, 66°49'S 70°25'E, 795-830 m, NMV F68688 (3); NMV
F76096 (1).

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87

Figure 13. SEM images for specimen of Taeniogyrus australianus (Stimpson, 1855) (Long Reef, Collaroy, Sydney; RBINS IG 31 459 ex AM
J20086). a, plates from calcareous ring; b, wheels and hook from body wall; c, wheels from body wall; d, hooks from body wall; e, rods from
tentacles.

P.M. O’Loughlin & D. VandenSpiegel

Figure 14. SEM images for specimen of Taeniogyrus maculatus (H. L. Clark, 1921) (Swansea Channel, E Australia; RBINS IG 31 459 ex AM
J21895). a, calcareous ring plates (with hook contaminants); b, wheels from body wall; c, hooks from body wall, d, montage photographs of
ossicles of Taeniogyrus purpureus (Lesson, 1830) (Strait of Magellan; USNM 1004241); rods from tentacle, and wheels and hook from body wall.

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

89

Diagnosis. Up to 50 mm long; 10 tentacles, 6 pairs of digits per
tentacle, digits increasing in length distally; tentacle ossicles
rods, thick, predominantly straight, short blunt branches
distally, rare centrally, rods 136-152 pm long; chiridotid wheels
with continuous series of teeth on inner rim; single dorsal
series of wheel aggregations, a few irregularly scattered wheel
clusters dorso-laterally; no wheel aggregations ventrally;
chiridotid wheels up to 90 pm diameter; sigmoid hooks
scattered in body wall, 232-272 pm long; single polian vesicle;
numerous gonad tubules arising from a common hub; single
ventral interradial series of ciliated funnels.

Distribution. Eastern Antarctica, MacRobertson Shelf, 109-
121 m; Prydz Bay Channel, outfall slope, 795-830 m.

Etymology. Named for Prydz Bay in eastern Antarctica near
the type locality.

Remarks. The ossicles in the type specimens show evidence of
erosion but are adequate for systematic assessment. The eastern
Antarctic Taeniogyrus prydzi sp. nov. is distinguished from the
western Antarctic Taeniogyrus antarcticus Heding by the
single polian vesicle and significantly longer sigmoid hooks
(see Table 3).

Taeniogyrus purpureus (Lesson, 1830)

Figure 14d; tables 2, 3

Holotliuria (Fistularia) purpurea Lesson, 1830: 155-156, pi. 52,
fig. 2— Rowe (in Rowe and Gates), 1995: 268.

Chiridota purpurea .— Jager, 1833: 16.— Brandt, 1835: 259.—
Dujardin and Hupe, 1862: 616.— Semper, 1867: 23.— Theel, 1886a:
35-36.— Lamp ert 1889: 851.

Chiridota australiana.— Theel, 1886a: 16 (non Chiridota

australiana Stimpson, 1855).

Chiridota studeri.— Lampert 1889: 849-850, pi. 24 fig. 12.—
Ludwig 1891: 359-360 (non Chiridota studeri Theel, 1886).

Trochodota purpurea .—Ludwig, 1898: 83-87, pi. 3 figs 43-45.—
Perrier, 1905: 76-77.-H. L. Clark, 1908: 123-124.-H. L. Clark,
1921: 166.— Pawson, 1964: 466,-Ekman, 1925: 149-150.-

Deichmann, 1947: 349.— Pawson, 1969a: 141.— Pawson, 1969b: 38,
map 6.— Rowe, 1976: 203-205, table 1.— Hernandez, 1981: 164, 166,
figs li, j, 4b, c.— Rowe (in Rowe and Gates), 1995: 268.— Smirnov,
1997: 16.

Material examined. Strait of Magellan, mouth of strait, 64 m, USNM
1004241 (2); Isla Bertrand, Puerto Grande, intertidal. Royal Society
1958 Expedition, USNM E16375 (1, no ossicles remaining).

Diagnosis (Pawson 1964). Tentacles 10, each with 2-6 pairs of
digits; wheels 130-180 pm diameter (80-144 pm this work,
USNM 1004241), scattered in body wall; sigmoid hooks 120-
130 pm long (120-160 pm this work, USNM 1004241),
scattered in body wall; tentacle rods average 78 pm long (56-
120 pm this work, USNM 1004241), bracket-shaped (rare in
this work), with dichotomously branching ends; colour
commonly purple.

Type locality. Puerto Soledad (Port Solitude), Falkland Is.

Distribution. South America, Strait of Magellan, 64 m (this
work); Falkland Is (type locality).

Remarks. In his key to the species of Trochodota, H. L. Clark
(1921) implied that Trochodota purpurea has discontinuous
series of teeth on the inner rim of the wheels. Smirnov (1997)
observed continuous series of teeth, and pointed out that
Hernandez (1981) illustrated a wheel of Trochodota purpurea
with continuous teeth. We confirm that observation here (Fig.
14d). Ossicle dimensions are given in Table 3. Holotliuria
(Fistularia) purpurea Lesson, 1830 is fixed in this work (above)
as type species of Trochodota Ludwig, 1891, but we then judge
Trochodota to be a junior synonym of Taeniogyrus Semper, 1867.

It is established above that Chiridota studeri Theel is a
junior synonym of Sigmodota contorta (Ludwig), and this is
listed in that synonymy. Lampert (1889, pp. 849, 850) discussed
material from the Strait of Magellan that he referred to Chiridota
studeri Theel, but his description of 10 tentacles, hooks and
wheels not in papillae clearly refers to Taeniogyrus purpureus
(Lesson), and this reference is listed in the synonym here.

We agree above with Ludwig (1898) that the Theel (1886a)
specimens of Chiridota australiana Stimpson were
Taeniogyrus purpureus (Lesson).

Subfamily Chiridotinae Ostergren, 1898 (sensu Smirnov 1998)

Diagnosis (Smirnov, 1998). Chiridotidae with 12 or 18
tentacles. Body wall ossicles wheels of chiridotid type gathered
into papillae, and/or rods. Radial plates of calcareous ring
perforated or with deep notch in anterior face for passage of
nerves. There are 4-30 polian vesicles.

Genera (Smirnov 1998). Chiridota Eschscholtz, 1829; Paradota
Heding (in Ludwig and Heding), 1935; Polycheira H. L. Clark,
1908.

Chiridota Eschscholtz, 1829
Synonymy. See Pawson 1964.

Diagnosis (Pawson 1964). Tentacles 12, digits 3-10 pairs,
terminal pair longest; polian vesicles 3-20; ossicles six-spoked
wheels collected into papillae with varying numbers of wheels
of diverse sizes; small curved rods with enlarged ends may be
present; minute miliary granules often present in longitudinal
muscles; lacking sigmoid hooks.

Type species. Chiridota discolor Eschscholtz, 1829

Type locality. Sitka; Sea of Okhotsk.

Chiridota pisanii Ludwig, 1886
Table 2

Chiridota purpurea.— Theel, 1886: 15, pi. 2 fig. 1 (= Chiridota
pisanii Ludwig, 1886; non Holotliuria ( Fistularia ) purpurea Lesson,
1830; see Ludwig 1898).

(For complete synonymy see Bohn in Altnoder et al. 2007).

Material examined. S Atlantic Ocean, Burdwood Bank, ICEFISH 2004
stn 1-OT1, 54.22°S 59.84°W, 93 m NMV FI 06959 (1); ICEFISH 2004
stn 1-OT2, 54.22°S 59.83°W, 93 m NMV F106963 (4); Patagonia,
52°12'S 67°19'W, 95 m. Discovery Expedition, William Scoresby stn
750, 19 Sept 1931, NHM 2010.110 (1, wheel cluster only); Argentina,

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P.M. O’Loughlin & D. VandenSpiegel

Figure 15. SEM images for holotype of Taeniogyrus prydzi sp. nov. (Prydz Bay; NMV F68690). a, hooks from body wall; b, wheels from body
wall; c, rods from tentacles.

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91

Tierra del Fuego, 54°00'S 67°24'W, 0 m, 1999, NMV F86016 (2); Cape
Horn, 56°20’S 67°10'W, 121 m. Discovery Expedition, William
Scoresby stn 388, 16 Apr 1930, NHM 2010.111 (1).

Diagnosis ( from Bohn in Altnoder et al. 2007). Up to 68 mm
long (130 mm in Ludwig 1898); tentacles 12, 4-7 pair of digits
per tentacle, longest distal ly, tentacle rods with variably
branched ends, sometimes central hub, 16-69 pm long;
calcareous ring with 5 radial 7 interradial plates, all radial
plates perforated for passage of nerve; polian vesicles 4-11 (-16
in Theel 1886); single ciliated urns at base of mesenteries,
numerous in mid-dorsal and left dorsal interradius, sparse in
right ventral interradius; chiridotid wheels with serrations on
inner side continuous, gathered into papillae in single series in
dorsal interradii, inconspicuous or lacking in ventral interradii,
43-147 pm diameter; miliary granules in longitudinal muscles
14-49 pm long.

Colour (preserved). Off-white to pink to reddish-brown.

Distribution. Pacific and Atlantic coasts of southern South
America (south of 42°S), Falkland Is, 0-102 m (Bohn in
Altnoder et al. 2007); Burdwood Bank, 93 m (this work); Cape
Horn, 121 m (this work).

Remarks. Chiridota pisanii Ludwig has not been reported
south of the Polar Front, and is not an Antarctic species. Bohn
(in Altnoder et al. 2007) has provided a comprehensive
systematic treatment with illustrations. He queried the identity
of the Challenger Falkland Is specimens determined by Theel
(1886) as Chiridota purpurea, and subsequently judged by
Ludwig (1898) to be Chiridota pisanii, since the wheel sizes
were larger (140-160 pm diameter) than his measurements for
Chiridota pisanii (43-147 pm diameter). Our ossicle
measurements for Burdwood Bank specimens are: wheels 80-
144 pm diameter; tentacle rods 48-80 pm long.

O’Loughlin (2002) initially failed to find hook ossicles in
specimens from Heard I. and identified them as Chiridota
pisanii. Hook ossicles were subsequently found, and the
material re-determined as Taeniogyrus contortus (see
O’Loughlin 2009).

Paradota Heding, 1935

Paradota Heding (in Ludwig and Heding), 1935: 150-151, fig.
14.— Gutt, 1990: 125-126.— Massin, 1992: 321-323.— Smirnov,
1998: 520.

Diagnosis. Tentacles 12, peltato-digitate, with 5 to 7 pairs of
digits, longest distally; rod ossicles in tentacles; lacking ossicles
in the body wall; 1 to 11 polian vesicles.

Type species. Achiridota ingolfi Heding, 1935.

Other species. Paradota weddellensis Gutt, 1990; Paradota
marionensis Massin, 1992.

Remarks. Heding (in Ludwig and Heding 1935) erected
Paradota for family Chiridotidae . Gutt ( 1 990) referred Paradota
to the Synpatidae Burmeister, 1837. Smirnov (1998) judged that
the morphology of genus Paradota was close to that of genus
Chiridota and referred Paradota to subfamily Chiridotinae.

Paradota weddellensis Gutt, 1990
Figure 16; table 2

Paradota weddellensis Gutt, 1990: 125-126, figs 11-14, table 3.—
Massin, 1992: 322-323.— O’Loughlin et al. 2009: table 1.

Material examined. Scotia Sea, South Orkney Is, US AMLR stn 104,
63°13.92'S 59°27.47'W, 759 m, NMV F168842 (5); South Sandwich Is,
351-393 m, USNM E49614 (20+); South Shetland Is, 59 m, USNM
E49620 (1).

Antarctic Peninsula, Joinville I., 265 m, USNM E49619 (1);
Palmer Archipelago, 126 m, USNM E49616 (1).

Ross Sea, NIWA expedition 2001, 66°49’S 162°37'E, 292 m,
NIWA 61095 (1); stn TAN0602/448, 66°56'S 162°57'E, 85 m, NIWA
49800 (1); NZ IPY-CAML stn TAN0802/272, 66°96'S 170°93’E, 658
m, NIWA 38869 (1).

Prydz Bay, ANARE 1987 stn 7, 68°40'S 77°12'E, 505-578 m,
NMV F76845 (1); NMV F76846 (1).

Heard I., ANARE AA92-01, 52°57’S 73°21'E, 159-176 m, NMV
F84977 (1); AA92-06, 53°13'S 73°40'E, 120 m, NMV F84979 (1);
AA92-08, 52°41’S 72°56'E, 215 m, NMV F84978 (1).

Diagnosis ( after Gutt 1990). Chiridotinid species up to 180
mm long; body wall thick; tentacles 12, peltato-digitate, 5-7
pairs of digits; calcareous ring lacking anterior and posterior
projections; 4 polian vesicles; tentacles with irregular rod
ossicles, many slightly bracket-shaped, some with branched
ends, some with projections mid-rod; body wall lacking
ossicles; longitudinal muscles with rare miliary granules.

Distribution. Circum-antarctic, south of Polar Front, 59-1191
m; Weddell Sea, 225-655 m (Gutt 1990); Scotia Sea, 59-759
m; Antarctic Peninsula, 126-265 m; Bellingshausen Sea,
Antarctic Peninsula, Peter I Island, 97-1191 m (O’Loughlin et
al. 2009); Ross Sea, 85-658 m; Prydz Bay, 505-578 m; Heard
I., 120-215 m.

Remarks. Paradota weddellensis Gutt is a large apodid with a
widespread polar distribution south of the Antarctic
Convergence. Paradota marionensis Massin, 1992 occurs
north of the Convergence at Marion and Prince Edward Is
(Massin 1992). We found rare miliary granules up to 40 pm
long in Ross Sea material.

Acknowledgments

We are grateful to the following for their gracious assistance:
Andrew Cabrinovic (provision of NHM registration numbers
and permission to donate specimens to RBINS for SEM study);
Ben Boonen (preparation of figures); Caroline Harding
(photography); Edward Tsyrlin (translation of Russian
literature); Gary Poore and Frank Rowe (helpful dialogue
around systematic issues); Gustav Paulay (assistance with
literature); Igor Smirnov (translation of Russian literature);
Katrin Linse (facilitation with BAS BIOPEARL loan and data
and donation of specimens); Leon Altoff and Audrey Falconer
(photography); Mike Reich (assistance with literature, and
translation of German literature); Niki Davey and Kareen
Schnabel (facilitation with NIWA specimen loan, and NIWA
Ross Sea specimens data); Olga Hionis (assistance with
literature); Rafael Bendayan de Moura (assistance with
literature); Steve Keable and Kate Attwood (facilitation with

92

P.M. O’Loughlin & D. VandenSpiegel

Figure 16. SEM images for specimens of Paradota weddellensis Gutt, 1990. a, calcareous ring plates: (from top left) anterior view, posterior view,
internal view, external view, section view (South Orkney Is; NMV F168842); b, rods from tentacles (Prydz Bay; NMV F76846).

A revision of Antarctic and some Indo-Pacific apodid sea cucumbers (Echinodermata: Holothuroidea: Apodida)

93

AM loan and donation of specimens); Susie Lockhart
(facilitation with US AMLR specimens and donations and
data); Toshihiko Fujita (assistance with literature). We
acknowledge the thorough work of the BIOPEARL scientists
who found the very small myriotrochid and taeniogyrid
specimens. We are appreciative of assistance with facilities
provided by Museum Victoria and the Smithsonian Institution,
and in particular with help provided by Chris Rowley (NMV)
and by David Pawson and Jennifer Hammock (SI). We are
grateful for access to specimens of the New Zealand
International Polar Year-Census of Antarctic Marine Life
Project. We are most grateful for the very helpful corrections
and suggestions made by Claude Massin, David Pawson and
Frank Rowe.

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l—l — I t I I

50 mm

Memoirs of Museum Victoria

Volume 67 31 December 2010

1 > Studies of Australian Hydrobiosella Tillyard: a review of the Australian species of the Hydrobiosella bispina Kimmins
group (Trichoptera: Philopotamidae)

David I. Cartwright

1 5 > A new species of Paraulopus (Aulopiformes: Paraulopidae) from seamounts of the Tasman Sea
Martin F. Gomon

19 > Taxonomic revision of the genus Ratabulus {Teleostei: Platycephalidae), with descriptions of two new species
from Australia

Martin F. Gomon and Flisashi Imamura

35 > Upper Devonian osteichthyan remains from the Genoa River, Victoria, Australia
Timothy Holland

45 > Three new species of the crangonid genus Metacrangon Zarenkov {Crustacea: Decapoda: Caridea) from
Australia

Tomoyuki Komai and Joanne Taylor

61 > A revision of Antarctic and some Indo-Pacific apod id sea cucumbers {Echinodermata: Holothuroidea: Apodida)
P. Mark O’Loughlin and Didier VandenSpiegel

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