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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)

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

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

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

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

- 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

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

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.

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

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).

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.

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.

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.

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).

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

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.

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Munoz-Quesada, F. J. and Holzenthal, R. W. 2008. Revision of the

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Australia. Australian Journal of Zoology 30: 271-325.

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Region. Dr W. Junk Publishers, Dordrecht. 286 pp.

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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.

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Michaelsen, W. and Hartmeyer. R. (eds). Die Fauna Sud west-

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

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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.

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

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

Anal -fin rays

8-9

Pectoral-fin rays

Pelvic-fin rays

Gill rakers

8+19-27

8-9 + 18-20 -

27-28

Pored lateral-line scales

48-49

Scales above lateral line

5.5

Scales below lateral line

3.5

Predorsal scales

19-21

Vertebrae

19 + 29

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

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)

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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

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

f ']

140 1

K-

t Vn

l */

■ n

, s

$> ,

, \ \ J

fLt

,rQ

• Ratabulus diversidens

A Ratabulus futviguttatus sp nov.

O Ratabulus megacephalus

□ Ratabulus ventralis sp.nm.

ft ^ .

‘Xx

* -JT A

A ^ -

/: A,

L-

--V

v-’

per

100 1» 140 160

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

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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

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.

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

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

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

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).

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

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).

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

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).

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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Magazine 29: 7-12 (in Japanese).

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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

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

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.

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

Bone of attachment

Orthodentine

Plication

Orthodentine

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

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

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

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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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.

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

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.

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

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;

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

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.

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

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.

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.

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

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

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. 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

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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.

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

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)

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,

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)

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.

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)

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)

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.

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.

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

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.

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)

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

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)

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

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

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

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

Hook

length

Tentacle rod

length

Tentacle

number

Scoliorhapis species

Soliorhapis theeli (type species)

Heding 1928

no data

absent

100-140

106

Scoliorhapis biopearli sp. nov.

S Shetland Is (this work)

6 mm

absent

168-184

112-120

Scoliorhapis massini sp. nov.

Shag Rocks (this work)

20 mm

absent

88-104

up to 80

Scoliorhapis ? massini sp. nov.

Falkland Is (this work)

50 mm

absent

120-176

120-160

Sigmodota species

Sigmodota contorta

(type species for Sigmodota )

Ludwig 1875

45 mm

140-160

no data

Ludwig 1898

45 mm

44—130

170-200

156-182

H.L. Clark 1921

no data

42-130

170-210

170

Ekman 1925

44 mm

45-120

140-170

no data

Pawson 1964

50 mm

42-130

140-200

170

Massin & Heterier 2004

45 mm

35-100

160-250

140-170

Bouvet I. (this work)

10 mm

40-88

224-240

120-128

Falkland Is (this work)

40 mm

40-128

136-200

80-184

S Georgia (this work)

45 mm

48-96

192-248

80-144

S Georgia (this work)

8 mm

56-72

no data

up to 128

S Orkney Is (this work)

9 mm

40-88

152-184

96-112

S Shetland Is (this work)

15 mm

56-104

168-176

96-112

Magellanic (this work)

40 mm

up to 136

up to 208

up to 184

Heard I. (this work)

no data

88-104

192-208

104-168

Summary

50 mm

35-136

136-250

80-184

Sigmodota dubia

Fisher 1907

60 mm

90-175

185-230

no data

Sigmodota magnibacula

Massin & Heterier 2004

72 mm

100-200

160-180

170-280

S Orkney Is (this work)

70 mm

112-160

192-216

256-296

S Orkney Is (this work)

15 mm

56-120

128-176

176-216

Ross Sea (this work)

25 mm

96-136

152-200

192-240

Ross Sea (this work)

73 mm

110-140

170-190

200-270

Wilkes Land (this work)

60 mm

up to 104

up to 192

up to 272

Prydz Bay (this work)

105 mm

80-186

176-192

184-320

Summary

105 mm

56-200

128-216

170-320

Taeniogyrus species

Taeniogyrus antarcticus

Heding 1931

no data

172-200

83-103

S Orkney Is (this work)

15 mm

64-80

200-208

128

S Orkney Is (this work)

7 mm

40-64

120-152

80-104

S Orkney Is (this work)

3 mm

48-72

112-120

128

S Orkney Is (this work)

2 mm

40-56

80-88

Shag Rocks (this work)

8 mm

48-64

144-168

no data

Shag Rocks (this work)

1 mm

48-64

64-80

64-72

Summary

15 mm

40-80

64-208

64-128

Taeniogyrus australianus

(type species for Taeniogyrus )

Australia (this work)

95 mm

48-88

112-136

72-104

Heding 1928

60 mm

70-100

up to 110

110

Summary

95 mm

48-100

110-136

72-110

Taeniogyrus maculatus

H.L. Clark 1921

26 mm

50-100

66-77

about 45

Newcastle (this work)

21 mm

48-104

80-96

56-72

Summary

26 mm

48-104

66-96

45-72

Taeniogyrus prydzi sp. nov.

Prydz Bay (this work)

7 mm

up to 90

232-240

136-144

Prydz Bay (this work)

50 mm

up to 90

256-272

136-152

Summary

50 mm

up to 90

232-272

136-152

Taeniogyrus purpureus

Ludwig 1898

33 mm

165-182

135-156

78-87

Ekman 1925

33 mm

70-180

80-150

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

Magellanic (this work)

15 mm

80-144

120-160

56-120

Summary

100 mm

70-182

80-160

56-120

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).

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

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).

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)

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).

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;

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

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).

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

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

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.

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).

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

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)

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,

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.

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

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

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)

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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