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CONTENTS

The New Zealand Cucumariidae (Echinodermata, Holothuroidea)

P. M. O'Loughlin and N. Alcock .1

A review of the cucumariid genus Psolidiella Mortensen (Echinodermata, Holothuroidea)

P. M. O'Loughlin .25

A review of the Tubificidae ( Annelida: Oligochaeta) from Australian inland waters

A. M. Finder and R. O. Brinkhurst .39

New Australian species of Oecetis allied to O. complexa Kimmins (Trichoptera: Leptoceridae)

A. Wells .77

Thaumatelsonine stenothoids (Crustacea: Amphipoda): Part 1

T. Krapp-Schickel .89

Rediagnosis of the endemic southern Australian genus Parastacilla Hale, 1924 (Crustacea:

Isopoda: Arcturidae) with descriptions of two new species

R. A. King .125

Levinebalia maria , a new genus and new species of Leptostraca (Crustacea) from Australia

G. K. Walker-Smith .137

Revision of the stegocephalid genera Phippsia and Tetradeion (Crustacea: Amphipoda)

with description of four new species

J. Berge and W. Vader .149

CONTENTS

The New Zealand Cucumariidae (Echinodermata, Holothuroidea)

P. M. O'Loughlin and N. Alcock .1

A review of the cucumariid genus Psolidiella Mortensen (Echinodermata, Holothuroidea)

P. M. O'Loughlin .25

A review of the Tubificidae (Annelida: Oligochaeta) from Australian inland waters

A. M. Pinder and R. O. Brinkhurst .39

New Australian species of Oecetis allied to O. complexa Kimmins (Trichoptera: Leptoceridae)

A. Wells .77

Thaumatelsonine stenothoids (Crustacea: Amphipoda): Part 1

T. Krapp-Schickel .89

Rediagnosis of the endemic southern Australian genus Parastacilla Hale, 1924 (Crustacea:

Isopoda: Arcturidae) with descriptions of two new species

R. A. King .125

Levinebalia maria, a new genus and new species of Leptostraca (Crustacea) from Australia

G. K. Walker-Smith .137

Revision of the stegocephalid genera Phippsia and Tetradeion (Crustacea: Amphipoda)

with description of four new species

J. Berge and W. Vader .149

Memoirs of Museum Victoria 58(1): 1-24 (2000)

THE NEW ZEALAND CUCUMARIIDAE (ECH1NODERMATA, HOLOTHUROIDEA)

P. Mark O’Loughlin' and Nicola Alcock 2

'Invertebrate Zoology, Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia

(moloughlin@parade.vic.edu.au)

"Department of Marine Science, University of Otago, Dunedin, New Zealand (n.alcock@niwa.cri.nz)

Abstract

O’Loughlin, P.M. and Alcock, N., 2000. The New Zealand Cucumariidae (Echinodermata,

Holothuroidea). Memoirs of Museum Victoria 58(1): 1-24.

Three new cucumariid holothurians, Pseudocnus sentus, SquamoCnus niveus and Squamoc-

nus lutens, are described from Stewart Island and Fiordland. Five New Zealand and Australian

cucumariid species, previously referred to the European genus Ocnus Forbes and Goodsir, are

assigned to other genera. Squamocnus brevidentis (Hutton) is a new combination. Australoc-

nus and Psolidocnus are new genera, and Australocnus calcareus (Dendy), Australocnus

occiduus (O’Loughlin and O’Hara), Psolidocnus amokurae (Mortensen), Psolidocnus

farquhari (Mortensen) and Psolidocnus sacculus (Pawson) are new combinations. Ocnus brev¬

identis var, carnleyensis (Dendy) is not recognised. The status of a syntype of Squamocnus

brevidentis (Hutton) is discussed. A neotype is established for Australocnus calcareus

(Dendy). Trachythyone bollonsi (Mortensen) is synonymised with Psolidiella nigra

Mortensen. The southern Australian Plesiocolochirus ignavus (Ludwig) is reported from New

Zealand. S. brevidentis and S. niveus sp. nov. are brood-protecting. A key is provided for the

New Zealand species of Cucumariidae.

Introduction

Recent field observations and collections by one

of us (Alcock) at Stewart Island and Fiordland in

southern New Zealand have yielded new species

of Pseudocnus Panning, 1949 and Squamocnus

O’Loughlin and O’Hara, 1992, and provided the

basis for this review.

The New Zealand cucumariid holothurians are

known mainly from the works of Hutton (1872,

1878), Dendy (1897, 1909), Ludwig (1898),

Dendy and Hindle (1907), Mortensen (1925),

Panning (1949, 1961, 1971), Dawbin (1950) and

Pawson (1963, 1968, 1970, 1983). A problem

developed historically with the identity of Ocnus

brevidentis (Hutton, 1872). The combined evi¬

dence of the original description, and a sub¬

sequent description by Dendy (1897) of the

probable type specimen in a very poor state of

preservation, points to the same species as Dendy

(1909) was to later describe as a variety ( brevi¬

dentis var. carnleyensis). When the variety was

described Dendy (see Dendy and Hindle, 1907)

wrongly assumed that another common and

related but undescribed species was brevidentis.

Mortensen (1925) followed the same thinking.

This common undescribed species is one of two

new Squamocnus species described here. A syn¬

type, of uncertain status, from Stewart Island

labelled Thyone brevidentis Hutton, held in The

Natural History Museum in London, is the straw-

berry-red cucumariid described by Dendy (1909)

as brevidentis var. carnleyensis but confirmed

here to be Hutton’s brevidentis.

Five cucumariid species and one variety from

New Zealand and Australia have at times been

referred to the initially European genus Ocnus

Forbes and Goodsir in Forbes, 1841 - O. brevi¬

dentis (Hutton, 1872), O. calcareus (Dendy,

1897), O. brevidentis var. carnleyensis (Dendy,

1909), O. farquhari (Mortensen, 1925), O. saccu¬

lus Pawson, 1983 and O. occiduus O’Loughlin

and O’Hara, 1992. A review of the assignment of

these species to Ocnus faces a further historical

problem as to which is the type species of Ocnus.

Panning (1949) revived the genus Ocnus, desig¬

nating O. brunneus (Forbes ms. in Thompson,

1840) as the type species. Mortensen (1927) had

synonymised O. brunneus with O. lacteus

(Forbes and Goodsir, 1839). Panning (1971)

accepted the synonymy by Cherbonnier (1951) of

O. brunneus with O. planci (Brandt, 1835), and

considered O. planci to be the type species. Rowe

(1970, 1995 in Rowe and Gates) followed

Mortensen’s synonymy, nominating O. lacteus as

the type species. McKenzie (1984) detailed the

history of the genus Ocnus ; recognised O. brun¬

neus as distinct from O. lacteus ; described a neo¬

type for O. brunneus ; and discussed the possible

synonymy of O. brunneus with O. planci.

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

McKenzie (1991) subsequently considered grow¬

ing evidence that O. planci and O. brunneus were

conspecific, and that characters used to separate

(). brunneus from O. lacieus were not reliable. O.

planci , O. lacteus and O. brunneus have all been

designated type species for Ocnus which is con¬

sidered here to be a European genus with diag¬

nostic characters inapplicable to the New Zealand

and Australian species.

Panning (1971) restricted Ocnus to five

species, included (). calcareus, excluded O. brev¬

identis and O.farqtthari, reassigned brevidentis to

Pentacta Goldfuss, 1820, but did not reassign O.

farquhari. O’Loughlin and O’Hara (1992) and

Rowe (in Rowe and Gates, 1995) continued to

assign brevidentis to Ocnus. Rowe (in Rowe and

Gates, 1995) reassigned occiduus lo Plesio-

colochirus Cherbonnier, 1946. Panning (1971:

36), Pawson (1983: 227), O’Loughlin and O’Hara

(1992: 237, 247) and Rowe (in Rowe and Gates,

1995: 279) questioned whether Ocnus was the

appropriate genus for New Zealand and Aus¬

tralian species. The five species listed above fall

into three groups. Ocnus brevidentis (Hutton) is

reassigned to Squamocnus O’Loughlin and

O’Hara, where two of the new species are

assigned. O. calcareus (Dendy) and O. occiduus

O’Loughlin and O’Hara are assigned to a new

genus. O. farquhari (Mortensen) and O. sacculus

Pawson are assigned to a second new genus, as is

Trachythyone amokurae (Mortensen, 1925).

Trachythyone bollonsi (Mortensen, 1925) is

synonymised here with Psolidiella nigra

Mortensen, 1925.

Plesiocolochirus ignavus (Ludwig, 1874),

known from across southern Australia, is reported

for New Zealand from NIWA specimens which

had been identified as Ocnus brevidentis and from

an unidentified MNZ specimen from Fiordland.

O’Loughlin (1994) listed 30 brood-protecting

cucumariid species, including the three New

Zealand species Trachythyone amokurae

(Mortensen), Psolidiella nigra Mortensen and

Ocnus sacculus Pawson. Two additional cases of

brood-protection by the New Zealand cucumariid

species Squamocnus brevidentis (Hutton) and

Squamocnus niveus sp. nov. are reported here.

Abbreviations for institutions are as follows:

AM, Australian Museum, Sydney, Australia;

MNZ, Museum of New Zealand, Te Papa Ton-

garewa, Wellington, New Zealand: NMV,

Museum Victoria, Melbourne, Australia; MPE,

Dr Th. Mortensen’s Pacific Expedition

1914-1916; BMNH, The Natural History

Museum, London; NIWA, New Zealand Institute

of Water and Atmospheric Research, Wellington,

New Zealand; USNM, US Museum of Natural

History, Smithsonian Institution, Washington,

USA; ZMUC, Zoological Museum, University of

Copenhagen, Denmark.

The following terms are defined:

Bent and curved. Elongate tentacle and tube

foot plates may be straight or bent (angular) in

one plane (surface) which itself may be fiat or

curved or concave (Fig. 31 illustrating straight and

curved, and Fig. 4i illustrating bent and curved).

Buttons. Thickened plates which are per¬

forated, regular to irregular in outline, small

(typically 0.08 mm long), discretely knobbed

(Figs 3i, 5c) or lumpy (Figs 6f, 6i) or smooth,

typically four perforations.

Cups. Thin plates which are perforated, con¬

cave, shallow (species in this paper) or deep, oval

to rectangular, small (typically 0.04 mm long),

cruciform (Figs 2g, 3f, 4d) or tripartite (Figs 5i,

5j) centrepiece, typically four large with fre¬

quently four small comer perforations, spinelcts

on rim and sometimes on centrepiece, sometimes

with bridging connections across rim.

Denticulate. With pointed to bluntly spinous

teeth-like spinelets on ossicles (Figs lh, lk).

Digitate. With finger-like spinelets on ossicles

(Figs If, 4d).

Multilayered ossicles. Ossicles which are nodu¬

lar, irregularly oval in outline, large (more than

0.2 mm long), comprising a perforated knobbed

base plate with additional layers built on both

sides creating low domes bilaterally (Figs 2k, 3j).

Sole. Flattened, delimited, modified ventral

body wall, bounded peripherally by tube feet

which do not extend in series to the introvert and

anus, and the place of attachment to the substrate.

Ventrum. Modified ventral body wall, bounded

laterally by the lateroventral series of tube feet,

not bounded peripherally by tube feet, and the

place of attachment to the substrate (Figs 3b, 6b).

Order Dendrochirotida Grube, 1840

Cueumariidae Ludwig, 1894

Remarks. Thirteen species of Cueumariidae are

now known for New Zealand. The two species

not discussed, Amphicyclus thomsoni (Hutton,

1878) and Neocucumella bicolumnata (Dendy

and Hindie, 1907), were discussed and illustrated

by Pawson (1963, 1970). Ten are endemic

species, while P ignavus is a common southern

Australian species, A. calcareus is reported for

Macquarie and Juan Fernandez Islands, and P.

nigra occurs in south-eastern Australia

(O’Loughlin, 2000). A South Australian speci¬

men, initially referred to N. bicolumnata, was

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

subsequently assigned to Neocucumella fracta

O’Loughlin and O’Hara, 1992. The continuing

absence of Pseudopsolus macquariensis (Bendy)

during recent intensive collecting at Stewart

Island reinforces the probability that it is a Mac¬

quarie Island species, and it is not included in the

key (Mortensen, 1925).

The following cucumariids are distinguished

from other New Zealand holothurian species by

having multiple-branching (dendritic) tentacles;

by having a cylindrical and not U-shaped body

form; by having a calcareous ring of ten pieces

which lack posterior prolongations; and by lack¬

ing a total or dorsal cover of imbricating scales.

Key to New Zealand Cucumariidae

1. Cups and/or rudimentary cup ossicles and/or concave crosses present in

body wall; tables and terminally spinous buttons absent.2

Cups and/or rudimentary cup ossicles and/or concave crosses absent from

body wall; tables or terminally spinous buttons present.10

2 . Multilayered (nodular) ossicles present in body wall.3

Multilayered (nodular) ossicles absent from body wall.9

3. Body with angular dorsolateral edges; tube feet on radii only; dorsal tube

feet surmount raised calcareous wreaths; some cups with bridges across rim;

live colour mottled red, mauve, brown, orange, cream and white

................. PlesiocoJochirus ignavus (Ludwig)

Body rounded dorsolaterally; dorsal tube feet densely or sparsely scattered

or absent; dorsal tube feet not surmounting raised calcareous wreaths; cups

all lack bridges across rim; live colour not mottled.4

4. Ventral tube feet radial series continuous to introvert; dorsal tube feet abun¬

dant and predominantly scattered; multilayered ossicles not macroscopically

evident as scales; body wall cups with predominantly more than 4 holes;

buttons with discrete rounded knobs.5

Ventral tube feet radial series not continuous to introvert; dorsal tube feet

sparse and scattered or absent; multilayered ossicles macroscopically evident

as scales; body wall cups with predominantly 4 holes; buttons with swellings

more than discrete knobs.7

5. Tube feet present on the ventral interradii; shallow concave perforated

knobbed plates about 0.1 mm long present in body wall; live colour

pale-spotted strawberry-red dorsally; lacking dark brown to black coloration

anteriorly and posteriorly. Squamocnus brevidentis ( Hutton)

Tube feet absent on ventral interradii; lacking concave perforated knobbed

plates; live colour white or brown or grey or yellow; dark brown to black

coloration present anteriorly and posteriorly.6

6. Cups in body wall with predominantly finely knobbed rarely semidigitate

spinelets; largest elongate tentacle plates often straight and curved; live

colour predominantly white. Squamocnus niveus sp. nov.

Cups in body wall frequently with semidigitate to digitate spinelets; largest

elongate tentacle plates bent and curved, not straight; live colour yellow

.. Squamocnus luteus sp. nov.

7. Tube feet scattered and sparse dorsally; abundant concave multiradiate

ossicles epidermally. Psolidocnus amokurae (Mortensen)

Tube feet absent dorsally; multiradiate ossicles absent.8

8. Ventral tube feet in double radial series; dorsal body wall scales frequently

towered, creating a knobbed surface. Psolidocnusfarquhari (Mortensen)

Ventral tube feet in single radial series; dorsal body wall scales not towered,

creating a smooth plated surface. Psolidocnus sacculus (Pawson)

9. Distinct sole; tube feet numerous and scattered dorsally, laterally, around

oral and anal cones; ventral body wall ossicles concave crosses and

rudimentary cups, perforated plates small, buttons not discretely knobbed;

dorsal colour blackish. Psolidiella nigra Mortensen

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Lacking a distinct sole; ventral tube feet in radial series; dorsal tube feet pre¬

dominantly in radial series; body wall ossicles predominantly knobbed but¬

tons with commonly 4 holes and 12 peripheral discrete knobs, cups, rare

large single-layered perforated plates; live colour white with dorsal tube tect

ends red... Australocnus calcareus (Dendy)

10. Table ossicles present in body wall, at least anteriorly and posteriorly; 20 or

25 tentacles...11

Table ossicles not present in body wall; 10 tentacles.12

11. Table ossicles abundant, regular, 8 perforations; 20 tentacles; tube feet in

double radial rows; tube feet discs and tentacles not white.

. Neocucumella bicolumnuta (Dendy and Hindle)

Table ossicles present only anteriorly and posteriorly, irregular, predomi¬

nantly more than 8 perforations; 25 tentacles; Uibe feet in more than 2 rows

radially; tube feet discs and tentacles white ...Amphicyclus thomsoni (Hutton)

12. 10 equal tentacles; body wall ossicles knobbed buttons spinous at one end;

multilayered ossicles absent. Pseudoants leonirtoides (Mortenscn)

8 large and 2 small ventral tentacles; knobbed buttons and multilayered

ossicles present, some with spinous ends. Pseudocnus sentus sp. nov.

Pseudocnus Panning

Pseudocnus Panning, 1949: 422.—Panning,

1962: 58.—Lambert, 1998: 474.

Diagnosis (emended). Cucumariidae with body

wall ossicles pear-shaped to irregular, single¬

layered, knobbed, perforated plates with one end

denticulate (laevigatus-gtoup)', sometimes with

knobbed buttons also present, and sometimes the

denticulate plates intergrading with or replaced by

multilayered ossicles some of which are den¬

ticulate (dubiosus-grovtp); sometimes buttons

only (cwni/Ms-group); cups and tables absent.

Remarks. The new species described here has

knobbed buttons, and multilayered ossicles some

of which have a denticulate end. The latter inter¬

grade with single-layered terminally digitate ossi¬

cles which are typical of Pseudocnus. The combi¬

nation of ossicle types in the new species places it

in the dubiosus -group of Pseudocnus categorised

by Panning (1962: 58-59). Descriptions and illus¬

trations of some members of the dubiosits-gwup

by Panning (1962: 66-69) indicate the presence

of multilayered ossicles. Rather than establish a

new monotypic genus, the relationship with Pseu¬

docnus is recognised and the diagnosis emended

to explicitly include multilayered terminally den¬

ticulate ossicles. Based on morphology and mito¬

chondrial DNA studies Lambert (1998) emended

the diagnosis of Pseudocnus to include species

with thick lobed non-denticulate buttons only,

and proposed a cwra/i/jf-group in addition to the

dubiosus- and laevigatus-groups of Panning

(1962: 58, 70). Pseudocnus now embraces species

with widely different ossicle types.

Pseudocnus sentus sp. nov.

Figure la-j

Material examined. Holotype (MNZ EC7483). New

Zealand, Stewart Island, Paterson Inlet, S side of Faith

Hope and Charity Group, granite or diorite rock sub¬

strate covered in pink coralline alga, 0-4 m, N. Alcock,

30 Mar 1998.

Paratypes. Type locality and date, NMV F82784 (1);

26 Sep 1998, MNZ EC7484 (5). NMV F82781 (6),

NIWA (5).

Comparative material. Syntypes of Pseudocnus

leoninoides (Mortensen, 1925) [ZMUC (207), MNZ

EC531 (16)], Auckland Islands, Masked Island, Cam-

ley Harbour, on rock wall with Me lobes ia. MPE, 30

Nov 1914 (Figure 1 k-l).

Diagnosis. Cucumariidae with 10 dendritic tenta¬

cles, ventral 2 smaller; tube feet in zig-zag to dou¬

ble radial rows ventrally, less regular double to

zig-zag rows dorsolaterally, rare interradially;

body wall ossicles irregular knobbed buttons,

single to multilayered (up to 0.6 mm long)

knobbed ossicles sometimes terminally denticul¬

ate or digitate and sometimes narrowed at the

denticulate end, rosettes in tentacles and introvert.

Description. Body up to 27 mm long, 8 mm dia¬

meter (preserved, tentacles fully withdrawn),

rounded to quadrangular to pentagonal in trans¬

verse section, elongate, anal cone upturned,

mouth slightly upturned, lacking distinct ven-

trum, oral valves well developed: body wall thin,

firm, flexible, calcareous, surface rough; 10 den¬

dritic tentacles, ventral 2 smaller; calcareous ring

with narrow parallel anterior projections, deep

posterior indentations (interradial) and notches

(radial), lacking posterior prolongations; 1 2

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

polian vesicles left lateral; small free madreporite

close to ring dorsally; microscopic anal scales.

Tube feet extending from introvert to anus,

close zig-zag to double radial rows ventrally, less

regular double to zig-zag rows dorsolaterally, rare

interradially, single very extensible tube feet at

introvert, tube feet not on introvert, 5 present

anally.

Body wall with multilayered ossicles, some¬

times imbricating anteriorly and posteriorly, some

denticulate at one end, up to 0.6 mm long; inter¬

grading with irregular finely to thickly knobbed

buttons, some digitate at one end or side, typically

0.1 mm long; tube feet with endplates, thin per¬

forated denticulate irregularly oval curved sup¬

port plates 0.1 mm long, elongate bent centrally

widened perforated knobbed support plates 0.24

mm long; introvert with numerous thin finely-

knobbed (sometimes with connecting bridges)

perforated non-denticulate plates typically 0.08

mm long with 4 central holes, intergrading with

open to densely branched and knobbed rosettes

typically 0,05 mm long, fewer smooth or knobbed

thick perforated non-denticulate oval to triangular

plates typically 0.17 mm long; tentacles with thin

perforated denticulate irregularly oval convex

plates 0.08 mm long, irregular elongate ossicles

0.16 mm long with large central perforations,

rosettes 0.04 mm long. Ossicle form consistent

through small to large specimens.

Colour (live). Body white, to white with brown

spots, to pale brown (grey preserved) with brown

spotting, to dark grey with black spotting; tenta¬

cles yellow, with or without dark brown mark¬

ings; intestine (preserved) cream, to cream with

brown spotting.

Reproduction. Gonad tubules long, unbranched;

prominent without discernible eggs in September.

Etymology. From the Latin sentus (rough),

referring to the surface of the body.

Distribution. New Zealand, Stewart Island,

Paterson Inlet, rocky shallows, 0-4 m.

Remarks. This new species is distinguished from

other species of Pseudocnus by having single and

multilayered, knobbed, sometimes terminally

denticulate or digitate body wall ossicles. Pseu¬

docnus leoninoides (Mortensen, 1925) was

reported by Dendy (1909) for the Auckland

Islands (as Cucumaria leonina Semper var.), by

Mortensen (1925) from Auckland and Campbell

Islands, and by Pawson (1965) from The Snares

islands (just south of Stewart Island). A specimen

from the Macquarie Seamount (NIWA stn D18),

identified as P. leoninoides , was redetermined by

the authors as Pseudocnus laevigatus (Verrill,

1876). Pawson noted (1965: 258) that leoninoides

could be expected for New Zealand, and while

Pseudocnus is reported here for Stewart Island it

is not P. leoninoides.

Numerous syntypes of Pseudocnus leoninoides

were examined (Figures 1 k-l). With single-lay¬

ered, typically 0.14 mm long, knobbed, terminally

denticulate body wall ossicles only, it belongs to

the laevigatus- group of Pseudocnus species. In

addition to lacking knobbed non-denticulate but¬

tons and multilayered sometimes denticulate

ossicles and tentacle rosettes, P. leoninoides has

10 equal tentacles, has radial tube feet on the

introvert, has a very thin indistinctly present cal¬

careous ring, and has a thin relatively soft body

wall. In all of these features P. leoninoides differs

from P. sentus.

In body form, tentacle number and form, tube

feet distribution, and ossicle combination of

multilayered ossicles and knobbed buttons with

an absence of cups, Pseudocnus sentus is similar

to Cucuvitrum rowei O’Loughlin and O’Hara,

1992. But the monotypic genus Cucuvitrum lacks

the distinctive pointed and digitate ossicle

spinelets of P. sentus.

Squamocnus O’Loughlin and O’Hara

Squamocnus O’Loughlin and O’Hara, 1992:

236-237.

Diagnosis (emended). Cucumariidae with cal¬

careous body wall; body rounded to slightly angu¬

lar dorsolaterally; lacking body wall growths on

radii; distinct ventrum with tube feet series

extending to introvert and frequently to anus, not

sole; 10 dendritic tentacles, ventral 2 smaller;

tube feet scattered dorsally and laterally some¬

times in irregular dorsolateral series, on ventral

radii sometimes extending onto ventral interradii,

absent on introvert. Body wall ossicles shallow

cruciform spinous cups (up to 0.05 mm long),

larger cups sometimes intergrading with shallow

concave finely knobbed plates (up to 0.13 mm

long), regular to irregular buttons with discrete

knobs, numerous large multilayered ossicles;

lacking connecting bridges across cups and

buttons; sometimes with tentacle rosettes.

Type species. Squamocnus aureoruber

O’Loughlin and O’Hara, 1992.

Species. S. aureoruber O’Loughlin and O’Hara,

1992, S. brevidentis (Hutton, 1872), S. niveus sp.

nov., S. luteus sp. nov..

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Distribution. Coastal New Zealand, New Zealand

subantarctic islands, and south-eastern Australia;

0-130 m.

Remarks. The genus Squamocnus comprises four

closely related species characterised by a rounded

dorsal body, scattered dorsal and lateral tube feet,

ventral tube feet series extending to introvert, anil

a body wall ossicle combination of large multi¬

layered ossicles, buttons with discrete knobs, and

cruciform spinous cups without bridges. The

presence of multilayered ossicles, cups that are

predominantly cruciform as distinct from triradi-

ate, and scattered dorsal and lateral tube feet dis¬

tinguish Squamocnus from Ocnus Forbes and

Goodsir.

Squamocnus brevidentis (Hutton) comb. nov.

Figure 2a-l

Colochirus brevidentis. Dendy, 1897: 40 41, pi. 5

figs 54—61.—Farquhar, 1898: 325.

Cucumaria brevidentis. —Perrier. 1905: 110 [uncer¬

tain S. brevidentis], —Dendy and llindle, 1907: 99-100

[some, if not all, non .S', brevidentis]. —Dawbin, 1950:

38, fig. 10 [part],

Cucumaria brevidentis (Hutton) var. carnleyensis

Dendy, 1909: 149-151. pi. 6: 2a—1. -Mortensen, 1925:

332-335, figs 24-25.

Ocnus brevidentis var. carnleyensis. —-Panning,

1949:437.

Ocnus brevidentis. —Pawson, 1968: 9, 11, 21-22,

25-27 [part].— 1970: 39-40, fig. 8, pi. 2(1) [part].—

Fenwick and Homing, 1980: 440, 443 [part].

O’Louahlin and O'Hara, 1992: 237. - Rowe (in Rowe

and Gates, 1995): 279.—Alcock, L999: 9-29.

Material examined. Syntype (labelled “? Co-type")

[BMNH 86.11.18.6.1 (1)]. New Zealand, Stewart

Island.

Other material. New Zealand, South Island, Fiord¬

land. Preservation Inlet, exposed rock surfaces, large

cluster colonies, 2-15 m, 30 Nov 1996, NMV F81958

(5) ; 30 Jul 1997, NMV F81957 (17); Kaikoura, 42°42'

S, 173°38' E, 100-112 m, 14 Dec 1982. MNZ EC7486

(1); 42°28' S. 173°40' E, 95-98 m, 21 Dec 1982, MNZ

EC6908 (1): 42°38’ S, 173°40' E. 120-130 m, 11 Dec

1982, MNZ EC691S (15); Foveaux Strait. 18 m, Feb

1952, MNZ EC6866 (I); Stewart Island. Paterson Inlet,

S side of Faith Hope and Charity Group, granite or dior-

ite rocky substrate covered in pink coralline alga, under

rocks, small groups, 0-5 m, 20 Aug 1997, NMV

F8I956 (6); 26 Sep 1998, MNZ EC7485 (3); NIWA

(3); Paterson Inlet shore and Ulva Island, MNZ EC533

(6) ; Bounty Platform, 49°40' S, 178°47' E, 63 m, 9 Nov

1962, NIWA stn A738 (4); Antipodes Islands, 73 m, 7

Nov 1962. NIWA stn A729 (I); SCUBA, 27 Nov.

1978, NMV F82779 (2).

Description of material. Body up to 55 mm long

(live, tentacles partly extended); body wall thick.

firm to hard; body domed dorsally, flat vcntrally,

distinct ventrum thin-walled in relaxed speck

mens, not sole; mouth anterior, slightly upturned,

5 weakly developed radial oral valves; upturned

anal cone, 5 microscopic anal scales; 10 dendritic

tentacles, ventral 2 smaller; calcareous ring solid,

slender tapering anterior projections and posterior

indentations radially and intcrradially, lacking

posterior prolongations; single left lateral poliafi

vesicle.

Tube feet up to 4 rows wide on radii of

ventrum, extending onto ventral interradii and

ventrolaterally, radial series extending to intro-

vert; double radial rows near introvert on oral

valves, absent on introvert, very extensible promk

nent single radial tube feet at introvert; tube feet

dorsally and laterally and around anal cone fewer,

evenly scattered, with low tubercle bases.

Body wall ossicles cups, small concave finely

knobbed perforated plates, buttons and multi¬

layered ossicles; small cups regular cruciform,

rarely with triradiate centrepiece, 0.025-0.04K

mm long, 4-8 holes, smallest cups with bare cru¬

ciform centrepiece and digitate spinelets on rim

all pointing up from cup; larger cups intergrade

with regular cruciform to irregular very shallow

concave plates; concave plates frequently with 8,

up to 26, holes large centrally decreasing in size

towards periphery, 0.09 0.13 mm long, finely

knobbed over whole plate, knobs pointing in all

directions; concave plates intergrade with but¬

tons; buttons with discrete knobs predominantly

irregular, about 1 in 4 may be regular with 4

holes, typically 4-7 very rarely up to 14 holes,

knobs bulbous and sometimes fused and irregular,

small buttons frequently with 1 oblong knob cen¬

trally less commonly 2 discrete knobs centrally,

smallest buttons typically 0.1 mm long; numerous

multilayered ossicles up to 0.9 mm long. Introvert

with typical cups and knobbed concave plates

only; concave plates frequently elongate, up to

0.13 mm long. Tentacle ossicles large elongate

narrow curved thickened perforated plates (holes

decreasing in size distally) up to 0.6 mm long,

irregular perforated denticulate convex plates typ¬

ically 0.06 mm long, narrow centrally-enlarged

perforated bent plates typically 0.15 mm long,

lacking rosettes. Tube feet ossicles endplatcs up

to 0.24 mm diameter, irregular denticulate per¬

forated curved plates typically 0.15 mm long,

elongate narrow centrally-widened curved bent

perforated plates typically 0.15 mm long, cups,

buttons. Anal ossicles typical body wall ossicles

and large perforated plates; perforated plates

thick, irregularly elongate or pear-shaped,

single-layered not knobbed at end(s), knobs and

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

secondary layers extensive, frequently bluntly

denticulate at narrow end, up to 0.8 mm long.

Colour (live). Body wall, dorsally and laterally

and around oral and anal cones, and tentacles

strawberry-red; ventrum cream to pale pink;

dorsal and lateral white spotting associated with

tubercles and tube feet; small dark brown irre¬

gular spots dorsally, laterally, ventrally; lacking

dark coloration or markings on tentacles or intro¬

vert or anteriorly or anally. Preserved colour

(alcohol) fades slowly to pale brown and finally

white.

Reproduction. Live male gonad tubules pink,

female tubules brownish red; elongate,

unbranched; eggs up to 0.75 mm long (March);

brood-protects during autumn/winter under body

(Alcock, 1999); not fissiparous.

Distribution (confirmed here). New Zealand.

South Island, Kaikoura, Fiordland, Foveaux

Strait, Stewart, Antipodes, Auckland and

Campbell Islands, Bounty Platform; 0-130 nr.

Remarks. There is no literature reference to a syn-

type, and the one held by BMNH is recorded with

a question mark. Locality is given, but no infor¬

mation about collector, date, identifier or donor.

The specimen is in good condition and is clearly

identifiable as the strawberry-red species and is

confirmed here as S. brevidentis. Hutton (1872)

described a brown body, pink tentacles with

yellowish tips, and tube feet all over the body and

more spread dorsally than ventrally for the type

specimen of 5. brevidentis. In describing what he

judged to be the same specimen Dendy (1897)

added that the tube feet were thickly and irregu¬

larly scattered over the ventral surface, the dorsal

tube feet were abundant and fairly uniformly scat¬

tered over the remainder of the body and twice the

size of the ventral ones, and the ossicles were

large reticulate nodules, irregular buttons, and

cups similar to S. calcaneus but with some more

fiat and irregular. Dendy (1897: fig. 58) illus¬

trated these last-mentioned distinctive ossicles for

the type specimen of S. brevidentis , and illus¬

trated (1909: pi. 6, 2h and 2k) similar ossicles for

S. brevidentis var. carnleyensis. Hutton’s and

Dendy’s characters refer to the strawberry species

described subsequently by Dendy (1909) as S.

brevidentis var. carnleyensis. Mortcnsen (1925)

followed Dendy (1909). The descriptions of var.

carnleyensis by Dendy (1909) and Mortensen

(1925) are descriptions of Hutton’s S. brevidentis.

There are white and yellow (white preserved)

undescribed species in New Zealand which are

similar to the strawberry 5. brevidentis, and one

or both have been assumed to be S. brevidentis by

Dendy and Hindle (1907), Dendy (1909),

Mortcnsen (1925) and Panning (1949, 1971). In

addition to differing from S. brevidentis in live

colour, both species (described below) have

double ventral radial series of tube feet with bare

interradii, and lack the unique shallow concave

knobbed plate ossicles illustrated by Dendy

(1909: pi. 6, 2h and 2k). References by Panning

(1949, 1971) to S. brevidentis are generally appli¬

cable to the strawberry, white and yellow species,

but the two specimens from Cook Strait described

and illustrated by Panning (1949: 437^138, fig.

32) are the white species described below.

Ludwig (1898) considered Colochirus cal-

careus Dendy, 1897 to be a junior synonym of C.

brevidentis, and was followed by Perrier (1905),

Dendy and Hindle (1907) and Dendy (1909).

Mortensen (1925) and subsequent authors recog¬

nised C. calcareus.

In assigning the species brevidentis to Pentacta

Goldfuss, 1820, and diagnosing the genus. Pan¬

ning (1971: 38-39) recognised that the genus was

not homogeneous and that brevidentis occupied

an uncertain place. And in fact, he was dealing

with a different species (see next species).

Recently Rowe (in Rowe and Gates, 1995: 271)

has discussed the diagnosis of Pentacta. Panning

considered appendages confined to the radii,

growths on the radii surmounted by tube feet or

papillae, bridges across knobbed ossicles, and

four or five-edged body form, to be characteristic

of Pentacta. None of these features occurs in S.

brevidentis which is assigned here to Squamocnus

as diagnosed above.

Pawson (1968, 1970) did not recognise var.

carnleyensis and in describing 5. brevidentis

referred to the strawberry species (Pawson, 1970:

ossicles fig. 8C' are characteristic of brevidentis)

and white species (Pawson, 1968: dark dorsal

colour occurs in the white but not red species),

and included a distribution around the North

Island where the red species has not been col¬

lected. Pawson (1968: 26-27) recorded S. brevi¬

dentis from Macquarie Island. Mortensen (1925:

335) had cast some doubt on the basis for this

record. A recent study of the echinodenn fauna of

Macquarie Island (O’Hara, 1999) did not confirm

the occurrence of S. brevidentis in the Macquarie

Island fauna. Mortensen (1925) reported S. brevi¬

dentis from the Auckland and Campbell Islands

as var. carnleyensis.

One of the specimens from the Antipodes

Islands collected on 27 November has eggs which

are 0.7 mm long in the gonad tubules, while no

specimen from southern New Zealand collected

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

in August and November has similar rnacro-

scopically discernible eggs. This indicates a prob¬

able variation in the reproductive cycle across the

geographical range of S. brevidentis. It was also

noted that regular buttons with four holes are

more common in specimens of S. brevidentis

from the subantarctic islands than from southern

New Zealand. These observations indicate that

speciation may be occurring across the geo¬

graphical range of this brood-protecting species.

•S’, brevidentis is distinguished from other

species of Squamocnus by its relatively large size

and pale-spotted strawberry-red colour (live), by

the presence of tube feet on the ventral interradii,

by the presence in the body wall of shallow

slightly concave and finely knobbed perforated

plates commonly 0.1 mm long, by the presence of

digitate spinelets on the smallest cups, and by a

habit of seasonal winter brood-protection under

the body (Alcock, 1999). Characteristic ossicle

form can be distinguished in juveniles as small as

5 mm long.

Squamocnus niveus sp. nov.

Figure 3a-l

Cucumaria brevidentis. —Dendy and Hindle, 1907:

99-100 [probably part].— Mortensen, 1925: 331-332.

fig. 26 a-b.—Dawbin, 1950: 38, fig. 10 [part],

Ocnus brevidentis. —Panning, 1949: 437-438, fig.

32.—Pawson, 1968: 9, 11.21-22,25-27 [part].—1970:

39-40, fig. 8, pi. 2(1) [part],

Pentacta brevidentis. —Panning, 1971: 38-39.

Material examined. Holotype (MNZ EC7487). Stewart

Island, Paterson Inlet, S side of Faith Hope and Charity

Group, granite or diorite rock substrate covered in pink

coralline alga, solitary under rocks, 0M m, N. Alcock,

30 Mar 1998.

Paratypes. Type locality and date, NMV F82783 (5),

NMV F83406 (1, brood-protecting); 21 Jan 1998, NMV

F82777 (3); 26 Sep 1998, MNZ EC7488 (6); NIWA

( 6 ).

Other material. North Island. Cape Maria van

Diemen. MPE, 4 Jan 1915, ZMUC (58); Slipper Island,

37° S, 176° E, MPE, 20 Dec 1914. ZMUC (11 ); Auck¬

land Islands. Masked Island. Camley Harbour, MPE, 3

Dec 1914. ZMUC (105); Bounty Platform. 49°40' S,

178°50' E. 37 m, 9 Nov 1962. NIWA stn A743 (1);

49°40 S, 178°47' E, 63 m, 9 Nov 1962, NIWA stn

A738 (I).

Description. Body up to 28 mm long (live, tenta¬

cles fully extended); body wall thick, firm to

hard; body tubular, domed dorsally, flat ventrally,

distinct ventrum, not sole; mouth anterior, slightly

upturned, 5 weakly-developed radial oral valves;

long upturned anal cone on many specimens; 5

microscopic anal scales; 10 dendritic tentacles,

ventral 2 smaller; calcareous ring solid, fine

tapering anterior projections radially and inter-

radially, posterior notches radially, wide

indentations interradially, lacking posterior

prolongations; single left ventral polian vesicle.

Tube feet in zig-zag to double rows on ventral

radii, extending from introvert to anus, ventral

interradii bare; absent from introvert; irregular

double dorsolateral rows or scattered dorsally and

laterally in midbody, double radial rows near

introvert, single very extensible radial tube feet at

introvert, bases of tube feet sometimes swollen

creating small tubercles.

Body wall ossicles cups, buttons and multi¬

layered ossicles; cups regular cruciform, rarely

with tripartite centrepiece, cups typically

0.028-0.056 mm long, typically 4-8 very rarely

up to 14 holes (very small peripherally), cup

spinelets dense and knobbed to bluntly denticu¬

late rarely semidigitate on larger cups, spinelets

always on centrepiece and pointing up or in all

directions from cup rim; cups intergrade with but¬

tons; buttons with discrete knobs regular and

irregular, commonly 3-6 rarely up to 14 holes,

buttons typically 0.08 mm long; about 1 in 4 but¬

tons regular, 4 uniform holes, 8-10 uniform large

peripheral knobs, 2 central knobs sometimes

larger; numerous multilayered ossicles up to 0.8

mm long. Introvert with typical cups and elongate

very shallow concave finely knobbed plates up to

0.14 mm long. Tentacle ossicles elongate, narrow,

straight or bent, curved, thickened, irregularly-

perforated plates up to 0.4 mm long; bent curved

perforated rods typically 0.15 mm long; irregular

curved and convex denticulate perforated plates

typically 0.06 mm long; lacking rosettes. Tube

feet with endplates up to 0.22 mm diameter, irreg¬

ular thin denticulate perforated curved to convex

plates typically 0.07 mm long, irregularly trian¬

gular to elongate centrally-widened bent curved

perforated plates typically 0.15 mm long, cups,

buttons. Anal ossicles typical body wall ossicles

and large perforated plates; perforated plates

thick, irregularly elongate or pear-shaped, part

single-layered not knobbed, extensive parts with

knobs and secondary layers, frequently bluntly

denticulate at end(-sj, rarely sharply denticulate,

plates up to 0.65 mm long.

Colour (live). Body white; large ossicles in

body wall create greyish spotting; tentacles pink,

rarely pale orange; dark brown to black on tenta¬

cle trunks, introvert, anteriorly and sometimes

dorsally and anally; sometimes fine black fleck¬

ing dorsally (dark markings persist on pre¬

served material); rare brown spotting dorsally and

ventrally.

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Reproduction. Gonads (preserved) orange to

white; long, thin, unbranched tubules; separate

sexes; seasonal autumn and winter internal brood-

protection in coelomic sacs; numerous eggs or

embryos 0.7-0.8 mm long in sacs in March;

coelomic embryos uniformly 1.0 mm long show¬

ing differentiation and ossicle development by

July; no brood-protection evident by September.

Etymology. From the Latin niveus (white as

snow), referring to the live body colour.

Distribution (confirmed here). New Zealand.

North, South, Stewart and Auckland Islands,

Bounty Platform. 0-63 m.

Remarks. This white species was assumed to be S'.

brevidentis by Dendy and Hindle (1907),

Mortensen (1925) and Panning (1949, 1971). In

recognising var. carnleyensis (the true brevi¬

dentis), Mortensen (1925) distinguished it from 5.

niveus (assumed to be S. brevidentis). Mortensen

(1925: 333) noted less scattered dorsolateral tube

feet on Masked Island material and expressed

doubt about referring this material to S. brevi¬

dentis. Panning (1949) described 5. niveus from

Cook Strait material (as Pentacta brevidentis).

Many of the small ZMUC specimens (MPE) from

Cape Maria van Diemen exhibit a deep midbody

constriction, suggesting the possibility of fissipar-

ity, but no anal body ends were present. The type

species of the genus, Squamocnus aureoruber , is

fissiparous. S. niveus is distinguished from the

other species in Squamocnus by its white colour

(live), grey to black anterior and posterior and

sometimes dorsal coloration, absence of ventral

interradial tube feet, absence in the body wall of

shallow concave finely knobbed perforated

plates, buttons which fairly commonly are regular

with four even holes and 8 10 peripheral and two

central knobs, and brood-protection in intra-

coelomic sacs. Characteristic ossicle form can

be distinguished in juveniles as small as 5 mm

long.

Squamocnus luteus sp. nov.

Figure 4a-i

Material examined. Holotype (MNZ EC7489). New

Zealand, Fiordland. Preservation Inlet, exposed rock

surfaces, amongst Squamocnus brevidentis colonies,

2 15 m, J. Duncan, 3 Feb 1998.

Paratypcs. Type locality and date, NMV F82776 (7);

type locality, 30 Jul 1997, NMV F8I955 (4); Stewart

Island, Paterson Inlet, S side of Faith Hope and Charity

Group, 2-4 m, 30 Mar 1998, NMV F83407 (I); 0-5 m,

26 Sep 1998, MNZ EC7490 (2).

Other material. Fiordland, Preservation Inlet,

Feb-Mar 1998, NIWA (8).

Description. Body up to 20 mm long, 6 mm wide

(preserved, tentacles extended); thin calcareous

body wall; body tubular; distinct ventrum, not

sole; mouth anterior, 5 weakly developed radial

oral valves; upturned anal cone, 5 microscopic

anal scales; 10 dendritic tentacles, ventral 2

smaller; calcareous ring solid, slender tapering

anterior projections and posterior indentations

radially and interradially, lacking posterior

prolongations; single left lateral polian vesicle.

Tube feet scattered dorsally and laterally in

midbody, bases slightly swollen, double radial

series near introvert, sometimes also evident in

irregular dorsolateral series; double to zig-zag

rows on ventral radii, extending from introvert to

anus, ventral interradii bare; absent on introvert; 5

radial anal tube feet.

Body wall ossicles cups, buttons, and multi¬

layered ossicles; cups shallow, predominantly

regular, cruciform, some triradiate, frequently

digitate to semidigitate spinelets or fine knobs on

rim pointing in all directions from rim and centre¬

piece, sometimes pointing up only on smallest

cups, always on centrepiece, cups 0.024-0.056

mm long, typically 0.032-0.040 mm long, 4-8,

very rarely up to 20 holes, holes very small

peripherally, smallest cups regularly oval to

rectangular, largest cups less regular around cru¬

ciform centrepiece; buttons predominantly irre¬

gular, 0.06-0.12 mm long, very rarely regular

with as few as 4 holes and 2 central and 8-12

peripheral knobs, commonly 5-7, ranging 3-16

holes, peripheral knobs relatively numerous and

usually discrete and not merging; numerous mul¬

tilayered ossicles up to 0.8 mm long. Introvert

with typical cups, some elongate up to 0.07 mm

long intergrading with some very shallow con¬

cave finely knobbed plates up to 0.07 mm long.

Tentacle ossicles elongate, narrow, curved, pre¬

dominantly bent, some centrally widened, per¬

forated plates up to 0.27 mm long; irregular

convex denticulate perforated plates typically

0.08 mm long; lacking rosettes. Tube feet with

convex endplates up to 0.16 mm diameter, irreg¬

ularly oval thin curved denticulate perforated

plates typically 0.08 mm long, irregularly oval to

elongate narrow centrally-widened curved per¬

orated knobbed plates typically 0.12 mm long,

cups, buttons. Anal ossicles typical body wall

ossicles and large perforated plates; perforated

plates thick, irregularly pear-shaped, part single-

layered not knobbed, part with knobs and sec¬

ondary layers, bluntly denticulate in places at

edges, plates up to 0.56 mm long.

Colour (live). Body and tentacles yellow; dark

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

grey to black markings frequently on tentacle

trunks, anteriorly and posteriorly. Yellow quickly

fades to white in alcohol; dark coloration persists.

Reproduction. Sexes separate; gonad tubules

unbranched; tubules with macroscopically distin¬

guishable eggs up to 0.6 mm long in February; no

evidence of fissipary or brood-protection in

February, July and October.

Etymology. From the Latin luteus (yellow),

referring to the live colour of the body.

Distribution. New Zealand, Fiordland, Preserva¬

tion Inlet; Stewart Island, Paterson Inlet. 0-15 m.

Remarks. After preservation in alcohol specimens

quickly become white, frequently with persistent

anterior and posterior dark grey to black mark¬

ings, and are indistinguishable in appearance

from .S’, niveus. S. luteus is distinguished from S.

niveus by its smaller size and yellow colour (live),

absence of a coelomic brood-protecting habit in

autumn-winter, larger cups frequently with digi¬

tate to semidigitate spinelets. and largest elongate

tentacle plates bent and curved but rarely straight

and curved. It is distinguished from S. brevidentis

by its smaller size and yellow colour (live),

absence of interradial tube feet ventrally, absence

of concave knobbed body wall plates, and pres¬

ence of dark coloration anteriorly and posteriorly.

Ossicle form is consistent from small to large

specimens.

Australocitus gen. nov.

Comparative material examined. Ocnus planci

(Brandt). Western Mediterranean off Banyuls-sur-mer,

50-85 m. NMV F82971 (6) (Figure 5g-l).

Diagnosis. Cucumariidae with thick, firm, friable,

body wall; body rounded to slightly pentagonal in

transverse section; 10 dendritic tentacles, ventral

2 smaller; tube feet in double rows on all 5 radii,

series extending ventrally from introvert to anus,

sparsely present on dorsal and lateral interradii,

absent on introvert. Body wall ossicles abundant

buttons with discrete knobs, cruciform cups, rare

plates; buttons frequently regular with 2 larger

central and 2 terminal holes, 12 peripheral knobs;

cups small shallow regular cruciform, finely

knobbed to spinous, 4—8 holes; rare large flat

elongate to irregularly oval, smooth or knobbed,

perforated, single-layered plates; lacking multi¬

layered ossicles.

Type species. Colochirus calcarea Dendy, 1897.

Species. A. calcareus (Dendy, 1897), A. occiduus

(O’Loughlin and O’Hara, 1992).

Etymology. From the Latin austral meaning

southern, with Ocnus , referring to the southern

hemisphere occurrence of this Ocnus- like genus.

Distribution of genus. Coastal New Zealand

Stewart Island. Australia, Macquarie Islanq

and south-western Australia. Juan Fernandez-

0-433 m.

Remarks. The Ocnus planci specimens examinee)

have a pentagonal body; no interradial tube feet;

irregular very small spinous cups with both crucu

form and triradiate centrepieces and weakly

developed rims with digitate spinelets; and irre,

gular buttons frequently with three or four central

knobs. Based on this material and the description^

of species of Ocnus by McKenzie (1984, 1991),

Australocnus differs from the European genus*

Ocnus Forbes and Goodsir, 1841 (in Forbes) by-

being rounded in transverse section, having somq

tube feet scattered on dorsal and lateral interradii,

and having predominantly regular cruciform cups*

and buttons with two central knobs. Australocnus

differs from the European genus Aslia Rowe,

1970 by having shallow cups with frequently

more than four perforations, not deep cups with

distinct rim and cruciform centrepiece with

unbranched amis. Australocnus differs from

Squamocnus by lacking multilayered ossicles,

and by having elongate buttons which frequently

have 12 small peripheral knobs and two larger

central holes.

Australocnus calcareus (Dendy) comb. nov.

Figure 5a-f

Colochirus calcarea Dendy, 1897: 38-40, pi. 5 figs

44 53.—Farquhar, 1898: 325.

Colochirus brevidentis.— Ludwig, 1898: 442-444,

pi. 26 figs 22-29 [non Squamocnus brevidentis (Hutton,

1872)].

Cucumaria calcarea. —Mortenscn, 1925: 335-337,

fig. 26c-d.—Dawbin. 1950: 38, pi. 2 fig. 11.

Ocnus calcarea, -Panning, 1949: 437.—Pawson,

1970: 39, pi. 1 fig. 5.—Fenwick and Homing, 1980:

443.—McKnight, 1984: 145.

Ocnus calcareus.— Panning, 1949: 438.—Pawson,

1963: 27.—1968: 21.—Panning. 1971: 30, 36.—

O’Loughlin and O’Hara, 1992: 247-248, fig. 7e-f.—

Rowe (in Rowe and Gates, 1995): 279.

Material examined. Neotype (MNZ EC749I). New

Zealand, Cook Strait, Makara, rocky shallows, 0-1 in,

O’Loughlin and Alcock, 20 Sep 1998.

Other material. Neotype locality and date, NMV

F83526 (I); Stewart Island, Paterson Inlet, S side of

Faith Hope and Charity Group, rocky substrate, 0-5 m,

30 Mar 1998, NMV F82782 (2); 26 Sep 1998, MNZ

EC7492 (6), NIWA (4), NMV F83404 (2); Cook Strait,

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Porirua Harbour, Whitireia Headland, rocky shallows, 2

May 1998, NMV F83405 (1) [photograph live NMV

62-8]; Fiordland, Preservation Inlet, Feb-Mar. 1998,

NIWA (2).

Description of material. Body up to 20 mm long,

5 mm wide (live, tentacles fully extended); body

wall friable; rounded dorsolateral and ventrolat¬

eral edges; anal cone upturned, oral end slightly

elevated; 10 dendritic tentacles, ventral 2 smaller;

tube feet in double radial rows ventrally extend¬

ing from introvert to anus, dorsolaterally in radial

double to zig-zag rows with a few present interra-

dially in midbody, absent on introvert, very exten¬

sible single radial tube feet at introvert on ends of

weakly developed oral valves, 5 present anally; 5

microscopic anal scales. Dorsal body wall ossi¬

cles predominantly buttons and cups, rare large

plates; buttons oval, elongate, with discrete

knobs, frequently with 2 central knobs larger than

12 peripheral ones, 2 middle holes larger than end

ones, buttons typically 0.08 mm long; shallow

concave cups with small knobs to blunt spines on

rim and centrepiece, mostly pointing in all direc¬

tions from cup rim, sometimes pointing up only,

cups with predominantly 4 holes (up to 8),

0.024-0.040 mm long; large single-layered plates

rare, irregularly oval to elongate to triangular,

thickened, perforated, knobbed, rarely with

bridges joining knobs to create very' limited sec¬

ondary layer development, up to 0.5 mm long.

Lacking multilayered ossicles and tentacle

rosettes.

Colour (live). Body white; tentacles yellow with

dark brown markings variably evident; no dark

markings anteriorly or anally; dorsal, and anterior

and posterior ventral, tube feet with distal ends

red.

Reproduction. Eggs in gonad tubules 0.7 mm long

in February; gonads present but eggs not evident

macroscopically in September.

Distribution. New Zealand, Slipper Island (37° S,

176° E), Cook Strait, Fiordland, Stewart Island;

Australia, Macquarie Island; Juan Fernandez

Islands. 0-433 m.

Remarks. No reference to type material has been

found for this species. A neotype is established

here from Cook Strait, the locality referred to in

the description of the species. Dendy (1897: 41)

noted an absence of “large reticulate nodules”

(multilayered ossicles) in A. calcareus but recog¬

nised their presence in S. brevidentis. Dendy and

Hindle (1907: 100) considered A. calcareus to be

juvenile specimens of S. brevidentis, noting the

presence of “large calcareus nodules” (multi¬

layered ossicles) in the large specimens but noting

them “very sparingly” in small specimens. Large

multilayered ossicles are always present in small

specimens of S. brevidentis, and the large ossicles

sparsely present in A. calcareus are single-layered

perforated plates. Mortensen (1925: 337) stated

that “larger plates” (not distinguishing perforated

plates and multilayered ossicles) were rare or

absent in A. calcareus, but present in S. brevi¬

dentis. Large multilayered ossicles are never

present in A. calcareus. and this characteristic

provides a diagnostic distinction between Aus-

tralocnus to which A. calcareus is assigned here

and Squamocnus O’Loughlin and O’Hara, 1992

to which S. brevidentis is assigned above.

Panning (1949: 438) was initially uncertain

whether A. calcareus was a species distinct from

S. brevidentis. Subsequently Panning (1971: 38)

reassigned brevidentis to Pentacta but with

reservations. Panning (1971: 30, 36) retained A.

calcareus in a restricted Ocnus.

The Juan Fernandez occurrence is based on a

re-determination as A. calcareus by Mortensen

(1925: 337) of material referred to S. brevidentis

by Ludwig (1898) who considered A. calcareus to

be a junior synonym of S. brevidentis. McKnight

(1984: 145) reported A. calcareus from Mac¬

quarie Island (71-433 m), but the authors were

not able to confirm this record. Mortensen (1925)

reported A. calcareus from Slipper Island.

A. calcareus is distinguished from the Aus¬

tralian species A. occiduus O’Loughlin and

O’Hara by its small size, red dorsal tube feet

when live, rare large perforated plates which are

knobbed, and absence of tentacle rosettes.

O’Loughlin and O’Hara (1992: 248) observed

swollen bases of tube feet (tubercles) in calcareus

specimens, but these were not evident in material

examined subsequently. There is no geographical

continuity for these similar species, neither

occurring in south-eastern Australia.

Australocnus occiduus (O’Loughlin and

O’Hara) comb. nov.

Ocnus calcareus. —Rowe, 1982: 446, fig. 10:32a.—

Marsh, 1991:472,477.—Marsh and Pawson, 1993: 295

[non Australocnus calcareus (Dendy, 1897)].

Ocnus occiduus O’Loughlin and O'Hara, 1992:

247-248, table 1, fig. 7a-d; pis lh, 9a-f.

Plesiocolochirus occiduus.- Rowe (in Rowe and

Oates, 1995): 279.

Material examined. See O’Loughlin and O’Hara, 1992:

247.

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Comparative material. Plesiocolochirus spinosus

(Quoy and Gaimard, 1833), NMV F45021 (3).

Description of material. Body up to 32 mm long

(tentacles withdrawn); rounded to slightly pentag¬

onal in transverse section; thin-walled ventrum;

double radial rows of tube feet extending from

introvert to anus, irregular rows dorsolaterally,

rare small dorsal and lateral ones interradially.

Dorsal body wall ossicles abundant buttons and

cups, rare plates; buttons with discrete knobs,

commonly regular with 12 peripheral knobs

uneven with midlateral and midterminal knobs

often smaller, 2 central knobs not normally larger,

largest buttons typically 0.09 mm long; shallow

cruciform cups with small knobs on rim and

centrepiece pointing in all directions, predomi¬

nantly 4-8 holes, cups typically 0.04 mm long;

large single-layered plates rare, irregularly oval to

elongate, smooth to lumpy, perforated, not

knobbed, up to 0.4 mm long; lacking multilayered

ossicles. Tentacles with rosettes.

Colour (live). White, grey on thin ventral surface

and sometimes laterally; tentacle trunks dark

brown to black; fine tentacle branches pale

yellow.

Distribution. Australia. Rottnest Island, Western

Australia, to Yorke Peninsula, South Australia;

0-30 m.

Remarks. Rowe (1995: 277, 279) referred this

species with reservations to Plesiocolochirus

Cherbonnier, 1946, considering it to be not con¬

generic with Ocnus Forbes, 1841 but possibly

requiring allocation to a new genus. The speci¬

mens of Plesiocolochirus spinosus, the type

species for Plesiocolochirus, have evenly dis¬

tributed radial and interradial tube feet, distinctive

small paired pointed posterior projections on the

radial plates of the calcareous ring, laterovcntral

tubercles, multilayered ossicles, and frequently

bridges across the rim of cups. Australocnus

occiduus has none of these leatures. A. occiduus

is distinguished from the New Zealand A. cal-

careus by its significantly larger size, absence ol

red coloration of dorsal tube feet, rare large plates

which are smooth to lumpy and not knobbed, and

the presence of tentacle rosettes.

Psolidocnus gen. nov.

Figure 61

Diagnosis. Cucumariidae with contiguous body

wall scales dorsally and laterally; body rounded in

transverse section with flattened ventrum, mouth

and anus at least slightly upturned or situated dor¬

sally; 10 dendritic tentacles, ventral 2 smaller;

tube feet in three radial series ventrally, series not

extending to introvert or anus, not continuous

around ventrum to create sole, absent or sparsely

present dorsally; body w r all ossicles large multi¬

layered sometimes towered ossicles (scales) up to

at least 0.8 mm long, irregular lumpy more than

discretely knobbed buttons, regular shallow cruci¬

form spinous cups up to 0.05 mm long with pre¬

dominantly 4 holes, sometimes multiradiate con¬

cave ossicles up to 0.03 mm long.

Type species. Cucumaria amokurae Mortensen,

1925.

Species. P. amokurae (Mortensen, 1925), P. far-

quhari (Mortensen, 1925), P. sacculus (Pawson,

1983).

Etymology. From the family name Psolidae, refer¬

ring to psolid-like characteristics, with Ocnus, the

genus to which two species were referred.

Distribution of genus. New Zealand. South Island;

Stewart Island; Auckland Islands. 9-660 m.

Remarks. Panning (1971: 36) did not include P.

farquhari in Ocnus, but did not reassign it. Paw-

son (1983: 227), O’Loughlin and O’Hara (1992:

237) and Rowe (in Rowe and Gates 1995: 279)

questioned whether Ocnus was a suitable genus

for one or both of the species P. farquhari and P.

sacculus. A pentagonal body form, radial rows of

ventral tube feet extending to introvert and anus,

numerous triradiate cups, and absence of multi¬

layered ossicles distinguish Ocnus (Figure 5g I)

from Psolidocnus.

Panning (1949: 426) assigned P. amokurae to

Trachythyone Studer, 1876 with a question.

O’Loughlin and O’Hara (1992: 237, 239) and

O’Loughlin (1994: 542) noted that Trachythyone

was an unsuitable genus for P. amokurae with its

multilayered ossicles. Panning (1949, 1964) diag¬

nosed Trachythyone as having cups and smooth

plates in the body wall, and the absence of multi¬

layered ossicles and lumpy to knobbed buttons

distinguishes Trachythyone from Psolidocnus. In

Psolidocnus the presence of large contiguous

scales dorsally and laterally, presence of buttons

and cups in the body wall, upturning of or dor¬

sally situated oral and anal ends, and ventral

radial tube feet series not extending to introvert

and anus are psolid-like features, but there is not

a distinct sole with peripheral tube feet as in the

Psolidae. Psolidocnus is distinguished from

Squamocnus and Australocnus by the ventral

radial tube feet series not continuing to the intro¬

vert or anus, by the form of the cups which are

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

predominantly four-holed and up to 0.05 mm

long, by the form of the buttons which are pre¬

dominantly lumpy as distinct from discretely

knobbed, and by the presence dorsally and later¬

ally of macroscopically distinguishable contigu¬

ous sometimes towered scales up to at least 0.8

mm long.

Psolidocnus is similar to Apsolidium O’Lough-

lin and O’Hara, 1992 in having ventral radial

series of tube feet not extending to introvert and

anus, and by having a body wall ossicle combina¬

tion of multilayered ossicles, lumpy to knobbed

buttons, and cups. Psolidocnus differs in lacking

a distinct sole; having significantly fewer tube

feet dorsally, laterally and ventrally; having very

large scale-like contiguous multilayered ossicles;

and having shallow cups. Apsolidium alvei

O’Loughlin and O’Hara, 1992 was assigned to its

genus with reservations since it lacks a distinct

sole, and the large plates with secondary layers

are not typical multilayered ossicles. The form of

the large plates and deep cups of alvei indicate

that it would be no more suitably assigned to

Psolidocnus.

Psolidocnus amokurae (Mortensen) comb. nov.

Figure 6a-g

Cucumaria amokurae Mortensen, 1925: 341-343,

figs 29, 30a.—Dawbin, 1950: 38, pi. 2 fig. 13.

Trachythyone amokurae. —Panning, 1949: 426.—

Pawson, 1970: 38.—O’Loughlin and O’Hara, 1992:

237, 239.—O’Loughlin, 1994: 539-542, fig. I.

Material examined. Three syntypes (ZMUC). New

Zealand, Auckland Islands, Camley Harbour, sandy

mud, 82 m, MPE, 6 Dec 1914; syntype (MNZ EC532),

type locality and date (1).

Other material. Stewart Island, Paterson Inlet, mud,

9-27 m, MPE. 17 Nov 1914, ZMUC (2); Kaikoura, 42°

S, 173° E, 69 m. 7 Dec 1982, MNZ EC6951 (2);

120-130 m, 11 Dec 1982, MNZ EC6907 (2), EC6922

(11), EC7493 (2); 100-112 m, 14 Dec 1982, MNZ

EC6925 (8); 95-98 m, 21 Dec 1982, MNZ EC7494 (1);

Kaikoura, 91 m, 14 Jun 1961, N1WA stn C653 (1,

brood-protecting); 556 m, 18 Oct 1965, NIWA stn

E434 (I).

Distinguishing features. Body up to 17 mm long,

11 mm wide (tentacles withdrawn); body calcare¬

ous, towered contiguous scales dorsally and later¬

ally, smooth contiguous scales ventrally; body

domed dorsally, flat ventrally, mouth antcrodor-

sal, anus posteriodorsal; 10 dendritic tentacles,

ventral 2 smaller; tube feet sparse dorsolaterally

between scales, present in 3 double radial rows on

flattened ventrum, ventral rows not continuing to

introvert or anus, not continuous around ventrum

to create sole, lateroventral row continuous, ven¬

trolateral row discontinuous. Dorsal body wall

ossicles large contiguous multilayered ossicles,

irregularly oval, up to 1.5 nun long, some bluntly

towered at one edge; small, concave, shallow

multiradiate ossicles in epidermis, up to 0.03 mm

long, arms sometimes bifid distally, some joined

at rim to form incomplete cups; concave cruci¬

form oval cups, normally 4, rarely up to 7, perfo¬

rations, knobbed cruciform centrepiece, rim with

semidigitate spinelets pointing up from cup rim,

cups typically 0.04 mm long; lumpy to knobbed

buttons, predominantly irregular, typically 6

(4—10) holes, typically 0.07 (0.07-0.12) mm long.

Colour. Yellow to red (live); white (alcohol)

(Mortensen, 1925).

Reproduction. Brood-protects four juveniles up to

2.2 mm long in dorsal coelomic chamber.

Distribution. New Zealand, Kaikoura; Stewart

Island, Paterson Inlet; Auckland Islands, Camley

Harbour. 9-556 m.

Remarks. O’Loughlin and O'Hara (1992) and

O’Loughlin (1994) remarked that with large

multilayered ossicles this species was not a

Trachythyone , but was close to both Squamocnus

and the Psolidae. O’Loughlin (1994) described

coelomic brood-protection, by a mode similar to

P. sacculus. Within the new genus P. amokurae is

unique in having small concave epidermal multi¬

radiate ossicles in addition to the common ossicle

combination. It is similar to P. farquhari in

having towered dorsal and lateral scales, but is

distinguished by having dorsal tube feet and the

multiradiate ossicles. It is similar to P. sacculus in

its mode of brood-protection, but is distinguished

by having towered scales, dorsal tube feet and

multiradiate ossicles.

Psolidocnus farquhari (Mortensen) comb. nov.

Figure 6h-i

Cucumaria farquhari Mortensen, 1925: 343-345;

figs 30b, 31 a-e—Dawbin, 1950: 38, pi. 2 fig. 12.

Ocnus farquhari .—Panning, 1949: 437.—Pawson,

1970: 39.-1983: 227-229.—O’Loughlin and O’Hara,

1992: 237.

Material examined. Syntype (ZMUC). New Zealand, 2

mi E of North Cape, hard bottom, 100 m, MPE, 2 Jan

1915.

Distinguishing features. Body up to 13 mm long

(extended); body cylindrical, slightly flattened

ventrally, anal cone upturned, mouth slightly

upturned; body calcareous, knobbed scales dor-

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

sally and laterally; 10 dendritic tentacles, ventral

2 smaller; tube feet in double series on 3 ventral

radii, not extending to introvert and anus, not con¬

tinuous around ventrum to form sole, absent dor-

sally and laterally. Dorsal body wall ossicles large

multilayered perforated ossicles (scales), up to 0.8

mm long, with towers; irregular lumpy to

knobbed buttons, 4-8 perforations, frequently

with large central swelling, typically 0.1 mm

long; cruciform concave cups, predominantly reg¬

ular with 4 holes, some with 4 additional small

comer holes, centrepiece lumpy, cups spinous to

digitate to knobbed, spinelets mostly pointing up

from cup rim, cups 0.032-0.048 mm long.

Colour. White, blackish dorsally especially ante¬

riorly and posteriorly (Mortensen, 1925).

Reproduction. Genital tubules unbranched, few

eggs linearly, ‘eggs’ up to 1.0 mm long. A few

ossicles, representing early stages of cup and

button development, were found in one ’egg’

from a gonad tubule, suggesting early embryo

development and vivipary.

Distribution. New Zealand, two miles east of

North Cape; 100 m.

Remarks. Mortensen (1925) reported a blackish

colour dorsally at the anterior and posterior ends.

The syntype examined is white and lacks any

black coloration. P. farquhari is similar to P.

amokurae in having towered scales. It is distin¬

guished from P. amokurae by lacking dorsal tube

feet and multiradiate ossicles. P. farquhari is dis¬

tinguished from P. sacculus by having towered

multilayered ossicles, and double rows of ventral

radial tube feet. The species is known from only

the two type specimens.

Psolidocnus sacculus (Pawson) comb. nov.

Figure 6j-k

Ocnus sacculus Pawson, 1983; 227-230. 2 figs.—

O'Loughlin and O'Hara, 1992: 237.—Rowe (in Rowe

and Gates. 1995): 279.—O’Loughlin, 1994: 540.

Material examined. Two paratypes (USNM E27994).

New Zealand, Taiaroa Canyon, 45°46' S, I71°05’ E.

600-660 m, 25 Mar 1974.

Distinguishing features. Body up to 17 mm long;

body calcareous, contiguous scales dorsally and

laterally; body cylindrical, slightly flattened ven¬

trum. mouth and anus slightly upturned; 10 den¬

dritic tentacles, ventral 2 smaller; tube feet in sin¬

gle rows on ventral radii, zig-zag midventrally.

not extending to introvert and anus, not continu¬

ous around ventrum to create sole, absent dorsally

and laterally except at introvert, single radial tube

feet at introvert. Dorsal body wall ossicles large

thin multilayered ossicles, irregularly oval, up to

1.0 mm long, evident tnacroscopically as scales

[about 9 longitudinally and 9 transversely

(excluding ventrum) in 9 mm long specimen];

lumpy to knobbed buttons, predominantly irregu¬

lar, 3-9 perforations, 0.11-0.17 mm long; cruci¬

form concave spinous cups, predominantly regu¬

lar with 4 holes, some with 4 additional small

corner holes, spinelets pointed to semidigitate,

pointing up only or in all directions from rim, dis¬

tinct cruciform centrepiece sometimes with fine

knobs, cups 0.03-0.05 mm long; ventral ossicles

similar to dorsal.

Colour (live). White (Pawson, 1983).

Reproduction. Gonads comprise long unbranched

tubules, sexes separate, long genital papilla

slightly anterior to dorsal tentacle pair on male

paratype dissected; brood-protects in 3 anterior

dorsal and lateral interradial closed internal

pouches (Pawson, 1983).

Distribution. Taiaroa Canyon, south-eastern New

Zealand; 600-660 m.

Remarks. P. sacculus is similar to P. farquhari in

lacking dorsal and lateral tube feet. It is distin¬

guished from P. farquhari by having multilayered

ossicles without towers, single rows of ventral

radial tube feet, and a closed coelomic sac mode

of brood-protection, it is distinguished from P.

amokurae by lacking dorsal tube feet, lacking

towers on the body wall scales, and lacking

multiradiate ossicles. The species is known from

only the 13 type specimens.

Plesiocolochirus ignavus (Ludwig)

Figure 4j-l

Cucumaria ignava Ludwig, 1874: 81.—Lampert,

1885: 142.—Theel, 1886: 107.

Pentacta australis. —H.L.Clark, 1946: 390, 392

(part) [non Plesiocolochirus australis (Ludwig, 1874)].

Trachythyone ignava - Panning, 1949:426.

Leptopentacta ignava. —Panning, 1966: 60-62, fig.

6 .

Pentacta ignava .—Rowe. 1982: 459, 466, fig.

10.30d.—O'Loughlin and O’Hara, 1992: 227, 229.

Pentacta sp. cf. minute - -Marsh, 1991: 471-472.—

Marsh and Pawson, 1993: 295, fig. 2B.

Plesiocolochirus ignava .- Rowe (in Rowe and

Gates. 1995): 278-279.

Material examined. New Zealand, North Island, Hau-

raki Gulf, off Tiri Tiri Island, 20 m, S. O’Shea, 10 Jun

1992, NMV F82775 (17); South Island, Fiordland,

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Dusky Sound, Beach Harbour, 11-18 m, W.H. Dawbin,

5 May 1950, MNZ EC6705 (1). Southern Australia,

abundant NMV material.

Comparative material. Plesiocolochirus spinosus

(Quoy and Gaimard, 1833), NMV F45021 (3); Lep-

topentacta grisea H.L.CIark, 1938, NMV F82972 (1).

Distinguishing features. Body up to 50 mm long;

body hard, box-like, prominent raised dorsolat¬

eral edges, distinct ventrum not a sole, lateral and

dorsal surfaces with lumps and ridges, smooth

ventrally, oral end with 5 pointed triangular flaps

(valves); 3 double radial rows of tube feet ven¬

trally, zig-zag rows dorsolaterally, all rows

extending from introvert to anus, no tube feet

interradially; dorsolateral tube feet and all tube

feet on tapering oral and anal ends surmount low

wreaths of encircling ridges of ossicles; radial

plates of the calcareous ring with no posterior

prolongations; body wall ossicles large multi¬

layered knobbed perforated scales up to 1.5 mm

long, regular and irregular knobbed perforated

buttons, shallow spinous cruciform cups of a

range of sizes and frequently with bridging

connections across rim.

Colour. Live body colour dorsally and laterally

mottled reddish-brown, brown, cream and white

with dorsally a few bright red markings and

mauve coloration; walls of tube feet orange on all

radii; white ventrally; tentacle trunks dark brown

with some white at bases, branches cream. Pre¬

served colour residual mottled dark reddish

brown on interradial dorsal and lateral surfaces,

white ventrally and on dorsolateral edges.

Distribution. New Zealand, Hauraki Gulf, Dusky

Sound; southern Australia. 0-36 m (Rowe in

Rowe and Gates, 1995).

Remarks. The specimens from Hauraki Gulf have

residual brown coloration, are less than 10 mm

long, and were initially identified as Ocnus brevi-

dentis. S. O’Shea reported (pers. comm.) observ¬

ing this species in north-eastern New Zealand,

and that it had red coloration. Morton and Miller

(1968: pi. 29-4) published a coloured photo of

Ocnus brevidentis but the mottled pale brown

with red and violet colours indicates P. ignavus.

This species is reported here for the first time for

New Zealand. It occurs abundantly in the rocky

shallows across southern Australia.

The generic placement of this species is prob¬

lematic. Plesiocolochirus spinosus (Quoy and

Gaimard), the type species for Plesiocolochirus

Cherbonnier, 1946, has evenly distributed radial

and interradial tube feet, distinctive small paired

pointed posterior projections on the radial plates

of the calcareous ring, and lateroventral tubercles.

P. ignava has none of these features. Leptopen-

tacta grisea H.L.CIark, the type species for Lep-

topentacta , has large posterior prolongations on

the radial plates of the ring, lacks connecting

bridges across the rim of cup ossicles, and lacks

body wall growths on any of the radii. P. ignavus

differs in these characters. In diagnosing Pentacta

Gold fuss, Panning (1971: 38-39) considered

appendages confined to the radii, growths on the

radii surmounted by tube feet or papillae, bridges

across knobbed ossicles, and four- or five-edged

body form to be amongst the characteristics. P.

ignavus does have these characters, but Rowe (in

Rowe and Gates, 1995: 271) has queried the diag¬

nosis of Pentacta. There are affinities and dis¬

crepancies for P. ignavus in relation to Leptopen-

tacta, Plesiocolochirus and Pentacta. A review of

the assignment of P. ignava and related species to

appropriate genera is beyond the scope of this

work.

Psolidiella nigra Mortensen

Psolidietla nigra Mortensen, 1925: 360-362, figs

42—43.—Dawbin, 1950: 35, pi. I fig. 3.—Panning,

1949: 415.-1957: 35.—1961: 192-194, figs 1-6.—

Hickman, 1962: 52.—Pawson, 1970: 36-38, fig. 7.—

Panning, 1971: 45.—-Fenwick and Homing, 1980: 441,

443.—O’Loughlin, 1994: 541-543, fig. 2.—Rowe (in

Rowe and Gates, 1995): 280.

Cucumaria bollonsi Mortensen, 1925: 345-346, fig.

32.—Dawbin, 1950: 38, fig. 9. syn. nov.

Trachythyone bollonsi. —Panning, 1949: 426.—

Pawson, 1970: 38-39.

Material examined. Holotype, 2 paratypes [ZMUC].

New Zealand. Stewart Island. Paterson Inlet, rocky

shore, 6 Apr 1924.

Other material. The Snares, Boat Harbour, covered

by plant detritus in rock pools, 18 Dec 1976, MNZ

EC4061 (10); AM J12281 (4).

Syntype of Trachythyone bollonsi (Mortensen,

1925). North Island, Cape Maria van Diemen, MPE, 4

Jan. 1915, ZMUC Eh 304-258 (1).

Distinguishing features. Body up to 28 mm long

(tentacles withdrawn), 11 mm diameter; body

rounded, distinct oval sole, oral cone and

upturned anal cone; oral cone wrinkled, folded,

pocketed on female specimens; 5 anal scales; 10

dendritic tentacles, ventral 2 smaller. Small tube

feet scattered dorsally, laterally, around oral and

anal cones; tube feet in 3 radial series ventrally,

not continuing to introvert and anus, bordering

sole to varying degrees anteriorly and posteriorly,

up to 4 rows wide lateroventrally, up to 2 rows

wide midventrally; absent from introvert. Dorsal

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

body wall ossicles absent or sparse thick smooth

buttons, typically 4 perforations, up to 0.1 mm

long (tube foot endplates and support plates pre¬

sent dorsally). Ventral body wall ossicles abun¬

dant thick smooth perforated plates, irregularly

oval, up to 16 perforations typically about 8, up to

O. 24 mm long; fewer buttons similar to dorsal

ones; numerous regular and irregular concave

crosses, arms frequently bifid and knobbed dis-

tally, typically 0.06 mm long, rarely partly or

fully closed around rim to form cups.

Colour (preserved). Body pale brown to very

dark brown, grey, black; sole pale brown

to cream; tube feet ends white; introvert and

tentacles dark brown.

Distribution. New Zealand, North Island (Cape

Maria van Diemen), South Island (Banks Penin¬

sula and Otago Peninsula), Stewart Islands and

The Snares; south-eastern Australia; rocky

shallows.

Remarks. O’Loughlin (1994) reported numerous

knobbed and concave crosses and rare cup ossi¬

cles in the type specimens of P. nigra , and Pan¬

ning (1961) illustrated these ossicles for P. nigra

material from Banks Peninsula. Mortensen (1925)

described and illustrated these ossicles for the

type specimens of T. bollonsi , but did not observe

them in P. nigra. The body wall ossicle combina¬

tion of rare cups, concave knobbed crosses,

smooth perforated plates and smooth buttons is

the same in P. nigra and T. bollonsi, as is the ten¬

tacle number and form, dorsal black and ventral

white colour, and habitat niche. The tube foot dis¬

tribution described for T. bollonsi is consistent

with the very small specimens being juveniles of

P. nigra.

The syntype of C. bollonsi which was exam¬

ined was 5 mm long, 2.5 mm in diameter, and a

large part of the dorsal body wall had been

removed. No ossicles were detected, and they

have presumably dissolved from the thin body

wall. The tube feet in the ventral radial series

were large, did not continue to the anus, were four

rows wide laterally and two rows wide mid-

ventrally, and single large ventral interradial tube

feet were present anteriorly and posteriorly creat¬

ing a distinct sole. Smaller tube feet were present

around a slightly upturned anal cone (oral cone

was fully withdrawn). There was some residual

brown colour evident anteriorly, posteriorly and

dorsally. These observations arc also consistent

with the small syntype of T. bollonsi being a

juvenile of P. nigra.

The type specimens of C. bollonsi are small

juveniles, and diagnostic ossicles could no longer

be detected. For these reasons the authors have

chosen to act under the “Principle of the First

Reviser” [ICZN Article 24(a)] and make C. bol¬

lonsi a junior synonym of P. nigra although C.

bollonsi has page precedence over P. nigra in

Mortensen (1925). O’Loughlin (1994) has

described the brood-protecting habit of P. nigra,

and has observed this species in south-eastern

Australia (O’Loughlin, 2000).

Acknowledgments

We are grateful for the assistance with materials

and resources given by Cynthia Aheam (USNM),

Dr Penny Berents (AM), Dr Claus Nielsen

(ZMUC), Ann Morgan and Ann Whittall

(BMNH), Rick Webber (MNZ), Dr Steve O’Shea

(NIWA), Sue Boyd, Tim Stranks and Chris Row-

ley (NMV), Dr Brian Stewart (University of

Otago), and Michael Mannach. We are apprecia¬

tive of the assistance with fieldwork provided by

Project A.W.A.R.E., Fiordland Travel, Divelog

NZ, and Fiordland Ecology Holidays. We are par¬

ticularly grateful to Dr David Pawson and Tim

O’Hara for their assistance and critical review of

the manuscript, and to Ben Boonen for the

presentation of the figures.

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NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Figure 1. a-j, Pseudocnussentus sp. nov. (ossicles from holotype EC7483). a, right lateral body of holotype (27 mm

long); b, midbody section; c, two plates of calcareous ring; d. button (0.08 mm long); e, digitate button (0.10 mm

long); f, digitate button (0.10 mm long); g, digitate plate with secondary layering (0.16 mm long); h, denticulate

multilayered ossicle (0.22 mm long); i, bridged introvert plate (0.09 mm long); j, tentacle rosettes,

k—1, Pseudocnus leoninoides (Mortenscn) (ZMUC syntype), denticulate plates (0.14 mm long).

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Figure 2. Squamocnus brevidentis (Hutton) (ossicles from F81957 and F82779 and N1WA A729). a, lateral body of

F81958 (36 mm long); b, left ventral; c, midbody section; d. two plates of calcareous ring; e, smallest cups (0.03

mm long); f, side of cup (0.03 mm long); g, medium cup (0.06 mm long); h, knobbed concave plate (0.1 mm long);

i, knobbed concave plate (0.1 mm long); j, knobbed concave plate and buttons (0.1 mm long); k, multilayered

ossicle (0.6 mm long); 1, tentacle plate (0.6 mm long).

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Figure 3. Squamocnus niveus sp. nov. (ossicles from paratypes F82777 and F82783). a, dorsal body of holotype,

EC7487 (20 mm long); b, ventral; c, dorsal view of ventral coelomic brood sacs and calcareous ring and empty

gonad upper left (14 mm long specimen, F83406); d, tnidbody section; e, two plates of calcareous ring; f, cup (0.04

mtn long); g, cups (0.05 mm long); h, buttons (0.09 mm long); i, buttons (0.08 mm long); j, multilayered ossicle

(0,6 mm long): k, curved and convex denticulate tentacle plates (0.06 mm long); 1, straight and curved elongate

tentacle plate (0.4 mm long).

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Figure 4. a-i, Squamocnus luieus sp. nov. (ossicles from paratypes F82776). a. left lateral body and tentacles of

holotype, EC 7489 (20 mm long): b. midbody section; c, two plates of calcareous ring; d, cup with semidigitate

spinelets (0.05 mm long); e, cup (0.05 mm long); f, button (0.08 mm long); g, buttons; h. multilayered ossicle (0.8

mm long); i. bent and curved elongate tentacle plate (0.25 mm long).

j-l. Plesiocoloehirus ignavus (Ludwig), j, anterior dorsolateral body of 48 mm long specimen showing radial

tubercle wreaths around tube feet (F76516); k, anterior oral valves; I. bridged cups (F82775).

NEW ZEALAND CUCUMARIIDAE (HOLOTHUROIDEA)

Figure 5. a-f, Australocnus calcareus (Dendy) (ossicles from F82782). a, dorsal body of F83405 (7 mm long); b,

cup (0.035 mm long); c, buttons (0.08 mm long); d, button and cups; e, plate (0.3 mm long); f, plate (0.5 mm long).

g-1, Ocnus planci (Brandt) (F82971). g, dorsal body (40 mm long); h, ventral; i, cups (0.025 mm long); j, cups

(0.025 mm long); k, buttons (0.08 mm long); 1, buttons (0.08 mm long).

P. MARK O’LOUGHLIN AND NICOLA ALCOCK

Figure 6. a-g, Psolidocnus umokurae (Mortcnsen) (ossicles from syntype. ZMUC and NIWA C653 specimens), a,

lateral body of syntype (9 mm long); b. ventral; c, four dorsal coelomic brood juveniles in 12 mm long specimen

(NIWA C653); d, concave multiradiate ossicles (0.02-0.03 mm long); e, multiradiate ossicle and cups (0.02-0.04

nim long); f, button (0.12 mm long); g. towered multilayered ossicle (1.2 mm long).

h-i, Psolidocnus farquhari (Mortensen) (ZMUC syntype). h. cup (0.04 mm long); i, button (0.1 mm long),

j-k, Psolidocnus sacculus (Pawson) (paratype E27994). j, cup (0.05 mm long); k. button (0.11 mni long).

1, Psolidocnus gen. nov. Drawing of ventral tube foot distribution.

Memoirs of Museum Victoria 58(1): 25-37 (2000)

A REVIEW OF THE CUCUMARIID GENUS PSOLID1ELLA MORTENSEN

(ECHINODERMATA, HOLOTHUROIDEA)

P. Mark O’Loughlin

Invertebrate Zoology, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

(moloughlin@parade.vic.edu.au)

Abstract

O’Loughlin, P.M., 2000. A review of the cucumariid genus Psolidiella Mortensen

(Echinodermata, Ilolothuroidea). Memoirs of Museum Victoria 58(1): 25-37.

The holothurian cucumariid genus Psolidiella Mortensen is reviewed. Psolidiella hickmani

sp. nov. and Psolidiella maculosa sp. nov. are described from southern Australia. Psolidiella

adhaerens Hickman is synonymised with Psolidiella nigra Mortensen. P. nigra and P. hick¬

mani are external brood-protecting species. The genus Psolidiella is restricted to New Zealand

and southern Australia. A key is provided for the described species of Psolidiella.

Introduction

The genus Psolidiella Mortensen, 1925 is known

principally from the works of Mortensen (1925),

Panning (1949, 1961, 1971), Hickman (1962),

PawSon (1969, 1970) and O’Loughlin (1994).

The initially monotypic genus Psolidiella was

established for the New Zealand species P. nigra

Mortensen, 1925. Panning (1949) referred both

Thyone speetabilis Ludwig, 1883, from the

Straits of Magellan, and Pseudocolochints mollis

Ludwig and Heding, 1935, from Bouvel Island,

to Psolidiella , but subsequently (Panning, 1957)

referred T. speetabilis to Hernioedema Herouard,

1929. Psolidiella adhaerens Hickman, 1962,

from south-eastern Tasmania, became a third

species referred to Psolidiella. Panning (1971)

finally included only the two species, P. nigra

from New Zealand and P adhaerens from Tas¬

mania, in Psolidiella thus excluding P. mollis but

not reassigning this species.

O'Loughlin (1994) and O’Loughlin and

Alcock (2000) anticipated a synonymy of Psoli¬

diella adhaerens Hickman, 1962, from south¬

eastern Australia, with Psolidiella nigra

Mortensen, 1925, from New Zealand. This syn¬

onymy is confirmed here. Specimens referrable

to Psolidiella have been collected from the rocky

shallows across southern Australia — from Tas¬

mania, Victoria, South Australia and Western

Australia. With a narrow range of variations,

specimens from the far eastern coasts of

Tasmania and Victoria are identifiable as the type

species P. nigra from New Zealand. Specimens

from the Derwent estuary, from both sides of

Bass Strait, and from the coast of South Australia

west to Robe, differ from P. nigra in size and

ossicle distribution and density, and represent a

new and second species. Specimens from the

southern coasts of Western Australia differ again

in details of form of ossicles, in the arrangement

of ventral tube feet, and in colour, and are a new

and third species. Two small specimens from

South Australia are not referrable to the three

species discussed below and indicate the pre¬

sence of a fourth and possibly a fifth species of

Psolidiella.

With the exclusion of P. mollis from Psoli¬

diella by Panning (1971) the genus it is not

circumpolar, as indicated by Pawson (1969),

but is restricted to New Zealand and southern

Australia.

Abbreviations of institutions are as follows:

AM, Australian Museum, Sydney, Australia;

MNZ. Museum of New Zealand, Te Papa Ton-

garewa, Wellington, New Zealand; NMV,

Museum Victoria, Melbourne, Australia; MPE,

Dr Th. Mortensen’s Pacific Expedition

1914-1916; TM, Tasmanian Museum, Hobart,

Australia; WAM, Western Australian Museum,

Perth, Australia; ZMUC, Zoological Museum,

University of Copenhagen, Denmark.

P. MARK O’LOUGHLIN

The following terms arc defined:

Bent and curved. Elongate tentacle and tube

foot plates may be straight or bent (angular) in

one plane (surface) which itself may be flat or

curved or convex (Fig. 3i illustrating straight and

curv ed, and Fig. 4k illustrating bent and curved).

Buttons. Thickened plates which are per¬

forated, regular to irregular in outline, small

(typically 0.08 mm long), discretely knobbed or

lumpy (Fig. 4c) or smooth (Fig. 3f), typically four

perforations.

Capitate. Form of cup spinelets which have a

rounded head and short neck (Fig. 3g).

Crosses. Small ossicles (typically 0.05 mm

long) consisting of four arms radiating from a

common centre or short rod, to varying degrees

bifid and distally knobbed (Figs 2d, 2f), flat or

concave, sometimes intergrading with cups if

concave with some distal ends joined, sometimes

intergrading with multiradiate ossicles.

Cups. Thin plates which are perforated, con¬

cave, shallow (species in this paper) or deep, oval

to rectangular, small (typically 0.04 mm long),

cruciform (Fig. 3g) or tripartite centrepiece, typi¬

cally four large with frequently four small comer

perforations, spinelets on rim and sometimes on

centrepiece, sometimes with bridging connections

across rim.

Denticulate. With pointed to bluntly spinous to

rounded teeth-like projections on the rim of

ossicles (Figs 2h, 3c).

Multilayered ossicles. Ossicles which are nodu¬

lar, irregularly oval in outline, large (more than

0.2 mm long), comprising a perforated knobbed

base plate with additional layers built on both

sides creating low domes bilaterally.

Multiradiate ossicles. Small ossicles (typically

0.05 mm long) consisting of three to five amis

radiating from a common centre or short rod.

including four-armed crosses, fiat or concave,

with the arms sometimes distally bifid and some¬

times knobbed (Figs 4d, 4g).

Sole. Flattened, delimited, modified ventral

body wall, bounded peripherally by tube feet

which do not extend in series to the introvert and

anus (Figs lc, 3b); the place of attachment to the

substrate.

Ventrum. Modified ventral body wall, bounded

laterally by the lateroventral series of tube feet,

not bounded peripherally by tube feet; ventro¬

lateral tube feet series either extend to introvert

and anus, or do not extend to introvert and anus

and create a sole-like ventrum (Fig. 4b); the place

of attachment to the substrate.

Order Dendrochirotida Grube, 1840,

restricted Pawson and Fell, 1965

Cucumariidae Ludwig, 1894

Psolidiella Mortensen, 1925

Psolidiella Mortensen, 1925: 359-360.—Panning,

1949:414-415.—Panning, 1957: 35.—Hickman, 1962;

50, 52.—Pawson, 1969: 37, pi. 22 map 1.—Pawson,

1970: 36.—Panning, 1971: 44—45.—Rowe (in Rowe

and Gates), 1995: 280.

Diagnosis (emended). Psolus-Mke form with thin-

walled body and distinct oval sole or sole-like

ventrum without an acute angular edge; upturned

anal cone; 10 dendritic tentacles, ventral 2

smaller; numerous small tube feet scattered dor-

sally, laterally, around oral and anal cones; tube

feet in 3 radial series ventrally, series not contin¬

uing to introvert and anus, sometimes bordering

sole to varying degrees anteriorly and posteriorly;

absent from introvert; body wall ossicles a com¬

bination of distally knobbed concave crosses or

multiradiate ossicles and sometimes cups typi¬

cally 0.05 mm long, and thickened perforated

irregularly oval buttons and plates less than 0.3

mm long; lacking multilayered ossicles and a

cover of scales; posterior intestine in left ventral

interradius.

Type species. Psolidiella nigra Mortensen, 1925

(monotypy).

Species referred to genus. P. nigra Mortensen,

1925; P. hickmani sp. nov.; P. maculosa sp. nov.

Distribution of genus. North and South Islands of

New Zealand, Stewart Island, The Snares;

southern Australia (Gabo Island to Perth). Rocky

shallows.

Remarks. Mortensen (1925) distinguished his

new genus by its distinct sole and Psolus-Mke

form, and gave significance to the location of the

posterior part of the intestine in the left ventral

interradius by which he distinguished it from a

similar genus Psolidium Ludwig, 1887 and placed

it in the Cucumariidae rather than the Psolidae.

The diagnostic significance of the Psolus-\ikc

form was maintained in subsequent diagnoses of

Psolidiella by Panning (1949, 1957. 1971) and

Pawson (1970). Panning (1949, 1957) continued

to recognize the location of the posterior part of

the intestine as being of diagnostic significance

but this character was then excluded in subse¬

quent diagnoses by Pawson (1970) and Panning

(1971). This original diagnostic character is

retained here as being true for the material exam-

REVIEW OF PSOLIDIELLA (HOLOTHUROIDEA)

ined but without judgement as to its universal reli¬

ability in distinguishing the Cucumariidac from

the Psolidae. It is relevant to note that in their

description of Pseudocolochirus mollis Ludwig

and Heding, 1935, temporarily referred to Psoli-

diella by Panning (1949), the authors reported

and illustrated the posterior part of the intestine in

the right ventral interradius.

All of the southern Australian specimens

referred below to two new species of Psolidiella

are similar to P. nigra in having a well-defined

very thin-walled sole or sole-like ventrum without

an acute angular edge, ventral radial tube feet

series not continuing to the introvert and anus,

small tube feet scattered dorsally and laterally,

up-turned anal cone, and non-calcareous thin

body wall with distally knobbed concave crosses

and small perforated plates but lacking scales.

The three ventral radial series of tube feet do not

continue to the introvert and anus in any of the

three species of Psolidiella. and in the type

species and new species from eastern Australia

tube feet are to varying degrees continuous

around the periphery of a sole. In the species from

western Australia the three ventral series of tube

feet are parallel with none occurring transversely

to create a distinct sole. This condition is referred

to here as a sole-like ventrum.

The presence of distally knobbed concave

crosses and sometimes cups in the body wall was

not reported by Mortensen (1925) or Panning

(1949, 1957) or Hickman (1962) or Pawson

(1970), although Panning (1961: fig. 6) did illus¬

trate distally knobbed crosses for specimens of P.

nigra which he examined. Panning (1971: 45)

referred to the presence of crosses (as retrogres¬

sive cups) in his final diagnosis of the genus. In

the light of a re-examination of type material, and

the availability of further material from southern

Australia referrable to Psolidiella , the presence of

concave crosses and sometimes cups is consid¬

ered here to be diagnostically significant. Distally

knobbed concave crosses are abundant in all

southern Australian material, while cups are pre¬

sent but rare in the type species and abundant in

south-eastern Australian material.

Panning (1949: 426, as bollonsoni) referred the

New Zealand species Cucumaria bollonsi

Mortensen, 1925 to Trachythyone Studer, 1876,

and O’Loughlin and Alcock (2000) have estab¬

lished T. bollonsi as a junior synonym of Psoli¬

diella nigra Mortensen, 1925. Panning (1971: 45)

noted a close relationship between Psolidiella

and Trachythyone. The body wall ossicle combi¬

nation of cups and smooth perforated plates, and

the tentacle form and number, indicate that the

genera are closely related. Psolidiella is distin¬

guished from Trachythyone by the former’s

distinct sole or sole-like ventrum with the

ventral radial tube feet series not continuing to the

introvert and anus, upturned anal cone, and con¬

cave crosses or multiradiate ossicles in the body

wall.

A single small dark grey-brown specimen from

Emu Bay on Kangaroo Island in South Australia

(NMV F59223), and single small dark blue-grey

specimen from Waterloo Bay on the Eyre Penin¬

sula in South Australia (NMV F59224), have the

body form of Psolidiella and have numerous con¬

cave, distally knobbed, multiradiate ossicles dor-

sally and ventrally. Neither has cups dorsally or

ventrally, and neither has buttons dorsally. They

do not belong to any of the three species

described below, and these South Australia

specimens represent one or two more species of

Psolidiella.

Key to species of Psolidiella

1. Dorsal body wall normally with tube foot ossicles only; knobbed crosses and

buttons absent or rare in dorsal body wall. Psolidiella nigra Mortensen

Dorsal body wall normally with tube foot ossicles, numerous crosses or

multiradiate ossicles, and numerous thickened buttons or plates.2

2. Dorsal body wall with numerous knobbed crosses, cups, smooth buttons and

plates; lacking rosettes; midventral radial tube feet series up to two wide;

lateral ventral radial series of tube feet joined to varying degrees anteriorly

and posteriorly to form a distinct sole. Psolidiella hickmani sp. nov.

Dorsal body wall with numerous crosses and multiradiate ossicles, lumpy

buttons and rosettes; lacking cups; midventral radial tube feet series up to

four wide; lateral ventral radial series of tube feet parallel and never joined

anteriorly and posteriorly to form a distinct sole.

. Psolidiella maculosa sp. nov.

P. MARK O’LOUGHLIN

Psolidiclla nigra Mortensen

Figures 1 a-d, f-k, 2a-l

Psolidietla nigra Mortensen, 1925: 360-362, figs 42,

43.—Panning, 1949:415.—Dawbiti, 1950: 35, pi. 1 fig.

3.—Panning, 1957: 35.—Panning, 1961: 192-194, figs

1-6.— Hickman. 1962: 52.— Pawson, 1969: 37, pi. 22

map 1.—Pawson. 1970: 36-38, fig. 7. —Panning, 1971:

45.—Fenwick and Homing, 1980: 441, 443.—

O’Loughlin, 1994: 541-543, fig. 2.—Rowe (in Rowe

and Gates) 1995: 280 --O’Loughlin and Alcock, 2000:

3, 15-16.

CUcumaria bollonsi Mortensen, 1925: 345-346, fig.

32.—Dawbin. 1950: 38, fig. 9.

Trachylhvone bollonsi. —Panning, 1949: 426.-—Paw¬

son, 1970: 38-39.

Psaiidiella adhaerens Hickman, 1962: 50-52, figs I-

16, pi. 1 fig. 1. Pawson, 1969: 37. pi. 22 map 1.—

Pawson, 1970: 36.—Panning. 1971: 45.—Dartnall.

1980: 12, 51. map 16(1).-Rowe. 1982: 458. 464.

Rowe and Vail, 1982: 222 (part).—O'Loughlin, 1984:

151 (part).— O’Loughlin and O’Hara, 1992: 227,

229.—O’Loughlin, 1994: 543-544, figs 3, 4 (part).—

Rowe (in Rowe and Gates) 1995: 280 (part), syn. nov.

Material examined. Holotype, 2 paratypes. New

Zealand. Stewart Island, Paterson Inlet, rocky shore,

S. Vallin, 6 Apr 1924 (ZMUC).

Other New Zealand material. The Snares, Boat Har¬

bour. covered bv plant detritus in rock pools, 18 Dec

1976, MNZ EC4061 (10): AM J12281 (4).

Syntype of Cucumaria bollonsi Mortensen, 1925.

North Island. Cape Maria van Diemen, MPE, 4 Jan

1915, ZMUC Eh 304-258(1).

Syntypes of Psolidietla adhaerens Hickman, 1962.

Australia, Tasmania, Pirates Bay, Eaglehawk Neck,

under stones near low tide level. V.V. and J.L. Hick¬

man. 19 Mar 1954. TM H2135 (I): 13 May 1954, H546

(5), H2008 (64), H2009 (microslide, ossicles), H2010

(microslide, ossicles); 9 Mar 1960, H2011 (microslide,

body mount).

Other material. Tasmania. Eaglehawk Neck. 22 May

1959. H2I36 (11); Adventure Bay. 5 Sep 1993, H2310

(1). Victoria. Gabo Island, rocky shallows, 16 May

1982, NMV F68267 (1); Mallacoota, rocky shallows,

21 Jan 1981, F68264 (1); Cape Conran, rocky shallows,

19 Apr 1976, F73809 (1).

Description of material. Body up to 36 mm long

(preserved, tentacles extended), 11 mm diameter;

body wall thin; body rounded, distinct very thin-

walled oval sole with rim, oral cone and upturned

anal cone; on female specimens oral cone wrin¬

kled, folded, pocketed, pockets sometimes

extending dorsally and laterally, rarely on male

specimens; 5 small spatulate anal scales; 10 den¬

dritic tentacles, ventral 2 smaller; calcareous ring

lacking posterior prolongations, 5 radial plates

with anterior notched taper and posterior notch, 5

interradial plates with anterior pointed taper and

posterior broad rounded indentation; single left

ventrolateral polian vesicle; posterior intestine in

left ventral interradius.

Close cover of small tube feet scattered dor-

sally, laterally, around oral and anal cones; tube

feet in 3 radial series vcntrally, not continuing to

introvert and anus, bordering sole to varying

degrees anteriorly and posteriorly, up to 4 rows

wide iateroventrally (paired double to zig-zag

rows), up to 2 rows wide midventrally; absent

from introvert; 5 radial tube feet anally.

Dorsal body wall predominantly with tube foot

ossicles only, rarely with a few buttons (in juve¬

niles), very rarely with a few concave knobbed

crosses or cups (in juveniles): tube foot ossicles

endplates, frequently fragmentary, up to 0.16 mm

wide, some with a narrow rim of support plates;

tube foot support plates irregularly oval to elon¬

gate, sometimes bent and curved, perforated with

large holes centrally, bluntly denticulate around

margin, typically 0.10 mm long, never thickened

rods; thick, smooth, irregularly oval buttons

rarely present dorsally, up to 8 (typically 4) per¬

forations, rarely marginally knobbed, frequently

incompletely formed, typically 0.10-0.16 mm

long.

Ventral body wall with perforated plates, but¬

tons, very rare spinous plates, knobbed concave

crosses, rare cups: plates abundant, thick, smooth,

rarely lumpy, irregularly oval, typically about 8

(up to 17) perforations, frequently thickly bluntly

denticulate around margin, up to 0.24 mm long;

thick plates intergrade with smaller dorsal body

wall type buttons; very rare irregular, spinous

(surface), marginally denticulate, perforated

plates; numerous shallow concave crosses,

regular and irregular, arms frequently bifid

and distally knobbed, knobs sometimes joined

to form incomplete cups, 0.04-0.12 mm long;

rare thick to thin shallow concave cups, rim

smooth to capitate, 0.04-0.10 (typically 0.06) mm

long.

Ventral tube feet with endplates up to 0.32 mm

wide, normally with thin rim of support plates,

typical ventral body wall plates: support plates

irregularly oval to elongate, frequently bent and

curved marginally denticulate, largest perfora¬

tions centrally, up to 0.19 mm long, very rarely

rod-like.

Tentacle ossicles numerous large thick per¬

forated narrow elongate plates and rods, straight

or bent, flat or curved, rarely branched, up to 0.50

mm long; some curved to convex, marginally

denticulate, perforated, irregularly oval to trian¬

gular, smooth plates up to 0.26 mm wide; bent

REVIEW OF PSOLIDIELLA (HOLOTHUROIDEA)

curved narrow denticulate perforated plates

0.08-0.18 mm long; thin perforated marginally

denticulate, irregularly round, convex plates

0.05-0.08 mm wide; sometimes rosettes up to

0.12 mm long.

Colour (preserved). Dorsal and lateral body off-

white to pale brown to very dark brown, grey,

black; oral and anal cones frequently darker than

dorsally; sole off-white to pale brown; tube feet

ends white; introvert and tentacles off-white to

dark brown.

Reproduction. Externally brood-protects differen¬

tiating embryos, up to 0.8 mm long, in pockets

and folds around the oral cone of female speci¬

mens (December to May; no June to August

material observed). Exhibits sexual dimorphism,

mature female specimens having pockets and

folds around the oral cone, mature male speci¬

mens normally having a smooth oral cone. The

three type specimens from Stewart Island (April)

are all males with a smooth oral cone. The speci¬

mens from The Snares (December) include one

brood-protecting female. All Eaglehawk Neck

lots (March, May) include specimens which have:

mature male and female gonads; a gonopore on

the oral disc anterior to the middorsal tentacle

pair; gonads with eggs ranging from small white

to large cream, up to 0.6 mm long; mature

females with distinct folds and pockets and small

tube feet around the oral cone; mature males with

smooth oral cones and some larger males with

some lesser degree of anterior pocketing; at least

one female with brood embryos in the anterior

pockets, one specimen (H2008) with about 100

brood embryos; some brood embryos showing the

beginning of differentiation, having numerous

crosses; brood embryos which are 0.6-0.8 mm

long. The Cape Conran specimen (April) is

brood-protecting.

Distribution. New Zealand, North Island (Cape

Maria van Diemen), South Island (Banks Penin¬

sula and Otago Peninsula), Stewart and The

Snares; far eastern coast of southern Australia

(Gabo Island, Mallacoota, Cape Conran, Eagle-

hawk Neck, Adventure Bay). Rocky shallows.

Remarks. O’Loughlin (1994: 544) and O’Lough-

lin and Alcock (2000) anticipated that P.

adhaerens might be a junior synonym of P. nigra.

A synonymy is formally established here. Hick¬

man (1962) based his new species P. adhaerens

on abundant material which was collected exclu¬

sively from Eaglehawk Neck on the eastern coast

of southern Tasmania, and distinguished P.

adhaerens from P. nigra on the grounds of differ¬

ing colour, possessing anal teeth, and lacking

ossicles in the walls of the tube feet. The type

material of both species has small anal teeth, and

the ventral and at least some dorsal tube feet have

a thin rim of support ossicles. The colour is vari¬

able, and material from New Zealand and eastern

Australia exhibits a similar colour range. Neither

Mortensen (1925) nor Hickman (1962) reported

the rare small concave distally knobbed crosses

and very rare cups which are present in the body

wall of the type material of both P. nigra and P.

adhaerens. The form and distribution and size of

the ossicles, the external morphology and size,

and the mode and seasonality of the distinctive

external brood-protective reproduction arc indis¬

tinguishable for the New Zealand and Australian

material referred to above.

Psolidiella hickmani sp. nov.

Figures ld-f, 3a—1

Psolidiella adhaerens .—Rowe and Vail, 1982: 222

(part).—O’Loughlin, 1984: 151 (part).— O’Loughlin,

1994: 541 (part), 543-544 (part), fig. 4.—Rowe (in

Rowe and Gates), 1995: 280 (part).

Material examined. Holotype. Victoria, Phillip Island,

Kitty Miller Bay. silt, rocky shallows, 0-1 m, J. Mona-

glc and M. O’Loughlin, 20 Apr 1987 (NMV F74912).

Paratypes. Type locality and date, NMV F68266

(14); 25 Oct 1987, F68259 (2); 23 Dec 1987, F68257

(4); 13 Feb 1988, F68260 (6); 22 Nov 1997, F65928

(7).

Other material. Victoria. Wilsons Promontory,

Obcron Bay, 7 Mar 1984. F6827I (I ): Cape Paterson.

29 Jan 1988, F68254 (4); Kilcunda, 26 Jan 1987.

F73815 (1); Harmers Haven, 27 Mar 1989, F57357 (1);

Phillip Island, Cowrie Beach. 27 Nov 1985, F68263

(11); Flinders, 16 Nov 1980, F68268 (I); Port Phillip

Bay. Altona. 8 Apr 1991. F7382I (2); Marengo, Hay ley

Point, 26 Mar 1977, F68253 (42); Crayfish Bay, 31 Dec

1980, F68270 (I); Armstrong Bay, 5 Apr 1998, F82973

(2). South Australia. Cape Northumberland, 16 May

1990. 1*74908 (1); Beachport, 9 Jan 1988, F74906 (5);

Robe. 10 Jan 1988, F74907 (2). Tasmania. Derwent

Estuary, Opossum Bay, 15 Nov 1982. F68258 (8); Bass

Strait, Lulworth, 22 Nov 1982, F74910 ( 2); mouth of

Tamar River, Greens Beach. 7 Mar 1981, F74911 (1).

Description of material. Body up to 17 mm long

(preserved, tentacles withdrawn), 11 mm diame¬

ter; body wall thin; body rounded, distinct very

thin-walled oval sole with rim, oral cone and

upturned anal cone; on female specimens oral

cone wrinkled, folded, pocketed, pockets some¬

times extending dorsally and laterally; 5 small

P. MARK O’LOUGHLIN

spatulate scales anally; 10 dendritic tentacles,

ventral 2 smaller; calcareous ring lacking pos¬

terior prolongations, 5 radial plates with anterior

notched taper and posterior notch, 5 interradial

plates with anterior pointed taper and posterior

broad rounded indentation; single left ventro¬

lateral polian vesicle; posterior intestine in left

ventral interradius.

Close cover of small tube feet scattered dor-

sally, laterally, around oral and anal cones; tube

feet in 3 radial series ventrally, not continuing to

introvert and anus, bordering sole to varying

degrees anteriorly and posteriorly, up to 4 rows

wide lateroventrally (paired double to zig-zag

rows), up to 2 rows wide midventrally; absent

from introvert; 5 radial tube feet anally.

Dorsal body wall ossicles crosses, cups, thick¬

ened plates, thin spinous plates, tube feet end-

plates and support ossicles: crosses numerous,

concave, some irregular and multiradiate, amis

frequently bifid and knobbed distally, knobs

sometimes joined to create incomplete rim or cup,

typically 0.04-0.06 mm long; cups, regular, shal¬

low, concave, rim smooth or with capitate

spinelets, typically 0.05-0.06 mm long, some

cups thick and large; numerous thickened smooth

perforated plates, irregularly oval, frequently

with small rounded denticulations marginally, up

to 16 perforations, up to 0.24 mm long, intergrade

with four-holed buttons; rare thin irregular perfo¬

rated plates, surface spines, marginally strongly

denticulate, up to 18 perforations, up to 0.19 mm

long.

Ventral body wall ossicles crosses, cups, thick¬

ened plates: rare knobbed crosses; abundant cups,

rim complete or incomplete, predominantly capi¬

tate spinelets on rim, cups typically 0.05-0.07

mm long; abundant thickened perforated plates

similar to dorsally, up to 14 perforations, up to

0.22 mm long, intergrade with rare buttons.

Tube feet ossicles cndplates, tube foot support

plates and rods, body wall ossicles: endplates up

to 0.32 mm diameter, fairly irregular perforations,

smallest centrally; support ossicles frequently

curved or bent thick perforated rods and narrow

elongate plates, up to 0.30 mm long.

Tentacle ossicles thickened, elongate, fre¬

quently bent and curved, perforated rods up to

0.32 mm long; frequently bent and curved elon¬

gate narrow perforated plates up to 0.22 mm long;

irregularly round, marginally denticulate, con¬

cave, thin perforated plates typically 0.08-0.12

mm diameter; some rosettes up to 0.07 mm loni>.

Colour (preserved). Dorsal and lateral body off-

white to pale brown to very dark brown, grey,

black; oral and anal cones frequently darker than

dorsally; sole off-white to pale brown; tube feet

ends white; introvert and tentacles dark brown.

Reproduction. Externally brood-protects differen¬

tiating embryos, up to 0.5 mm long, in pockets

and folds around the oral cone of female speci¬

mens (October to April). Exhibits sexual dimor¬

phism, mature female specimens having pockets

and folds around the oral cone, mature male spec¬

imens normally having a smooth oral cone. Lots

from Oberon Bay, Cape Paterson, Kitty Miller

Bay, Marengo, Armstrong Bay and Opossum

Bay all include at least one externally brood-

protecting female, one female from Opossum Bay

(November) having about 40 brood embryos in

oral pockets.

Distribution. Australia, Tasmania (Opossum

Bay in Derwent Estuary, and Bass Strait coast);

Victoria (west of Wilsons Promontory) to South

Australia (west to Robe). Rocky shallows, 0-1 m.

Etymology. Named in recognition of the contribu¬

tion to our knowledge of Tasmanian holothurians

by the late Prof. V.V. Hickman (formerly of the

University of Tasmania).

Remarks. Psolidiella hickmani differs consis¬

tently from P. nigra by having numerous thick,

smooth, perforated plates and buttons, and numer¬

ous knobbed concave crosses, in the dorsal body

wall. No specimens of P. hickmani from Victoria

have been observed with more than three brood

embryos in pockets, while one from southern Tas¬

mania has about 40 brood embryos in pockets.

Specimens from southern Tasmania are up to 17

mm long (tentacles withdrawn) and from coastal

Victoria are up to 14 mm long (tentacles with¬

drawn). Although based on limited evidence,

these observations reflect a pattern previously

noted by Materia et al. (1991) for the cucumariid

holothurians Staurothyone inconspicua (Bell,

1887) and Neoamphicyclus lividus Hickman,

1962. In S. inconspicua more numerous co-

elomic juveniles are brood-protected in south¬

ern Tasmanian specimens that in Victorian

specimens, and in both species mature adults in

southern Tasmania are larger than on coastal

Victoria.

Psolidiella maculosa sp. nov.

Figures Id, f, 4a I

Psolidiella sp. Marsh, 1991: 473, 477.

Materia! examined. I lolotype. Western Australia, Perth,

Marmion Marine Park, Mettams Pool, sandy pockets on

reef flat, midtidal, B.R. Wilson, 3 Jan 1987 (WAM

Z4781).

REVIEW OF PSOLID1ELLA (HOLOTHUROIDEA)

Paratypcs. Type locality and date. WAM Z4782 (2);

Cottesloe, reef off Rosendo Street, in sand amongst

Caulerpa holdfasts, 0-1 m. 6 Feb 1985, WAM Z4931

(13); Trigg Island, with Cauterpa, 22 Nov 1969, NMV

F59222 (2); Quaranup, boat ramp, intertidal sand near

Posidonia, 20 Jan 1988, WAM Z4932 (1).

Other material. Western Australia, Duke of Orleans

Bay, Little Wharton Beach, off algae, 0-2 m, 19 Dec

1991, NMV F66253 (I).

Description of material. Up to 27 trim long (pre¬

served. tentacles withdrawn, anal cone extended),

8 mm diameter; body wall thin; oral extension

and anal cone upturned; oral cone lacking folds

and pockets; 10 dendritic tentacles, ventral 2

smaller; distinct introvert, lacking tube feet; dis¬

tinct sole-like ventrum, narrower than body, not

extending to introvert or anus; 5 microscopic

spatulate anal scales; calcareous ring solid, lack¬

ing posterior prolongations, anterior tapering pro¬

jections radially and interradially, posterior

notches radially, wide indentations interradially;

single left ventrolateral polian vesicle; posterior

intestine in left ventral interradius.

Close cover of large and small tube feet scat¬

tered dorsally and laterally and around oral and

anal cones; 3 parallel radial series of tube feet

ventrally, series up to 4 irregular rows wide on

lateral ventral radii, irregular 2-A rows wide on

the midventral radius, radial series not extending

to introvert and anus, rows not joined anteriorly

and posteriorly to border a distinct sole; paired

radial tube feet at introvert; 5 radial tube feet

anally.

Dorsal body wall with epidermal multiradiate

ossicles and crosses, buttons, rosettes, tube foot

ossicles: numerous multiradiate ossicles includ¬

ing crosses, frequently concave, frequently irreg¬

ular, 3-5 arms frequently branched and distally

knobbed, up to 0.06 (typically 0.04) mm long;

numerous buttons, oval to irregular, predomi¬

nantly 4-7 holes, frequently with thickenings or

lumps or knobs, buttons up to 0.17 (typically

0.13) mm long; numerous rosettes up to 0.07 mm

long; cups not present.

Ventrally multiradiate ossicles and crosses sim¬

ilar to dorsally; rosettes; buttons more irregular

than dorsally, intergrading with small thick lumpy

perforated plates, up to 12 holes, frequently

slightly knobbed marginally, similar size to

dorsally, perforations smaller; cups not present.

Tube feet with endplates up to 0.28 mm wide,

perforations frequently smaller centrally; elon¬

gate, narrow, thick, straight or bent, sometimes

branched, perforated support rods up to 0.28 mm

long; irregular elongate perforated support plates,

straight to bent, frequently curved, up to 0.18 mm

long.

Tentacles with abundant plates and elongate,

narrow ossicles, thickened, irregular, perforated,

straight or bent, flat or curved, up to 0.45 mm

long; convex perforated plates, irregularly oval to

rectangular, typically 0.07-0.13 mm long,

marginal perforations smallest; abundant rosettes,

typically 0.05 mm long.

Colour (preserved). Body dark blue-grey to

brown with white spots, sometimes pale ven¬

trally; tube feet ends white; small white calcare¬

ous spots around tube feet, sometimes between

tube feet dorsally and laterally.

Reproduction. The material, collected in Novem¬

ber to February lacks mature female gonads.

Long, thin, smooth, white gonad tubules are pre¬

sent. There are no pockets or folds around the oral

cone of any specimens in the material examined.

Distribution. South-western Australia, from Duke

of Orleans Bay to Perth in Western

Australia; 0-2 m.

Etymology. From the Latin maculosa (spotted),

referring to the body appearance.

Remarks. A body wall ossicle combination of

numerous distally knobbed crosses and multira¬

diate ossicles, lumpy buttons and rosettes, and an

absence of cups, distinguishes this species from

P. nigra and P. hickmani. Distinctive small white

spots in the dark body wall (in addition to the

white tube feet ends, and created by aggregations

of ossicles), midventral tube feet series up to four

rows wide, and the absence of any interradial tube

feet anteriorly and posteriorly delineating a dis¬

tinct sole, also distinguish P. maculosa from the

other two species. There are no mature female

specimens in the material available, and it is not

possible to determine whether or not P. maculosa

has the same external brood-protecting habit as

P. nigra and P. hickmani.

Acknowledgements

I am grateful for the assistance, variously with

fieldwork and materials and resources, provided

by Dr Margot Jensen (ZMUC); Helen Clark

(MNZ); Dr John L. Hickman (formerly of the

University of Tasmania); Alison Green and Eliz¬

abeth Turner (TM); Loisette Marsh, Shirley

Slack-Smith and Dr Jane Fromont (WAM); Sue

Boyd, Tim Stranks and Chris Rowley (NMV);

and Susan Lockhart, Jane Monagle, Christine

Rowland and Peter Tutera. I am particularly

grateful to Dr David Pawson and Tim O’Hara for

their helpful comments, and to Ben Boonen for

the presentation of the figures.

P. MARK O’LOUGHLIN

References

Bell, F.J., 1887. Studies in the Holothuroidea. — VI.

Descriptions of new species. Proceedings of the

Zoological Society of London 35: 531-534, pi. 45.

Dartnall, A., 1980. Tasmanian echinoderms. Fauna of

Tasmania Handbook 3: 82 pp., 36 figs, 5 pis, 18

maps. University of Tasmania: Hobart.

Dawbin, W.H., 1950. A guide to the holothurians of

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REVIEW OF PSOLID1ELLA (HOLOTHUROIDEA)

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invertebrates of southern Australia. Part 1. South

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antarktischen Mecre und zwei neue Seeigel

von den Papua-Inseln, gesammelt auf der Reise

SMS Gazelle urn die Erde. Monatsberichte

der Koniglichen Preussischen Akademie der

Wissenschaften zu Berlin. Pp. 452—465.

P. MARK O’LOUGHLIN

Figure 1. a-c, views of Psolidiella type species P. nigra (H2135, 26 mm long) with oral cone brood embryos, a,

dorsal; b, lateral, showing oral and anal cones; c, ventral, showing sole with two rows of niidventral tube feet,

d, cucumariid calcareous ring of Psolidiella species, e, brood-protection of embryos (white, round, 0.5 mm long)

around oral cone of P. hickmani (F68258, 15 mm long), f, dorsal view of alimentary canal of Psolidiella species

from mouth (m) to anus (a) with posterior intestine left (1) ventral, g-k, body mount showing form and distribution

of ossicles of Psolidiella type species P. nigra (H201I). g, dorsal; h, ventral interradial; i, ventral tube feet; j,

tentacle trunk; k, tentacle ends.

REVIEW OF PSOLIDIELLA (HOLOTHUROIDEA)

OSor

s&QdRSs

oO'

*XJs

Figure 2. a-l, Psolidiella nigra Mortensen. a, lateral view of holotype (ZMUC, 24 mm long), b—1, ossicles

(measurements of lengths), b, ventral plate (0.19 mm, paratype); c, ventral buttons and plates (0.13—0.20 mm,

paratype); d, ventral crosses (0.06-0.08 rnrn, EC4061); e, ventral cups (0.06 mm, EC4061); f, ventral crosses

(0.05-0.06 mm, H2008); g, ventral tube foot support plates and endplate (endplate 0.32 mm, H2009); h, tentacle

plate (0.24 mm, holotype); i, curved tentacle plate (0.13 mm, paratype); j, elongate tentacle plate (0.15 mm,

EC4061); k, elongate tentacle plate (0.26 mm, EC4061); I, tentacle rosettes (0.10-0.12 mm, holotype).

P. MARK O’LOUGHLIN

,jDCJ m

Figure 3. a—1, Psolidiella hickmani sp. nov. a, lateral view of holotype (11 mm long), b, ventrolateral view of

specimen showing sole with two rows ol midventral tube feet (F68258, 16 mm long), c 1, ossicles from types

(measurements of lengths), c, dorsal plate (0.21 mm) and crosses (0.05 mm ); d, dorsal button (0.13 mm) and crosses

(0.05-0.06 mm); e, dorsal spinous plate (0.17 mm); f, dorsal buttons (0.11-0.13 mm); g, dorsal cups (0.05 mm); h,

dorsal tube foot endplate (0.11 mm); i, dorsal tube foot support plate (0.12 mm); j, ventral plates (0.14-0.18 mm);

k, ventral tube foot ossicles (plate 0.13 mm, cross 0.06 mm); I, tentacle ossicles (0.06-0.24 mm).

REVIEW OF PSOllDlELLA (HOLOTHUROIDEA)

Figure 4. a-1, Psolidiella maculosa sp. nov. a, lateral view of holotype (19 mm wide shown); b, ventral view of

paratype showing sole-like ventrum with four rows of midventral tube feet (18 mm shown), c—1, ossicles from types

(measurements of lengths), c, dorsal buttons (0.10-0.15 mm); d, dorsal multiradiate ossicles (0.03 mm); e, dorsal

rosette (0.07 mm); f, ventral plates (0.09-0.15 mm); g, ventral multiradiate ossicles (0.04-0.05 mm); h, ventral tube

foot plate (0.18 mm) and rod (0.24 mm); i, ventral tube foot ossicles (button 0.11 mm); j, tentacle plate (0.16 mm);

k, tentacle rod (0.32 mm); I. convex tentacle plate (0.06 mm) and rosette.

Memoirs of Museum Victoria 58(1): 39-75 (2000)

A REVIEW OF THE TUBIFICIDAE (ANNELIDA: OLIGOCHAETA) FROM

AUSTRALIAN INLAND WATERS

Adrian M. Pinder 1 and Ralph O. Brinkhurst 2

'Department of Conservation and Land Management, Wildlife Research Centre,

PO Box 51, Wanneroo, Western Australia 6946, Australia (adrianp@calm.wa.gov.au)

2 205 Cameron Court, Hermitage, Tennessee 37076, USA (OUgol@aol.com)

Abstract

Pinder, A.M. and Brinkhurst, R.O., 2000. A review of the Tubificidae (Annelida: Oligochaeta)

from Australian inland waters. Memoirs of Museum Victoria 58(1): 39-75.

The Australian limnic representatives of the cosmopolitan oligochaete family Tubificidae

are reviewed. The four Australian species of Telmatodrilus Eisen, 1879 are placed in new

genera ( Breviatria and Biprostatus ) and these are considered to belong to Rhyacodrilinae

rather than Telmatodrilinae. Four new species, Rhyacodrilus megaprostatus. Aimidrilus

nharna, Breviatria arvensis and Biprostatus duplex are described and Rhyacodrilus fultoni

Brinkhurst, 1982 is transferred to Aimidrilus Finogenova, 1982. Limnodrilus daparedeianus

Ratzel, 1868 and Aulodriltts limnobius Bretscher. 1899 are recorded in Australia for the first

time. The Australian fauna now consists of 29 described species, including 11 of the most

widespread species of cosmopolitan genera ( Tubifex Lamark, 1816. Potamothrix Vejdovsky

and Mrazek, 1902, Limnodrilus Claparede, 1862, Aulodrilus Bretscher, 1899, Rhyacodrilus

Bretscher. 1901, Bothrioneurum Stoic, 1886 and Branchiura Beddard, 1892). Antipodrilus

Brinkhurst, 1971, with five species, and the two new rhyacodriline genera, with six species,

are essentially Australian genera, except that one species of Antipodrilus and possibly one of

Biprostatus also occur in New Zealand. Seven endemic species belong to more widespread

genera ( Rhizodrilus Smith. 1900, Rhyacodrilus and Aimidrilus). The Australian tubificid fauna

appears to have biogeographic affinities with northern hemisphere and marine tubificids.

Introduction

The first tubificid to be described from Australia

was Branchiura pleurotheca Benhatn, 1907 from

Blue Lake. Mt Kosciusko. Further tubificid

records and descriptions from Australia were not

published until Brinkhurst (1971a, b) reviewed

the aquatic oligochaete faunas of the world

and Australia. In these publications, Branchiura

pleurotheca was synonyntised with Rhyacodrilus

coccineus (Vejdovsky, 1875), several other new

species and a new genus ( Antipodrilus ) were

described and the presence of numerous cos¬

mopolitan species was recorded. Subsequent

records and descriptions of new species w'ere pro¬

vided by Brinkhurst and Fulton (1979) and

Brinkhurst (1982, 1984).

In this paper we list previous records and add

many new ones for most species, improve some

descriptions, describe four new species and

address problems relating to the taxonomic

position of Australian species of Telmato¬

drilinae by moving them to two new rhyaco¬

driline genera. This contribution is intended to

provide a foundation for future work, comple¬

menting the recent review of the Australian

Phreodrilidae (Pinder and Brinkhurst, 1997). Our

definition of the Tubificidae is conservative,

excluding the Naididae which are generally

acknowledged to be part of the same taxon

(Brinkhurst, 1994; Erseus, 1990b), although a

formal proposal to merge the families has yet to

be made.

Materials and methods. Most specimens were

stored in ethanol, often after fixation in formalin.

They were then whole-mounted, dissected or

serial sectioned. Dissected or whole-mounted

specimens were usually first stained in

Grenacher’s borax carmine. Sectioned speci¬

mens were cut at 6 mm and stained in Mayer’s

haematoxylin and eosin.

Descriptions are limited to brief diagnoses,

with important distinguishing characteristics,

except for new species, or where an original

description can be substantially amended.

The term chaetae is used in preference to setae,

following a determination at the International

A. M. PINDER AND R. O. BRINKHURST

Symposium on Aquatic Oligochaete Biology,

Presque Isle, Maine, USA, 1997. Roman num¬

erals refer to segment numbers and two arabic

numerals separated by a slash refer to the septum

(or intersegmental furrow) separating two adja¬

cent segments. Body lengths are for preserved

specimens and are indicative of the length of

mature specimens only.

Where details of the holotype are not given, as

is the case for some of the earliest described cos¬

mopolitan species, this information is unknown

and not listed in Reynolds and Cook (1976, 1981,

1989, 1993).

Institutions and collections are abbreviated as

follows: AMS, Australian Museum, Sydney,

NSW; AMP, Adrian Pinder collection; AWT,

Australian Water Technologies, Sydney, NSW;

BGMJ, Barrie Jamieson collection, Brisbane,

Old; CALM, Wildlife Research Centre, Depart¬

ment of Conservation and Land Management,

Perth, WA; ERISS, Environmental Research

Institute of the Supervising Scientist. Jabim, NT;

IFC, Inland Fisheries Commission, Hobart, Tas..

(all collections now in QVM); MDFRC, Murray

Darling Freshwater Research Centre, Albury,

NSW (J. Hawking); MNHN, Museum National

d'Histoire Naturelle, Paris, France; MON, Ecol¬

ogy and Evolutionary Biology, Monash Univers¬

ity, Melbourne, Vic.. (P.S. Lake); MUR, Aquatic

Ecosystems Research, Environmental Science,

Murdoch University, Perth, WA (J. Davis);

NMV, Museum Victoria, Melbourne, Vic.; NTM,

Museum and Art Gallery of the Northern Terri¬

tory, Darwin, NT; QVM, Queen Victoria

Museum, Launceston, Tas.; ROB, Ralph

Brinkhurst collection; SAM, South Australian

Museum, Adelaide, SA; TAS., Tasmanian

Museum, Hobart, Tas.; UCA, University of

Canberra, Canberra, ACT (R. Norris);

USNM, National Museum of Natural History,

Washington, USA; WAM, Western Australian

Museum, Perth, WA; ZMUH. Zoological

Museum, University of Hamburg, Germany.

Abbreviations used in the figures arc as

follows: a, atrium; c, copulatory chamber; e, ejac¬

ulatory duct; f, female funnel; g, gut; gd, gut

diverticulum; gt, glandular tissue; mf, male fun¬

nel; n, nerve cord; o, ovary; p, prostate; pa, papil¬

lae; pc, penial chaetae; pe, peritoneal tissue;

s, spermatheca; sc, spermathecal chaetae; sd.

spermathecal duct; t, testis; v, vas deferens.

Where possible, localities are given with lati¬

tude and longitude, Australian map grid (AMG)

1:100 000 coordinates (four digit map number

followed by three eastings and three northings) or

Tasmap I: 500 000 coordinates (three eastings

and three northings). Australian states and terri¬

tories are abbreviated as follows: Australian

Capital Territory (ACT), New South Wales

(NSW), Northern Territory (NT), South Aus¬

tralia (SA), Queensland (Qld), Victoria (Vic.),

Tasmania (Tas.) and Western Australia (WA).

Tubificidae

Diagnosis. Genitalia bilaterally paired. One pair

of testes and 1 pair of ovaries, usually in X and XI

respectively. Spennathecae mostly in the testicu¬

lar segment. Male funnels on posterior septum of

testicular segment leading to vasa deferentia, atria

and male pores in the ovarian segment. Female

funnels leading to pores on posterior furrow of

ovarian segment. Chaetae from 1 to many per

bundle, hair chaetae usually restricted to dorsal

bundles or absent. Reproduction normally sexual.

Tubificinae

Diagnosis. Spermathecal pores in the middle of

the testes segment in most species. In all but a few

genera, sperm organised as spermatozeugmata of

a specific type, with non-fertile sperm encapsulat¬

ing fertile sperm. Atria each with a single stalked

prostate gland (prostate rarely absent). Pendant

penes usually present, often with cuticle of penis

sac thickened to form a penis sheath. Sperma¬

thecal chaetae, when present, usually single,

enlarged, grooved distally and with large ehaetal

glands. Penial chaetae normally absent, but when

present then usually of similar form to the sper¬

mathecal chaetae. Coelomocytes not large and

abundant.

Remarks. The spermatozeugmata found in tubi-

Hcines arc of a distinct form, with fertile sperm

cells enclosed by a cortex of bound infertile

sperm cells (Braidotti and Ferraguti, 1982;

Ferraguti et al„ 1994; Jamieson, 1992), distinct

from the looser sperm agglomerations found in

the rhyacodrilinae.

Tubifex Lamarck

Type species. Lumbricus tubifex Muller, 1774.

Diagnosis. Hair chaetae usually present dorsally,

accompanied by pectinate chaetae, but poly¬

morphic forms occur. Vasa deferentia long, often

of two widths. Atria comma-shaped, with vasa

deferentia opening close to the apex anteriorly,

each with a large stalked prostate gland. Pendant

penes present, each with a thin tub-shaped sheath.

Genital chaetae not modified. Spermatozeugmata

present.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

Tubifex tubifex (Muller)

Lumbricus tubifex Muller, 1774: 27 (in part).

Tubifex rivuiorum Lamark, 1816: 225.

Tubifex tubifex. Michaelsen, 1900: 48.—

Brinkhurst, 1971a: 453, figs 8.1A-D, 8.3G-J.—

Brinkhurst, 1971b: 10, fig. 2A.—'Timms, 1973: 11.—

Timms, 1977: 170.—Timms, 1978: 435.—Timms,

1981: 188.—Timms, 1982: 126.—Marshall and

Winterboum, 1979: 205.

Material examined. NSW: Wingecarribee Creek,

Bcrrima, 34°29'S 150°20'E, 7 Nov 1991, 4 specimens

(AWT). Tas.: stream 4.9 km above Barrow Creek, 26

Nov 1978, 1 on slide (ROB); creek E of The Needles,

13 Nov 1978, 2 on slides (ROB): Arthur R., 31 Oct

1978, 1 on slide (ROB). Vic.: Beechworth Trout Hatch¬

ery, May 1994, 1 on slide (AMP) and Sep 1994, 2 on

slides (NMV F77644-F77645); Thomson R. at Thomp¬

son Valley Rd, 24 Nov 1976, 2 on slides and 3 in

alcohol (NMV F77646-F77648); Phytotelmata in fig

tree. South Yarra, Melbourne, several on slide (MON).

WA: Soak NW of Corrigin, 32°10'38”S 117°49'08"E,

21 Oct 1997, several on slides and in alcohol (CALM);

Paperbark Swamp, E of Corrigin, 32°24'58 'S

118°05'52' E, 8 Oct 1997, 1 on slide (CALM); Farm

dam near Corrigin, 32°19'44 "S 118°02'24"E, 7 Oct

1997, 2 on slides (CALM).

Other Australian records. Qld: Downfall Creek

(Brinkhurst, 1971b). SA: A creek near Adelaide, no

date, I on slide (AMP). Vic.: Lakes Bong Bong,

Bridgewater, Purrumbete and Elusive (Timms, 1973,

1981, 1977). WA: Lake Monger, Perth (Brinkhurst,

1971b). Tas.: Lake St Clair (Timms, 1978).

Distribution and habitat. Cosmopolitan, includ¬

ing New Zealand (Brinkhurst, 1971b; Marshall

and Winterboum, 1979; Timms, 1982). Although

widespread in Australia, T. tubifex is not com¬

monly encountered. In the northern hemisphere

T. tubifex is commonly found in marginal sites,

such as those with heavy organic pollution or

intense sedimentation or in highly oligotrophic

waters (Brinkhurst, 1996), In sites with heavy

organic pollution this species is usually associated

with Limnodrilus hoffineisteri Claparede, 1862

and these are the dominant oligochaetes or even

the dominant benthic invertebrates (Brinkhurst,

1996). Studies of gross organic pollution are less

common in Australia (as are ecological studies in

which oligochaetes are speciated) and the Aus¬

tralian records of T. tubifex span a wide variety of

habitats. This species can tolerate salinities up to

10 ppt (Chapman et al., 1982) and mature worms

can encyst to survive short periods of drought

(Kasterand Bushnell, 1981).

Diagnosis. Length 20-200 mm. Anterior dorsal

bundles with 1-6 hair ehaetae and 3-5 pectinate

chaetae with irregular intermediate teeth. Hair

ehaetae and pectinate chaetae reduced posteriorly.

Ventral chaetae bifid, 6-10 per bundle anteriorly,

with the upper teeth thinner than but not much

longer than the lower, posteriorly with the upper

teeth relatively shorter and fewer chaetae. Poly¬

morphic forms with hairs short and bifid chaetae

dorsally, or bifid chaetae both dorsally and

ventrally, may be found, especially where con¬

ductivity is high. Vasa deferentia elongate,

divided into thin and thick sections, entering the

comma-shaped atria on the concave side, near

the where the prostate glands join. Penis sheaths

tub-shaped but often indistinct.

Limnodrilus Claparede

Limnodrilus Claparede, 1862: 221, pi I figs 1-3, pi.

Ill fig. 12, pi. IV fig. 6.—Michaelsen, 1900: 42.—

Stephenson, 1930: 747.—Brinkhurst, 1971a: 463.

Type species. Limnodrilus hoffineisteri

Claparede, 1862.

Diagnosis. Dorsal and ventral chaetae bifid, hairs

absent. Vasa deferentia long, entering short bean¬

shaped atria apically. Prostates large, entering

atria by short stalks. Atria connect, via long ejac¬

ulatory ducts, to pendant penes lying free within

thick penis sheaths (formed by the thickened cuti¬

cle of the penis sac wall) of varying lengths, some

longer than the body width. No modified genital

chaetae. Spermatozeugmata present.

Remarks. Mature specimens of the three species

present in Australia are readily separable by

examining the penial sheaths. These are usually

visible through the body wall of cleared and slide

mounted specimens.

Limnodrilus hoffineisteri Claparede

Limnodrilus hoffineisteri Claparede. 1862: 248, pi. 1,

figs 1-3, pi. Ill, fig. 12, pi. IV, fig. 6.—Brinkhurst,

1971a: 464, figs 8.3M, O, 8.4C, H, I, 8.5E.-

Brinkhurst, 1971b: 112, fig. 2B.—Brinkhurst and

Fulton, 1979: 1.—Fulton, 1983a: 778.—Fulton. 1983b:

792. -Timms, 1978: 428.—Timms, 1982: 126.—

Marshall and Winterboum, 1979: 205.

Clitellio (Limnodrilus) hoffineisteri. —Valliant, 1890:

424.

Tubifex hoffineisteri. —Ditlevsen, 1904: 422.

Material examined. ACT: Murrumbidgee R.,

1987-1988, returned to R. Norris (UCA). NSW:

Georges R. at Liverpool Weir, 1995, 1 on slide (AMS

W7876); Meggaritys Creek at Warradale Rd, 33“54'S

150°36’E, 9 Feb 1993, I on slide (AWT); Rushy Bill-

abong, 36°02’S 146“42'F, 5 Sep 1984, 1 on slide

(MDFRC); Ryans Billabong, 36°06’S I46"58’E, 30 Jun

1981, I on slide and I in alcohol (MDFRC); Orphan

School Creek, Kings Park, 3 May 1996, 1 on slide

(AWT). Qld: Bulimba Creek, Brisbane, 4.8 km from

mouth, 4 on slides (AMS W18081). SA: Valley Lake,

A. M. PINDER AND R. O. BRINKHURST

Mt Gambier, 37°50'S 140°46'E, 26 Feb 1978, 1 on slide

(AMS W18084). Vic.: La Trobe R. at Moe—Willow

Grove Bridge Rd and 100 m downstream of Moe Drain,

AMG 8121-346773 and 8121-359769 respectively, Jul

1980, 2 on slides (NMV F77626-7); Mitta Mitta R., 2

km downstream of Dartmouth Dam, 31 Jan 1975 and

Nov 1975, 3 on slides (NMV F77628, F77629, F77631)

and 4 km N of Eskdale, 1 Feb 1975, 1 on slide (NMV

F776230). WA: Munday Swamp, Perth, 22 Nov 1989,

1 on slide (WAM 41-95); Fire dam on Seven Day Rd,

Warren R. catchment. 34°19'S 1 !5"58'E, 4 in alcohol,

1 on slide (WAM 39-95 and 40-95); stream across

Bussell Highway, 34°08'32"S 115°06'41 "E, 15 Sep

1996, 1 on slide (AMP).

Ollier Australian records. Tas.: Arthurs Lake and Lake

Sorrel (Fulton, 1983ab); Great Lake (Brinkhurst and

Fulton, 1979); Lakes St Clair, Dove, Sorell, Crescent,

Leake and Toombs (Timms, 1978).

Distribution and habitat. Cosmopolitan, includ¬

ing New Zealand (Brinkhurst, 1971b; Titnms,

1982; Marshall and Winterbourn, 1979). This is

the most commonly encountered Limnodrilus and

is found in many habitat types, reaching very high

abundance in organically enriched sites, often

with T. tubifex (Brinkhurst, 1965b). It can tolerate

salinities up to 10 ppt (Chapman et al., 1982).

Diagnosis. Length 20-35 mm. Anterior bundles

with 3-10 chaetae with upper teeth varying in

length from slightly shorter than, to longer than,

the lower and usually thinner than the lower.

Fewer chaetae posteriorly. Penis sheaths up to 20

times longer than broad, ental ends slightly flared,

ectal ends either with elaborate hood opening at

right-angles to shaft or sometimes a plate with

scalloped edges.

Limnodrilus udekemianus Claparede

Limnodrilus udekemianus Claparede, 1862: 243, pi.

1, figs 4-5, pi. Ill, figs 13, 13a, pi. VII, fig. 1 —

Brinkhurst, 1971a: 467, figs 8.4A-B.—Brinkhurst,

1971b: 113, fig. 2E.—Timms, 1978: 428.—Marshall

and Winterbourn, 1979: 205.

Clitellio (Limnodrilus) udekemianus .—Vaillant

1890: 425.

Tubifex udekemianus. —Ditlevsen. 1904: 422.

Material examined. NSW: Dairy drains, Taree, I Apr

1987 (AMS). Qld: Bulimba Creek, Brisbane, 4.8 km

from mouth, 2 on slides (immature, but with character¬

istic chaetae) (AMS W18082). SA: Valley Lake, Mt

Gambier, 37°50'S 140°46’E, 26 Feb 1978, 1 (immature,

but with characteristic chaetae) on slide (AMS

W18083). WA: Katanning Creek at Katanning sale

yards, 33°42'S 117°34'E, 16 Sep 1997 I on slide

(CALM).

Other Australian records. Qld: Moggill Creek, Bris¬

bane; Downfall Creek, Chermside. Tas.: Lake Pedder

(all previous from Brinkhurst, 1971b), Lake St Clair

and Dove Lake (Timms, 1978). WA: Lake Monger,

Perth, uncertain identification (Brinkhurst, 1971b).

Distribution and habitat. Cosmopolitan, includ¬

ing New Zealand (Brinkhurst, 1971b; Mar¬

shall and Winterbourn, 1979). Often, though

not exclusively, found in organically polluted

situations.

Diagnosis. Length 20-90 mm. Anterior chaetal

bundles with 3-8 bifid chaetae with the upper

teeth much thicker and longer than the lower,

fewer chaetae posteriorly with teeth more nearly

equal. Penis sheath usually only up to 4 times as

long as wide with a simple plate-like hood usually

reflected back over the shaft.

Limnodrilus claparedeianus Ratzel

Limnodrilus claparedeianus Ratzel, 1868: 590, pi.

XLII, fig. 24.—Michaelsen, 1900: 45.—Brinkhurst,

1971a: 468, figs 8.3Q, R.—Brinkhurst, 1971b: 113,

fig. 2F.

Tubifex claparedeianus. —Ditlevsen, 1904: 422.

Material examined. Vic.: Lake Thirlemere, 8 Sep 1981,

2 on 3 slides (NMV F81432).

Distribution and habitat. Cosmopolitan

(Brinkhurst, 1971b), occurring in a wide range

of habitat types.

Diagnosis. Length 30-60 mm. Anterior bundles

with 4-9 bifid chaetae, some with the upper teeth

much longer than and as thick or somewhat

thicker than the lower, fewer chaetae posteriorly.

Penis sheaths up to 50 or even 80 times longer

than broad with small triangular plates set at an

angle to the shaft dislally.

Remarks. This is the first and only record of this

species from Australia. The shape of the penis

sheath is normally diagnostic except in North

America, where variants between this species and

Limnodrilus cervix Brinkhurst, 1963 can be

observed.

Potamothrix Vejdovsky and Mrazek

Potamothrix Vejdovsky and Mrazek, 1902: 24.—

Brinkhurst, 1971a: 486.—Holmquist, 1985: 311.—

Finogenova and Poddubnaja, 1990: 55.

Euilyodrilus Brinkhurst, 1962: 329.

Type species. Potamothrix moldaviensis

Vejdovsky and Mrazek, 1902.

Diagnosis. Hair chaetae present in dorsal bundles

of most species. Vasa deferentia short, joining

elongate tubular atria apically, prostate glands

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

rudimentary or absent. Pendant penes small, with¬

out cuticular sheaths. Large grooved genital

chactae associated with spermathecal pores.

Spermatozeugmata present.

Remarks. While the genus is largely holarctic, and

mostly palearctic, some of the many Ponto-

Caspian species appear to have been introduced to

North America, Africa and South America (Peru)

and others are spreading through Europe (G. Mil-

brink, pers. comm.). The single species occurring

in Australia, Polamothrix bavaricus (Oschmann,

1913), is a cosmopolitan species, although it may

have been introduced to some parts of its current

range, e.g., South America (Brinkhurst and

Marchese, 1989).

Polamothrix bavaricus (Oschmann)

Tubifex (llyodrilus) bavaricus Oschmann, 1913: 559,

figs 1-5.

llyodrilus bavaricus. —Stephenson, 1930: 746.

Euilyodrilus bavaricus. —Brinkhurst, 1962: 321.

Polamothrix bavaricus. -Brinkhurst, 1971 a: 492,

figs 8.12H.I.—Brinkhurst, 1971b: 113, fig. 2G.—

Timms, 1981: 188.—Marshall and Winterbourn, 1979:

205.

Holotype. Type locality unknown (ZMUH 7704).

Materia! examined. NSW: Gowrie Creek, 54 km down¬

stream of sewage treatment plant, 27°20'S 151“27'E,

28 Sep 1995, few in alcohol (AWT). Tas,: Big Water-

house Lake, 40°53'20"S 147’36'50'E, 1 Mar 1997, 2

on slide (AMP). Vic.: Curdies R„ 38°20S 143”08 E, 7

and 29 Feb 1992, 2 on slides (NMV F77641-2). WA:

Malaga Wetlands, Perth, 2 Nov 1989, 2 on slides

(WAM 76-95, 77-95); Herdsman Lake, Perth, 2 and 13

Nov 1989, 13 on slides, 8 in alcohol (slides WAM

55-95, 67-95, alcohol 54-95. 78-95): Lakes Novver-

gup, 3I°38’I6"S 115°44'42"E, Yonderup, 31°34'S

1I5 0 41'E, and Carabooda, 3P’37'S 1I5“43'E. 8 on

slides (WAM 68-95 to 75-95); Coogee Springs, Perth,

9-14 Nov 1989, several on slides (MUR); Shark Lake,

N of Esperance. SS^'S 12I"51'E, 5 Sep 1996, many

specimens in alcohol (AMP); Congelin Dam,

32°49'07”S 116°53'05"E, 26 Oct 1997, 3 on slides

(CALM).

Other Australian records. Vic.: Lake Purrumbete,

Camperdown (AMS W4275) (Timms, 1981). WA:

Corio Pool, Rottnest Island (Brinkhurst, 1971b).

Distribution and habitat. Eurasia, North, Central

and South America, Australia and New Zealand

(Brinkhurst, 1971a; Brinkhurst and Marchese,

1987; Coates and Stacey, 1994; Marshall and

Winterbourn, 1979). Polamothrix bavaricus is

quite widespread in southern Australia, occurring

in both lcntic and lotic sites and often very

abundant when present.

Diagnosis. Length 15-35 mm. Dorsal anterior

bundles with 1-5 hair ehaetae and 2-5 pectinate

chaetae. Ventral ehaetae with teeth equally long

but the upper thinner than the lower. Sperma¬

thecal chaetae relatively stout, about equal in

length to the somatic ventral chaetae, with trian¬

gular blade-like ectal half. Vasa deferentia short

joining tubular atria apically. Prostate glands

absent. Penis sheaths absent.

Antipodrilus Brinkhurst

Antipodrilus Brinkhurst, 1971a: 523.—Brinkhurst

and Wetzel, 1984: 50.

Type species. Tubifex davidis Benham, 1907.

Diagnosis. Dorsal chaetal bundles with or without

hairs. Vasa deferentia long and narrow, joining

atria apically or subapically. Atria globular, nar¬

rowing to form ejaculatory ducts that terminate in

pendant penes (or protrusible pseudopenes?).

Apparent penis sheaths (formed by thickened

cuticular lining of the ectal part of the male duct)

present in some species. Single stalked prostate

gland attached apically each atria. Spermathecae

with elongate spermatozeugmata. Spermathecal

chaetae thin with grooved elongate tips.

Remarks. Antipodrilus is known only from Aus¬

tralia (all five species) and New Zealand (one

species, A. davidis). In a cladistic analysis of the

subfamily by Brinkhurst (1991), Antipodrilus

appeared to group with two genera that are widely

distributed in the northern hemisphere: Psam-

moryctides Hrabe, 1964 and Haber Holmquist,

1978. These northern genera have protrusible

pseudopenes, with a crumpled cuticular lining to

the inverted ectal part of the male duct, which

then forms a penis sheath when the penis is

everted. Antipodrilus magelensis Brinkhurst,

1984 and possibly Antipodrilus timmsi Brink¬

hurst, 1971, appear to have similar crumpled

sheaths within the ectal part of the male duct and

so the genus was coded as having protrusible

pseudopenes in Brinkhurst (1991). Unfortunately

further Antipodrilus specimens were not available

for this study. Antipodrilus davidis (Benham,

1907) does appear to have pendant penes (though

these may be protruded pseudopenes) and the

nature of the penes is not clear for the other two

species (Antipodrilus multiseta Brinkhurst and

Fulton, 1979 and Antipodrilus plectilus Brink¬

hurst and Fulton, 1979). Further anatomical study

is required to confirm the presence of pseudo¬

penes and to generally improve descriptions of

Antipodrilus species.

A. M. PINDER AND R. O. BRINKHURST

Antipodrilus davidis (Benham)

Tubifex davidis Benham, 1907: 252, figs 1-6.—

Hrabe. 1962: 307.—Hrabe, 1966: 74.

Etdlyodrilus heuscheri (Bretscher).—Brinkhurst,

1963: 49 (in pari).

Antipodrilus davidis. —Brinkhurst, 1971a: 523.—

Brinkhurst, 1971b: 112, fig. 2b.—Brinkhurst. 1982:

1.—Timms, 1979: 121—Timms, 1982: 126.—

Marshall and Winterboum. 1979: 205.

Holotype. Blue Lake, Mt Kosciusko, NSW. 1906.

sectioned on 4 slides (AMS G5610 11).

Material examined. NSW: Holotype, as above. SA: The

Bubbler Spring, near Lake Eyre. 3 Dec 1974, 2 on

slides, 1 in alcohol (SAM E2823-E2825); Coopers

Creek, southwest branch. 27 Mar 1987, 1 on slide

(SAM E2817); Freeling Springs, 1987 (AMS

W198716); Davenport Spring Bore, near Hermit Hill,

Lake Eyre South. 21 Sep 1989. 5 on slides (SAM

E2818-E2821). W'A: Spectacles Lake (North Eye),

Brownmans Swamp, Lake Mariginiup and Thomson

Lake, Perth. 30 Oct and 3-9 Nov 1989. 11 on slides. 8

in alcohol (WAM 42-95 to 47-95, 49-95 to 53-95 and

81-95).

Other records. NSW: Mt Kosciusko lakes (Timms,

1979). Following records all from Brinkhurst (1971b).

SA: Lakes Edward and Leake, Glencoe. Vic.: Lakes

Surprise and Mumblin, western Victoria. Tas.:

Cambridge. 30 Aug 1966 (TAS. K221). Also recorded

from New Zealand (Brinkhurst, 1971b; Marshall and

Winterboum. 1979; Timms. 1982).

Distribution and habitat. Southern Australia and

New Zealand (Brinkhurst, 1971b; Marshall and

Winterboum, 1979; Timms, 1982). Favours lentic

habitats.

Diagnosis. Length 25-40 mm. Anterior dorsal

bundles with 2-5 non-hispid hair chaetae and 2-4

pectinate chaetae with teeth approximately equal.

Ventral chaetal bundles with 3-5 bifid chaetae

with the upper teeth longer and thinner than the

lower anteriorly and teeth equal or the upper

shorter posteriorly. Spermathecal chaetae single,

grooved or with short lower tooth (probably when

partially developed), about equal in length to

somatic ventral chaetae, rarely absent. Vasa def-

erentia long, atria small, narrowing abruptly to

fonn long ejaculatory ducts leading to (pendant?)

penes.

Antipodrilus timmsi Brinkhurst

Antipodrilus timmsi Brinkhurst, 1971b: 112, fig.

12c.—Brinkhurst and Wetzel, 1984: 50.—Timms,

1981: 187.—Timms, 1983: 168.

Holotype. Lake Bullenmcri, Vic., 13 Aug 1969, 5 on

slides (AMS W4276-W4278, typus amissus).

Material examined. NSW: Lake Jillamatong, near

Cooma, 3 Apr 1986, many in alcohol (NMV F77614).

Vic.: Type locality and date, 5 on slides (ROB); Lake

Colangulac, 2 on slides (NMV 77612—F77613).

Ollier Australian records. Vic.: several lakes in western

Victoria (Timms, 1981, 1983).

Distribution and habitat. Lakes in southeastern

mainland Australia. Has been collected in sites

with up to 25 ppt salinity, in both sandy and

muddy sediments and at depths of up to 65 m

(Timms, 1981, 1983).

Diagnosis. Length at least 14.5 mm (length of

largest fragment). Anterior dorsal bundles with

2-5 serrate hairs and 3-5 pectinate chaetae with

the upper teeth slightly longer than the lower,

fewer posteriorly with upper teeth becoming

longer and pectinations less apparent or absent.

Ventral bundles with 3-5 chaetae anteriorly with

the upper teeth markedly longer and somewhat

thinner than the lower, fewer chaetae posteriorly

with upper teeth less markedly longer than the

lower. Spermathecal chaetae single, grooved and

narrowly blade-shaped distally, slightly longer

than somatic ventral chaetae. Vasa deferentia

long, atria small and almost spherical, narrowing

abruptly to form ejaculatory ducts. Penes present,

possibly with thickened inner cuticle.

Remarks. The crumpled cuticle within the penes

is indicative of a protrusible penis, but the struc¬

ture of this requires closer examination from fresh

specimens.

Antipodrilus magelensis Brinkhurst

Antipodrilus magelensis Brinkhurst, 1984: 142, figs

1 - 2 .

Holotype. Mudginberri Billabong, Magela Creek, NT 4

May 1981, 1 on slide (USNM 80692).

Material examined. NT: Holotype, as above, and

paratypes from type locality as well as Georgetown and

Bowerbird Billalxmgs, Magela Creek, 11 on slides, 19

in alcohol (USNM 80693-80697, 80700) and several

on slides (ROB); Nankeen Billabong, Magela Creek, 1

immature on slide (ROB); South Alligator R„ Oct 1988

and Dec 1989, 1 on slide (NMV F7761I) and 2 on

slides (ERISS). SA: Charles Angus Bore, 15 km N of

Alberri Creek railway siding, Lake Eyre South. 2 Dec

1974, 1 on slide (SAM E2815); Coopers Creek, north¬

west branch, 27 Mar 1987. I on slide (SAM E2816).

Distribution and habitat. Northern Territory

(coastal riverine sites) and South Australia (few

arid inland sites). The new South Australian spec¬

imens arc the first to be recorded outside of the

Northern Territory.

Diagnosis. Length up to 9 mm. Anterior bundles

with 4-6 bifid chaetae with teeth equal in length

and breadth, diminishing in number posteriad to 2

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

per bundle with upper teeth becoming shorter and

thinner than the lower. Spermathecal chaetae

single with grooved tips, about equal in size to

somatic ventral chaetae. Vasa deferentia moder¬

ately long, joining atria subapically. Short atria

terminate in coiled ejaculatory ducts, which lead

to (protrusible?) penes with crumpled cuticular

lining forming apparent penis sheaths.

Antipodrilus multiseta Brinkhurst and Fulton

Antipodrilus multiseta Brinkhurst and Fulton, 1979:

2, figs 6-8.—Brinkhurst and Wetzel, 1984: 50.

Fulton, 1983a: 778.—Fulton. 1983b: 792.

Holotype. Great Lake, Tas., 1975 (QVM:14:316).

Material examined. Tas.: Holotype and paratypes from

type locality and date, four on slides

(QVM:14:316-319); Lake Soreli. 3 on slides, uncertain

identification (ROB).

Ollier Australian Records. Tas.: Great Lake, Arthurs

Lake and Lake Soreli (Fulton, 1983ab).

Distribution and habitat. Known only from large

lakes of Tasmania’s central plateau. Collected at

depths of up to 17.5 m and in both silt/sand and

clay sediments (Fulton, 1983b).

Diagnosis. Length up to 50 mm. Chaetae 12-15

per bundle in II, gradually diminishing in number

to 3 or 4 posteriorly. Anterior chaetae with blunt

teeth of equal thickness, the upper slightly longer,

posterior chaetae with upper teeth shorter than the

lower. Spermathecal chaetae single, thin with

grooved-tips, about equal in size to somatic

ventral chaetae. Vasa deferentia moderately long,

atria and prostate glands small, ejaculatory ducts

long and thin, leading to small penes. Posterior

end of body tapering to a coiled tail. Living with¬

in tubes of fine sand grains (at least in the type

locality).

Antipodrilus plectilus Brinkhurst and Fulton

Antipodrilus plectilus Brinkhurst and Fulton,

1979: 2, figs 1-5. — Brinkhurst and Wetzel, 1984:

50.—Fulton, 1983a: 778.—Fulton. 1983b: 792.

Holotype. Great Lake, Tas., 1975 (QVM: 14:309).

Material examined. Tas.: Holotype and paratypes from

type locality and date (QVM: 14:309-315); Lake Soreli,

2 Feb 1981, 7 on slides (ROB).

Other Australian records. Tas.: Great Lake and Arthurs

Lake (Fulton, 1983a, b).

Distribution and habitat. Known only from large

lakes of Tasmania’s central plateau. Collected at

depths of up to 17.5 m and in both silt/sand and

clay sediments (Fulton, 1983b).

Diagnosis. Length at least 20 mm. Thin, red

worms that may be found in clusters of up to 50

specimens. Dorsal and ventral bundles with 7-9

chaetae anteriorly, reduced in number posteriorly

to 3 or fewer per bundle. Chaetae of II with upper

teeth shorter and thinner than the lower, chaetae

of other preclitellar bundles with teeth equally

long and with teeth equal in width or the upper

teeth thinner than the lower. Posterior chaetae

with upper teeth shorter and thinner than the

lower. Spermathecal chaetae single, thin and

groove tipped, twice as long as somatic ventral

chaetae. Vasa deferentia moderately long, atria

elongate, distended where vasa deferentia and

prostates attached. Ejaculatory ducts moderately

long.

Remarks. The type specimens have all chaetae

bifid, but those from Lake Soreli have pectinate

chaetae posteriorly. Further investigation may be

required to substantiate the unity of this taxon,

although chaetal polymorphism is known for

other oligochaetes (Chapman and Brinkhurst,

1986, 1987).

Aulodrilits Bretscher

Aulodrilus Bretscher, 1899: 388.—Michaelsen,

1900: 55.—Stephenson, 1930: 752.

Type species. Aulodrilus limnobius Bretscher,

1899.

Diagnosis. Dorsal bundles with or without hair

chaetae. Often with gonads replicated and/or gen¬

ital apparatus shifted forwards due to asexual

reproduction. Vasa deferentia short, atria globu¬

lar, bean-shaped or cylindrical with solid prostate

glands. Penes pendant without cuticular sheaths.

Penial chaetae (but not spermathecal chaetae)

may be modified. Sperm stored as loose bundles,

spermatozeugmata absent.

Remarks. Asexual reproduction is common in

Aulodrilus (Brinkhurst, 1984) and most Aus¬

tralian records are of immature worms. However,

the three species found in Australia have distinc¬

tive chaetae that permit identification of immature

specimens. Finogenova and Arkhipova (1994)

provide scanning electron micrographs of the

distinctive chaetae of the following species.

Aulodrilus limnobius Bretscher

Aulodrilus limnobius Bretscher, 1899: 388, figs

8.23G-H.—Brinkhurst, 1971a: 524, figs 8.24G, H. -

Finogenova and Arkhipova, 1994: 8, figs 1-2, 4-5, 19.

Material examined. NSW: Murray R. at Heyward’s

Gauge, 36°06'S 147 o 01'E, 6 Jan 1988, 1 on slide

(MDFRC). Vic.: Wentworth R., 17.7 km upstream of

A. M. PINDER AND R. O. BRINKHURST

Tabberabbera, 37“31'S 147°23'E, 2 Oct 1975, 1 in

alcohol (NMV F77618J; Mitta Mitta R.. 2 km down¬

stream of Dartmouth Dam wall, 5 Nov 1976, 1 in alco¬

hol (NMV F776I5); Branjce Creek, Goulbum Valley,

36°42’S 148°20'E, 23 Apr 1992, 1 on slide, 3 in alcohol

(NMV F77616, F77617).

Distribution and habitat. Recorded from Africa,

Europe, Asia and North and South America

(Brinkhurst, 1971a) and now southeastern

Australia.

Diagnosis. Length 12-15 mm. Anterior dorsal

and ventral chaetae similar with up to 10 bifid

chaetae with upper teeth shorter and thinner than

lower, the chaetae of first bundles often shorter

and thicker than the rest. Chaetae of mid and pos¬

terior segments with lateral wings which appear

as narrow oar-shaped blades when viewed

frontally, with the upper teeth visible as a small

projection beyond the blade. The wings can be

discerned as a ridge on the lateral aspect of the

chaetae. Vasa defcrentia long, entering long

cylindrical atria apically. Pendant penes large.

Genitalia may be shifted forwards.

Remarks. These are the first Australian records of

this otherwise widespread species.

Aulodrilus phiriseta (Piguet)

Naidium phiriseta Piguet, 1906: 218.

Aulodrilus phiriseta .—Piguet and Bretschcr, 1913:

118, figs 1-2.—Brinkhurst, 1971a: 524, figs

8.23J-N.—Brinkhurst, 1971b: 114, fig. 21.—

Brinkhurst, 1984: 147.—Timms, 1982: 126.—Marshall

and Winterboum, 1979: 205. —Finogenova and

Arkhipova, 1994: 1 I. figs 11-13, 20-22.

Aulodrilus japonicus Yamaguchi, 1953: 298, fig. 12,

pi. VII. figs 5-7.— Finogenova and Arkhipova. 1994: 8,

figs 3, 6.9, 10.

Material examined. No new specimens examined.

Other Australian records. NT: Georgetown and Mudg-

inberri Billabongs, Magela Creek, Kakadu National

Park (Brinkhurst, 1984). Qld: Yule Creek, Yanda Yarra

(Brinkhurst, 1971b).

Distribution and habitat. Cosmopolitan

(Brinkhurst, 1971b) but in Australia only known

from the north so far.

Diagnosis. Length 10-25 mm. Anterior dorsal

bundles with up to 8 short hair chaetae and up to

10 bifid chaetae with the upper tooth shorter and

thinner than the lower and frequently replicated.

Anterior ventral chaetae up to 16 per bundle with

very thin, short upper teeth. Vasa deferentia long,

entering ovoid atria apically. Atria with large

lobate prostate gland, terminating in pendant

penes. Genitalia may be shifted forwards.

Remarks. All Australian specimens observed so

far have had replicate upper teeth on the dorsal

chaetae. Specimens with replicate teeth were

originally described as a separate species, Aulo¬

drilus japonicus Yamaguchi, 1953, but this

species was synonymised with A. pluriseta by

Brinkhurst (1971a). Some Russian specimens,

identified as A. japonicus by Finogenova and

Arkhipova (1994), had replicate upper teeth, male

pores in X and there were other minor differences

in the form of the male ducts. Replicate teeth on

these specimens were shown to be lateral, occur¬

ring either side of the main teeth. These may be a

separate species, but forward shifting of the

reproductive organs is commonly associated with

asexual reproduction and other A. phiriseta

(including other specimens described by Finogen¬

ova and Arkhipova (1994) have testes as far ante¬

rior as V (Brinkhurst, 1971a). Furthermore, the

replicate teeth are essentially a form ol pectina¬

tion which has been shown to vary in other tubifi-

cids, including as a response to environmental

conditions (Chapman and Brinkhurst, 1986,

1987).

Aulodrilus pigueti Kowalewski

Aulodriluspigueti Kowalewski, 1914: 625, fig. 12.—

Brinkhurst, 1971a: 526, fig. 8.231.— Brinkhurst, 1971b:

114, fig. 2J.—Brinkhurst, 1984: 147. —Finogenova

and Arkhipova, 1994: 13, figs 14—17.

Material examined. NSW: Ryans Billabong (station

61), 36°06 S I47"58E. 30 Aug 1978, 1 on slide

(MDFRC): Manning R. at Wingham Bridge, 3F’52'S

152°23'E, 29 May 1996, 1 immature (AWT). NT: Main

stream in Compartment L of CSIRO Kapalga Research

Station, 20 Aug 1992, 10 in alcohol (NMV F77619).

Vic.: La Trobe R. below Moe Drain, AMG 8121

359769. 3 Jul 1979 and Jul 1980, 7 immature in alcohol

(NMV F77621-F77624): Steavenson R„ 19 Feb 1991,

1 immature in alcohol (NMV F77625); Branjee Creek,

Goulbum Valley, 23 Apr 1992, I immature in alcohol

(NMV F77620). WA: Geegully Creek, Kimberley

region, 18°13’34''S I23"44'36'E, 5 May 1995, few

immature in alcohol (CALM): Meda R. near Lennard

R. Gorge, 17°20'58"S 124°30'27”E, 20 May 1996, 2

mature in alcohol (CALM).

Other Australian records. NT: Georgetown, Mudgin-

.berri and Nankeen Billabongs, Magela Creek, Kakadu

National Park (Brinkhurst, 1984). WA: Pine Hill Rock,

near Balladonia, locality uncertain (Brinkhurst, 1971b).

Qld: Pond near Tully (Brinkhurst, 1971b).

Distribution and habitat. Cosmopolitan

(Brinkhurst, 1971b) and widespread in Australia,

occurring in a variety of habitats.

Diagnosis. Length 2-28 mm. Dorsal anterior bun¬

dles with 2-5 hairs and 4-5 (rarely to 10) other

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

chaetae which are either simple-pointed or bifid

with the upper teeth shorter and thinner than the

lower. Beyond Vll the bifid dorsal chaetae

become oar-shaped when viewed laterally (i.e. in

the same plane as the teeth), often with rudimen¬

tary teeth visible on the end of the oar blade. Dor¬

sal chaetae or just hair chaetae may be absent

from some or all of II—VII. Ventral bundles with

3-6 bifid chaetae, with upper teeth shorter and

thinner than the lower, not becoming oar-shaped.

Large spoon-shaped penial chaetae beside each

male pore on XI in mature specimens. Vasa def-

erentia short, atria bean-shaped. Pendant penes

open via a median inversion of the body wall.

Remarks. The chaetae of A. limnobius are also

oar-shaped, but more narrowly so than in A.

pigueti and the blade-like wings of the former are

in a plane perpendicular to the orientation of the

teeth rather than in the same plane.

Rhyacodrilinae Hrabe

Diagnosis. Spermathecal pores close to anterior

border of their segment (usually the testes bearing

segment, normally X). Spermathecal ampullae

containing loose or loosely bundled sperm. Atria

with prostate glands diffuse or formed into dis¬

tinct glands, rarely absent. Pendant penes gener¬

ally absent; protrusible or eversible pseudopenes

often present. Penial chaetae often modified,

often numerous and typically arranged lanwise or

in a row, with the blunt or bifid ectal ends close

together. Spermathecal chaetae usually not modi¬

fied. Coelomocytes usually large and abundant,

rarely absent.

Remarks. Without doubt, the most problematic

Australian freshwater tubificids are the four

species placed in the Telmatodrilinac by

Brinkhurst (1971a) and Brinkhurst and Fulton

(1979) and two similar new species to be

described below. The Telmatodrilinae is diag¬

nosed by the presence of multiple prostate glands,

each attached to the atria by short stalks. Telma-

todrilus multiprostatus Brinkhurst, 1971 and Tel-

matodrilus pectinatus Brinkhurst, 1971 were

placed within the Telmatodrilinae because the

prostate cells are grouped into several glands,

each of which appear to have a discrete connec¬

tion to the atrial lumen. Later, two other Aus¬

tralian species, Telmatodrilus (Alexandrovia)

papillatus Brinkhurst and Fulton, 1979 and Tel¬

matodrilus? (Telmatodrilus?) bifidus Brinkhurst

and Fulton, 1979, were tentatively placed here for

the same reasons, although T. bifidus has only two

prostates per atrium and these are not stalked. Of

the two new species described below, one has

prostate tissue similar to that of T. multiprostatus

and the other has a pair of unstalked prostate

glands, like that of T. bifidus.

Prior to the description of the Australian forms,

the subfamily Telmatodrilinae consisted of three

or four northern hemisphere species in two genera

(or subgenera); Telmatodrilus Eisen, 1879 from

western USA and Canada, and Alexandrovia

Hrabe, 1962 from Alaska and Russia. This sub¬

family is generally considered to be closely

related to the Tubificinae, based on cladistic anal¬

yses using the few characters available (Erseus,

1990b; Brinkhurst, 1994).

The Australian species differ from the holarctic

forms in many respects. Their atria are ovoid to

pear-shaped with apical prostates, rather than

tubular with prostates along their length. In two of

the Australian species the prostates lack stalks,

instead being attached rather broadly to the ante¬

rior and posterior walls of the atria (Figs 9E, 11 A,

B). The Australian species have simple male

pores, usually on the inner aspects of ventrolateral

folds of the body wall, whereas the northern

forms have pendant penes. Grooved spermathecal

chaetae are found in two of the northern species

and most of the Australian species, but all except

one of the Australian forms also have modified

penial chaetae which are absent in the northern

forms, Coelomocytes are present in the two new

species but have not been observed in the other

Australian forms and appear to be absent in the

holarctic species.

Before discussing the subfamilial relationships

of these Australian species we should indicate

that we intend to assign them to two new genera,

described below. Breviatria gen. nov. will include

B. multiprostatus, B. pectinatus, B. papillatus and

B. arvensis sp. nov. and Biprostatus gen. nov. will

include B. bifidus and B. duplex sp. nov. These

names are mentioned here to facilitate the

following discussion.

Erseus (1990b), in a cladistic analysis of the

Naididae and subfamilies of Tubificidae, showed

the two subfamilies Tubificinae and Telmatodrili¬

nae (based on the northern species) as sharing

plesiomorphic character states for modified sper¬

mathecal chaetae, absence of penial chaetae,

unmodified oesophagus, absence of multiple

large granular coelomocytes, lack of atrial cilia-

tion, and absence of posterior prostates. Tubifi¬

cinae and Telmatodrilinae share the apomorphic

states for penes and spermatozeugmata present.

While these subfamilies share the presence of

stalked prostates, the former has one per atrium,

the latter many, and the analysis suggested that

A. M. PINDER AND R. O. BRINKHURST

the stalked prostates of these two groups had

‘evolved independently from the diffuse prostate

condition in the tubificid ancestor’. These sub¬

families are also separated in that analysis by the

apomorphic state of the spermathecal pore posi¬

tion in midsegment in Tubificinae (anterior in the

Telmatodrilinae). Erseus (1990b) found that the

telmatodrilines and tubificines formed a sister

clade to another containing the phallodrilines,

marine and freshwater rhyacodrilines and the

Naididae, which share the presence of numer¬

ous coelomocytes and penial chaetae. The cod¬

ings in Erseus’ analysis depend on the coding of

the outgroup (Phreodrilidae) adopted and are not

necessarily accepted here.

We reran the analysis with the Australian forms

as a separate group (AUS) using PAUP 3.1 and

the original options. AUS was coded as lacking

spermatozeugmata, atrial cilia and penes, but pos¬

sessing coelomocytes, penial chaetae and multi¬

ple prostates (the coding was thus 10100 01101

000). Three equally parsimonious trees of 24

steps (Cl 0.708) were obtained (unpublished

data). In all of these, the Australian species appear

within a clade which also included Phallodrilinae

and marine and freshwater Rhyacodrilinae, sepa¬

rate from a Tubificinae-Telmatodrilinae-Limno-

driloidinae clade. This was predictable because of

coding for the presence of coelomocytes and

penial chaetae, and implies that the multiple

prostates of AUS are a convergent development

to those of the (northern) Telmatodrilinae. Within

this group, AUS was associated with the Phallo¬

drilinae and marine Rhyacodrilinae, rather than

the freshw'ater Rhyacodrilinae and associated

Naididae, because of the lack of hair chaetae and

the apical vasa deferentia.

Brinkhurst’s (1994) cladistic analysis had the

broader aim of investigating the evolutionary

relationships within the whole of the Clitellata

and the characters and choice of states used

reflected this. Again, Telmatodrilinae was coded

based on the northern species, on the assumption

that these were plesiomorphic within the sub¬

family. A clade including Tubificinae, Telmato¬

drilinae and Limnodriloidinae, recognised by pos¬

session of spermathecal chaetae (not unique) and

stalked prostates, loss of penial chaetae (presence

of which was considered plesiomorphic for the

family), and organised bundles of sperm in the

spermathecae, separated from a clade containing

the Rhyacodrilinae and Naididae. As discussed

below, the Phallodrilinae was associated with

the tubificine clade rather than that including

Rhyacodrilinae.

This analysis was also rerun using PAUP 3.1

and the original options and weightings and with

all characters unordered. AUS was coded as lack¬

ing spermatozeugmata but having coelomocytes,

penial chaetae and stalked prostates (the coding

was thus 10110 lllll 50010 01000 012). Twelve

trees of equal length (88, Cl 0.568) were pro¬

duced (unpublished data). In this rerun, AUS

appeared within a clade with the Tubificinae,

northern Telmatodrilinae, Phallodrilinae and

Limnodriloidinae, which formed a sister group to

the clade including Rhyacodrilinae and Naididae.

The appearance of AUS and Phallodrilinae with

the northern Telmatodrilinae, Tubificinae and

Limnodriloidinae, rather than with the Rhyaco¬

drilinae (as in Erseus’ study), appears to result

from the fact that this analysis saw penial chaetae

as plesiomorphic for the whole family and that the

prostates were coded differently by Erseus.

The lack of consistency in the above analyses

requires that we consider some of the character

states in detail.

Prostate. Traditionally, the prostate tissue of

Rhyacodrilinae has been described as diffuse

(e.g., Brinkhurst, 1971a), meaning that the

prostate cells form a continuous layer over the

atrium, with each cell communicating with the

atrial lumen independently. More recently, this

view (which focused on the freshwater forms) has

been modified following the description of

numerous marine fonns with prostate tissue that

is more clumped. Thus, Baker and Brinkhurst

(1981) describe the Rhyacodrilinae as “having

prostate glands forming a diffuse glandular cover¬

ing; the cells communicating with the atrial lumen

independently or in small clusters”. Erseus (1984)

describes the subfamily as having “Prostates dif¬

fuse or irregularly scattered in two or more clus¬

ters of cells broadly attached to the atrial surface,

or absent”. Some rhyacodri lines, including the

marine Ueronidrilus Erseus and Jamieson, 1981

and Heterodrilus Pierantoni, 1902, the freshwater

monotypic Macquaridrilus Jamieson, 1968 and

the Lake Baikal endemic Rhyacodriloides

Chekanovskaya, 1975, have some or all of the

prostate cells formed into two or more glands

(Ijrseus, 1981; Erseus and Jamieson, 1981;

Jamieson, 1968; Martin and Brinkhurst, 1998).

There are therefore precedents for including Bre-

viatria and Biprostatus in Rhyacodrilinae in

respect to the form of the prostates. Although

most of the rhyacodrilines mentioned above have

the prostate clumps broadly attached to the

atrium, rather than narrowly stalked, the same is

true for the two Biprostatus species. Furthermore,

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

some Heteroclrilus (H. inermis (Erseus, 1981) and

H. rapidensis Erseus, 1997) do have prostate

glands that narrow into stalks like Breviatria. The

Phallodrilinae is the only other subfamily to have

more than one prostate gland per atrium and so

should be mentioned here. Like Biprostatus , phal-

lodrilines have a pair of prostates on each atrium,

but these are almost always stalked and typically

placed ental and ectal in position on the atria

(Erseus, 1992), unlike the symmetrically placed

prostate tissue of Biprostatus. A few phallodri-

lines ( Pacifidrilus Erseus, 1992 and Pirodrilus

Erseus, 1992) do have the prostates placed sym¬

metrically on the atria but this appears to be a

consequence of the small size of the atria. There

is a need for more knowledge on the histology

and morphogenesis of prostate tissue in various

tubificids to more adequately use prostate

characters in phylogenetic work, as highlighted

by Gustavsson and Erseus (1997).

Coelomocytes. An abundance of coelomocytes

(a rhyacodriline trait) has been found in both of

the new species described below but their pres¬

ence in the older Australian taxa cannot be estab¬

lished because of the condition of the scarce

material. They have recently been observed in

rhyacodri lines such as TorodrilusC ook, 1970 and

Rhizodrilus Smith, 1900, in which they were pre¬

viously thought to be absent (Erseus, 1994).

Coelomocytes have not been observed in Rhyaco-

driloides, but that genus is nonetheless now asso¬

ciated with the Rhyacodrilinae (Baker and

Brinkhurst, 1981; Martin and Brinkhurst, 1998).

Coelomocytes have not been reported for any of

the northern telmatodrilines (Holmquist, 1974)

and, with few exceptions, are not abundant in the

Phallodrilinae (Erseus, 1992).

Genital chaetae. Groove-tipped genital chaetae

are common in the Tubificinae and Limnodriloid-

inae and some northern Telmatodrilinae. These

are generally few in number (usually I or 2 per

bundle) and lie in enlarged chaetal glands, usually

on the spermathecal segment. A few rhyacodri-

lines, some Rhizodrilus and Rhyacodriloides , do

have similar chaetae (Baker and Brinkhurst,

1981; Erseus, 1990a; Martin and Brinkhurst,

1998), although these are on the male pore seg¬

ment in the latter. The superficially similar

chaetae of the rhyacodriline Torodrilus , appear to

be a separate apomorphic development (Erseus,

1994). More typical in the Rhyacodrilinae and

Phallodrilinae is modification of the penial

chaetae associated with the male pores. These

have bifid or simple (often club-shaped) tips,

arranged in a row or fanned out with the tips close

together, always on the male pore segment.

Chaetae of the male pore segment are not modi¬

fied in the northern Telmatodrilinae and only

rarely modified in the Tubificinae (two Vari-

chaetadrilus Brinkhurst and Kathman, 1983 and

the few species of Krenedrilus Dunmicka, 1983).

Biprostatus and Breviatria are unusual within the

family in possessing both grooved chaetae on the

spermathecal segment (absent in two Breviatria)

and modified penial chaetae on the male pore seg¬

ment (absent in one Breviatria). Krenedrilus

(Tubificinae) and some Rhizodrilus (Rhyaco¬

drilinae) are the only other tubificids to have both

types of genital chaetae. This means the genital

chaetae may be of little help in classifying the

new genera, unless grooved spermathecal chaetae

are seen as plesiomorphic for the family (almost

identical chaetae do occur in the Phreodrilidae)

and modified penial chaetae are seen as an apo¬

morphic development in the ancestral rhyacodri¬

line, as tentatively suggested by Erseus (1990b).

In that case, the presence of penial chaetae in the

new genera might suggest their inclusion within

the Rhyacodrilinae.

Somatic chaetae. Somatic chaetae are generally

difficult to use in higher oligochaete taxonomy

because of a lack of consistency among and

within taxa. Both the northern telmatodrilines and

Breviatria and Biprostatus have some unusual

chaetae, but there are no chaetal features that

would indicate a close relationship between the

Australian and northern species. Neither do the

chaetae provide evidence of a relationship to any

other particular group of tubificids.

Sperm. Some Breviatria and Biprostatus

appear to have sperm in the spermathecae

arranged into numerous bundles with the nuclei

aligned at one end, others have the sperm cells in

loose masses, both arrangements typical of the

Rhyacodrilinae and Phallodrilinae. The sperma-

tozeugmata more typical of the Tubificinae, with

fertile sperm encapsulated by infertile sperm

(Braidotti and Ferraguti, 1982; Ferraguti et a!.,

1989; Ferraguti et aL, 1994), are absent. Of the

northern telmatodrilines, Alexandrovia species

have some kind of sperm bundles (Holmquist,

1974), which require ultrastructural analysis to

investigate homology with tubificine sperma-

tozeugmata, and Telmatodrilus have loose sperm

in the spermathecae. This character is thus

ambiguous in terms of relationships.

Spermathecal pores. In general, tubificines

have spermathecal pores near the middle of the

segment (Brinkhurst, 1991) while rhyacodrilines

and phallodrilines have these pores at or just

A. M. PINDER AND R. O. BRINKHURST

behind the anterior septa (Baker and Brinkhurst,

1981; Erseus, 1992). All Breviatria, at least one

of the two Biprostatus, and the northern telmato-

drilines have anterior spermathecal pores but,

since this may be the plesiomorphic state for the

family (Erseus, 1990b; Brinkhurst, 1991), it indi¬

cates little about the relationship between these

worms.

Penes. Pendant penes may constitute an apo-

morphy for tubificines and northern telmatodri-

lines (Erseus, 1990b; Brinkhurst, 1994), although

they are also present in some Phreodrilidae,

Absence of penes in Biprostatus and Breviatria

may be a plesiomorphic state shared by rhyaco-

drilines and phallodrilines.

In summary, the argument that Bre\’iatria and

Biprostatus belong within the Telmatodriiinae

simply because they have prostate cells formed

into multiple glands is not strong, particularly

considering the lack of knowledge about prostate

homology and the number of rhyacodrilines that

are now known to have clumped prostate tissue.

Breviatria and Biprostatus have several other fea¬

tures, including lack of pendant penes and sper-

matozeugmata and presence of penial chaetae and

coelomocytes (in at least some species) which

indicate a closer relationship with the Rhyaco-

drilinae. Therefore, as anticipated by the discus¬

sion of these and other problem taxa by Martin

and Brinkhurst (1998), we now propose to

consider the Australian former telmatodrilines to

be properly associated with Rhyacodrilinae,

described below as two new genera.

While this is undoubtedly not the last word on

the position of these worms, we believe that this

move represents a more parsimonious classifi¬

cation, pending further ontogenetic and ultra-

structural analyses of some characters (especially

prostate glands and organised sperm) and further

phylogenetic work on the rhyacodrilines. It does

seem that the Rhyacodrilinae is paraphylctic

(Erseus, 1990b; Brinkhurst, 1994) and we may be

trying to force the Australian ex-telmatodrilines

into a group that has no phylogenetic reality. A

thorough study of rhyacodriline phylogenetics is

yet to be undertaken but Erseus (1990b) can¬

vasses the possibility that the Rhyacodrilinae will

have to be split into a number of smaller groups to

achieve monophyly, perhaps with each raised to

family level. If this were to be given effect then

the Australian ex-telmatodrilines may form one

(or more) such groups.

Rhyacodrilus Bretscher

Rhyacodrilus Bretscher, 1901: 205.—Stephenson

1930: 742.—Brinkhurst, 1971a: 544.

Diagnosis. Dorsal chaetal bundles with or without

hair chaetae. Vasa deferentia moderately long,

entering atria subapically, atria usually bulbous or

rounded, usually covered with a diffuse layer of

prostate cells. Protrusible pseudopenes present or

absent. Penial chaetae usually modified,

numerous with blunt tips held close together.

Coelomocytes usually large and abundant.

Rhyacodrilus coccineus (Vejdovsky)

Tubifex coccineus Vejdovsky, 1875: 193_

Michaelsen. 1909: 32, figs 55-58.

Ryodrilus coccineus. —Stoic, 1886: 656.

Rhyacodrilus coccineus. —Michaelsen, 1909- 31 _

Brinkhurst, 1971a: 537, figs 8.31A-D.—Brinkhurst

1971b: 115, fig, 2K.

Branchiura pletirotheca Benham. 1907: 256 nl

XLVI. figs 7-12. ’

Holotype: Type locality unknown (ZMUH 6948).

Material examined. NSW: Blue Lake, Mt Kosciusko

36°24 24 S 148“18'50"E. Jun 1906, several on slides

(2 sectioned) (AMS W5612-W5613). WA: Power

Rock, Balladonia, 1 on slide, uncertain id. (ROB).

Distribution and habitat. Holarctic, Australia and

South America (Brinkhurst, 1971a; Brinkhurst

and Marchese, 1989). This species is not

commonly collected in Australia.

Diagnosis. Length 10-35 mm. Anterior dorsal

bundles with 3-5 hair chaetae and up to 5 pecti¬

nate chaetae, hairs missing in many posterior seg¬

ments. Ventral chaetae 2-5 per bundle with the

upper teeth thinner and not much longer than the

lower (but longer in Australian specimens

described as B. pletirotheca). Ventral chaetae of

XI with 3—5 club-headed penial chaetae per

bundle, equal in size to somatic ventTal chaetae.

Spemiathecal pores ventrolateral (lateral in the B.

pletirotheca specimens). Vasa deferentia not

coiled around atria, which are globular and

covered with a diffuse layer of prostate cells.

Coelomocytes large and abundant.

Remarks. Benham (1907) described Branchiura

pletirotheca from Blue Lake. Mt Kosciusko, but

Brinkhurst (197lab) considered this to be syn¬

onymous with R. coccineus. The long upper teeth

of the anterior ventral chaetae and the lateral posi¬

tion of the spermathecal pores of Benham’s mate¬

rial may prove to be diagnostic of a distinct

species once intra-specific variation in R. coc¬

cineus is better documented. Further specimens

Irom the type locality of B. pletirotheca (Blue

Lake, Mt Kosciusko) are required to substantiate

the synonymy. The Power Rock specimen is dis¬

sected but not stained and little can now be seen

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

of the genital anatomy. The most recent accounts

of species with hair chaetae are Ohtaka (1995)

and Timm (1990). The Tasmanian specimens

from Southwell R., mistaken as R. coccineus by

Pinder and Brinkhurst (1994), are attributable to

Ainudrilus fultoni Brinkhurst, 1982. Other worms

attributed to this species by Pinder and Brinkhurst

(1994) are described below as a new taxon.

Rhyacodrilus megaprostatus sp. nov.

Figure 1

Non Rhyacodrilus coccineus (Vejdovsky, 1875).—

Pinder and Brinkhurst, 1994: 94.

Material examined. Holotype: Curdies R„ Vic.,

(38°20'S 143°08E), 7 Feb 1992, G. Quinn (MON), 1

mature on slide (NMV F81869).

Paratypes. Vic: Grassy Creek, Otway Ranges,

38“28'S 144°00’E, 6 Jun 1989, 1 mature on slide (NMV

F81868): Lake Purrumbete, 38°17'S 143°14'E, 31 Jan

1992, G. Quinn (MON), 1 mature on slide (NMV

F81866); Lake Purrumbete, no date, A. Glaister

(MON), 1 mature on slide (NMV F81867); Branjee

Creek. Goulburn Valley, 36°42'S I48"20E, 23 Apr

1992, A. Pinder, 1 mature on slide (NMV F81870).

Distribution and habitat. All of the above locali¬

ties are in central southern Victoria. Collected

from lentic and lotic sites and from a variety of

Figure 1, Rhyacodrilus megaprostatus sp. nov.: A, tips of anterior dorsal chaetae with section of hair; B, tip of ven¬

tral chaeta of II; C, tip of posterior ventral chaeta; D, atrium with prostate tissue and penial chaetae of holotype; E,

spermatheca of NMV F81868. Scales: A-C, 0.01 mm; D, 0.04 mm; E, 0.03 mm.

A. M. PINDER AND R. O. BRINKHURST

habitats including sandy sediment and rocks and

bricks used for colonisation studies.

Etymology’: Named for the size of the prostate

gland in relation to the small atrium.

Description. Length 5-9 mm, diameter when

slide mounted about 0.3 mm. Number of

segments 40-70. Coelomocytes abundant.

Hair chaetae in preclitellar dorsal bundles nor¬

mally 3-4 per bundle (1 long and 1 short in one

Lake Purrumbete specimen), 270-300 pm long.

Hairs accompanied by 3-6 pectinate chaetae,

100-110 pm long, with upper teeth slightly

shorter than lower and few pectinations (Fig. 1 A).

Fewer dorsal chaetae posteriorly. Anterior ventral

bundles with 3 5 chaetae (those from Curdies R.

with as many as 7), 90-140 pm long, with upper

teeth thinner than lower, teeth about equal in

length on 11 (Fig. IB), upper teeth slightly shorter

than lower posteriorly (Fig. 1C). Penial chaetae in

XI bluntly simple-pointed, 90-100 pm long, 1-3

per bundle, projecting medial to the line of the

somatic ventral chaetae (Fig. D).

Paired male and spermathecal pores in line with

ventral chaetae.

Atria small, spherical (Fig. ID), opening into

smaller spherical chamber with a lobed pore.

Atria enclosed by a voluminous prostate gland

(Fig. I A), concealing the vas deferens-atrial

union. Prostate not closely applied to the sides of

the atrium. Ovaries large and obscuring the vasa

deferentia. Male funnels observed in some speci¬

mens, often closely associated with the sper-

mathecae and a part of vasa deferentia folded and

adhered to funnels. Spermathecae ovoid (Fig. IE)

containing sperm arranged in distinct masses.

Remarks. The apparent attachment of the prostate

at the top of the atrium needs to be confirmed

from sectioned material as this may indicates that

the prostate is not as diffuse as is normal in

Rhyacodrilus. Within the genus, the large volume

of prostate tissue compared to the atria is unique

to this new species.

According to the key to Rhyacodrilus in

Brinkhurst (1971a) and notes in the update by

Brinkhurst and Wetzel (1984), R. megaprostatus

is closest to R. coccineus, from which it differs

mainly in the volume of the prostate. Several

other species with hair and pectinate chaetae have

been described more recently. In Rhyacodrilus

okamikae Giani and Rodriguez, 1988 the pecti¬

nate chaetae have short upper teeth, the ventral

chaetae have long upper teeth and there is only a

single median spermatheca. Rhyacodrilus supu-

tensis Timm, 1990 has lyre-shaped pectinate

chaetae and tubular atria. Rhyacodrilus hiemalis

Ohtaka, 1995 has voluminous spermathecae with

narrow ducts, more numerous chaetae and a less

voluminous prostate gland.

Rhyacodrilus bifidus Brinkhurst

Rhyacodrilus bi/idus Brinkhurst, 1982: 3, pi. 6.

Holotype: Blue Lake, Mt Kosciusko plateau, NSW

(QVM: 14:232).

Material examined: NSW: Holotype and paratype from

type locality. 36°24'24"S I48°18'5(f E, 7 Feb 1977, 5

on slides (QVM: 14:232-233) and 7 on slides (ROB);

Lake Cootapatamba, Mt Kosciusko plateau,

37 n 27'58"S I48"15'48"E, 7 Feb 1977, 1 on slide

(ROB). Vic.: Thomson R. at Thomson Portal Rd, 3 on

slides (NMV F81671-81873); Wentworth R„ 17.7 km

upstream of Tabberabbera, 37“31 'S 147°23'E, 2 Oct

1975, 1 on slide, tentative identification (NMV

F81874). New Zealand: Lake Matheson, Canterbury

Highlands, 2 Nov 1978, 2 on slides (ROB); Lake Gault.

Canterbury Highlands, 5 Nov 1978, 2 on slides (ROB).

Distribution and habitat. Known only from the

above lakes and rivers of the far southeast of

mainland Australia and the lakes on the South

Island of New Zealand (Brinkhurst, 1982),

although the identification of the latter could not

be confirmed as the genitalia are no longer visible

in the specimens examined. Australian specimens

have been collected from coarse sediments in

rivers: sand and gravel, often between cobbles

and pebbles (Ahern and Blyth, 1979; Malipatil

and Blyth, 1982) but possibly in finer sediments

from the Mt Kosciusko sites (Timms, 1979).

Diagnosis. Length up to 7 mm. Ventral and dor¬

sal bundles with 3-8 chaetae anteriorly, bifid with

upper teeth 2-3 times as long as, but thinner

than, the lower. Penial chaetae 6-10 bifid per

bundle in XI, about 2 times longer than somatic

chaetae and slightly curved ectally. Vasa deferen¬

tia as long as atria. Atria 4 times longer than

broad, ending in wide bulbs with narrow pores on

the anterior face of penis sacs which open to the

exterior via narrow pores. Abundant prostate

tissue attached to atria subapically. Coelomocytes

not abundant.

Remarks. The reproductive structures of the

Wentworth R. specimen are not visible and so the

identification cannot be confirmed. There are

only three penial chaetae per bundle in XI, but

these are large and bifid and the somatic chaetae

are mostly four or five per bundle with the upper

teeth much longer than the lower. Similarly, the

identification of the whole-mounted New Zealand

specimens could not be confirmed.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

Ainudrilus Finogenova

Ainudrilus Finogenova, 1982: 1255.—Erseus, 1990a.

Vadicola Baker, 1982: 3232.

Type species. Ainudrilus oceanicus Finogenova,

1982.

Diagnosis. Flair chaetae present or absent.

Chaetae of penial segment modified, chaetae of

spermathecal segment slightly modified in one

species. Vasa defcrentia usually wide and at least

partially granulated, entering atria subapically.

Atria variably shaped but more or less erect or

directed posteriad, usually consisting of an

ampullae, often with spacious lumen contain¬

ing sperm, usually leading to thick, and often

equally large and well developed, ejaculatory

ducts, often giving the appearance of a bi-partite

atria. Prostate absent. Penes absent. Spcrmathecae

with distinct, and often complex ducts. Sperm

loose in ampullae. Coelomocytes large and

abundant.

Remarks. The lack of prostate on the atria distin¬

guishes Ainudrilus from almost all other rhyaco-

drilines, bar some other highly unusual forms

such as Jolydrilus Marcus, 1965 and Epirodrilus

Hrabe, 1930 (see Brinkhurst, 1971a; Baker and

Brinkhurst, 1981), and one freshwater Rhyaco-

drilus. The latter, Rhyacodrilus simplex (Benham,

1903), lacks prostate tissue but also differs from

other Rhyacodrilus in having vasa deferentia that

coil around the atria and which join the atria api-

cally. Further study of this species may lead to it

being moved back to a resurrected Taupodrilus ,

which was established by Benham (1903) for its

reception.

Ainudrilus billabongus (Brinkhurst, 1984),

Ainudrilus stagnalis (Erseus, 1997) and the fol¬

lowing new species and new combination are the

only freshwater representatives described so far.

There arc eight marine species, from Hong Kong,

Northern Australia, the Pacific Ocean and the

Caribbean (Erseus, 1990ab, 1997; Finogenova,

1982).

Ainudrilus billabongus (Brinkhurst)

Rhyacodrilus billabongus Brinkhurst, 1984: 144,

fig. 3.

Ainudrilus billabongus. —Erseus, 1990a: 265.

Hololvpe. Bowerbird Billabong, Magela Creek, NT

(USNM 80698).

Material examined: NT: Paratype, from type locality, 1

on slide, 21 Aug 1981 (ROB); Magela Creek, 24 Feb

1988, 1 on slide (ER1SS).

Distribution and habitat. Recorded only from

lentic and lotic sites of the Magela Creek system.

Northern Territory.

Diagnosis. Length at least 7.5 mm, all specimens

incomplete. Coelomocytes large and abundant.

Anterior chaetae bifid with teeth of equal length,

4-6 per bundle anteriorly, reduced in number pos¬

teriorly. Up to 6 simple-pointed penial chaetae in

each ventral bundle of XI, strongly recurved at the

tip and about 2 times longer than somatic chaetae.

Vasa deferentia wide and glandular, about twice

the length of the muscular upright atria, entering

the atria basally. Atria leading directly to small

circular pores. Spermathecal ducts long and

narrow, sharply set of from the globular ampullae.

Remarks. The original description of A. bill¬

abongus made mention of a bundle of sperm lying

outside of the atrium. Most fully mature Ain¬

udrilus seem to maintain sperm in the atrium or

expanded ejaculatory ducts so the sperm bundle

of the holotype illustrated in Brinkhurst (1982)

was most likely extruded from the atrium during

dissection.

Ainudrilus stagnalis Erseus, 1997

Ainudrilus stagnalis Erseus, 1997: 9, figs 2E-F.

Holotype. Billabong associated with Stephens Creek,

Port Darwin West Arm. NT (NTM W00020).

Material examined. No new specimens examined.

Distribution and habitat. Known only from the

freshwater type locality, in fine sand (Erseus,

1997).

Diagnosis. Length at least 3.2 mm (only specimen

incomplete). Coelomocytes abundant. Hair

chaetae absent. Anterior chaetae bifid with upper

tooth much longer than lower, 4-6 per bundle

anteriorly, 3-4 posteriorly. A single bifid chaeta,

not larger than other ventral chaetae, near each

male pore. Vasa deferentia not described. Atria

with non-muscular ampullae and thick muscular

ejaculatory ducts terminating in simple pores

medial to the ventral chaetal line. Spermathecae

absent in the only known specimen.

Remarks. Erseus (1997) suggested that the chaeta

near each male pore is not a modified genital

chaeta, but that such might be developed at a later

stage.

Ainudrilus fultoni (Brinkhurst) comb. nov.

Figure 2

Rhyacodrilus fultoni Brinkhurst, 1982: 2, pi. 1-5.

Holotype. Lake Sorell, Tas. (QVM: 14:230).

A. M. PINDER AND R. O. BRINKHURST

A B

Figure 2, Ainudrilus fultoni (Brinkhurst) from Lake Sorell: A, somatic chaetae; B, penial chaetae; C, genitalia of

QVM: 14:4202. Scales: A-B, 0.01 mm; C, 0.05 mm.

Material examined. Tas.: Holotype and paratypes.

type locality, several sectioned specimens and others in

alcohol, 2'Feb 1981 (QVM: 14:230-231 and ROB);

Southwell R., 41°34 - 50 "S I45"44’04"E. 20 Dec 1988,

2 on slides (QVM: 14:4201 4202); Ouse R. at Marlbor¬

ough Highway, 41°59'22”S I46°38'43"E. 16 Mar

1997, I on slide (AMP); Lake Perry. 43°1310"S

146”45I7"E. I I Mar 1997, 2 on slides, 3 in alcohol

(AMP).

Distribution and habitat. Known only from lakes

and rivers in Tasmania. The Ouse R. and Lake

Perry specimens were collected from sand

between cobbles in shallow water.

Diagnosis. Length up to 15 mm. Coelomocytcs

large and fairly abundant. Chaetae all bifid with

teeth equally long, 10-13 per bundle anteriorly,

fewer posteriorly (Fig. 2A). Ventral bundles of XI

with 7-9 straight penial chaetae with blunt,

slightly curved tips, about equal in length to

somatic ventral chaetae (Fig. 2B). Short vasa

defercntia enter atria subapically (Fig. 2C). Atria

consisting of a thick walled ental portion with a

narrow lumen, which leads into a thinner walled

ectal portion with a wide lumen which usually

contains sperm in mature specimens (Fig. 2C).

Spcrmathecae with ampullae full of loose masses

of sperm, ducts short and broad opening through

large pores (Fig. 2C).

Remarks. Specimens from Southwell R, were

attributed to R. coceineus by Pinder and Brink-

hurst (1994) but conform to the description of A.

fultoni. As these specimens are dissected, rather

than sectioned like the type material, the repro¬

ductive structures arc illustrated here to give a dif¬

ferent perspective (Fig. 2C). The atrium consists

of a thick-walled portion in which the lumen

can be seen in only one dissected specimen and a

thinner walled octal chamber (essentially an

ejaculatory duct), with a sperm bundle within the

latter in examined specimens.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

It seems preferable to transfer R. fultoni from

Rhyacodrilus to the expanding Ainudrilus, which

includes most species without prostate glands,

whereas all Rhyacodrilus (except R. simplex,

discussed above) have diffuse prostate tissue.

Ainudrilus fultoni can be distinguished from its

freshwater congeners by the more numerous

somatic and penial chaetae.

Ainudrilus nharna sp. nov.

Figure 3

Material examined. Holotype: From Frankland R. at

Roe Rd ford. Western Australia, 34"41'02''S

116°5ri3 "E, 9 Sep 1996, A. Pinder and R. Brinkhurst,

serially sectioned specimen on three slides (WAM

1-99).

Paratypes; Collection details as for holotype, 2 spec¬

imens serially sectioned on slides and others in alcohol

(WAM 2-99 to 4-99).

Other material: WA: Collection details as for types, 7

on slides, including 1 serially sectioned (AMP); Lake

Wilson at track off Scott Rd. 34°25'23"S

115°07’52"E, 14 Sep 1996, 2 on slides (AMP);

Thomas Spring, Augusta, 34°2rOO"S 115°09'35"E,

17 Sep 1996, A. Pinder and R. Brinkhurst, 5 on slides

(AMP); stream crossing Thomson Rd, approximately 5

km N of Granite Peak, 34"40'48"S 116°42'13 "E, 12

Sep 1996, A. Pinder and R. Brinkhurst, 1 on slide

(AMP); northern tributary of Collier Creek on

Cemetary Rd, 34“58'30"S 116°45'12 "E, 11 Sep 1996,

A. Pinder and R. Brinkhurst, 1 on slide (AMP); Boyn-

diminup Rd, Lake Muir wetland system, WA, 34°30'S

116“35 E, Jul and Oct 1993, P. Horwitz (ECU), 3 on

slides and others in alcohol (WAM 90-97, 91-97,

Figure 3, Ainudrilus nharna sp. nov.: A, tip of dorsal chaeta; B, tip of ventral chaeta; C, tip of penial chaeta; D, gen¬

italia, based mostly on holotype. Scales: A-C, 0.01 mm; D, 0.1 mm.

A. M. PINDER AND R. O. BRINKHURST

92- 97, 98-97 and 99-97); Peaty seepage in logging

coup, Johnsons Rd, upper tributary of Deep R.. 34°41 'S

116°36'E, Jan 1993, P, Horwitz (ECU), 4 on slides

(WAM 95-97 to 97-97, 93-97); creek in sandy shrub

land at Bevan Rd, Kent R. catchment, 34°39'S

116°58 E, P. Horwitz (ECU). Oct 1993. 1 on slide

(WAM 94-97); Beedelup Brook South, 34°22'39 "S

115“56 E, Sep 1993. K. Trayler (MUR). 1 on slide

(WAM 36 98); Beedelup Brook South. 34”22'50"S

I15“55 E, Oct 1994, K. Trayler (MUR), 1 on slide

(WAM 37-98); Carey Brook at Pile Rd, 34‘ > 21'29"S

I15“54'29"E, 16 Sep 1996, A. Pindcr and R.

Brinkhurst, I on slide and 3 in alcohol (AMP); Lake

Walbyring, 32°50'I9 "S 117°35'27”E, 18 Dec 1996,

M. Smith (CALM), I on slide (WAM 38-98) and 2 on

slides (AMP).

Distribution and habitat. Ainudrilus nharna

appears to be widespread in the southwest of WA,

occurring in springs, lakes, and both flowing and

still reaches of streams and rivers. Sediments at

collecting sites ranged from peat to mud to coarse

sand. Specimens from Lake Walbyring, a brack¬

ish to saline lake, were collected in Dec 1996

when salinity in the lake was 2.8 ppt, but not in

Oct 1997 when salinity was 20 ppt (unpublished

data).

Etymology: From nharna , worm in Nhungar

aboriginal language.

Description. Length about 9 mm. Number of

segments 75-90. Width at clitellum (slide-

mounted worms) 0.3-0.5 mm. Coeiomocytes

present.

Clitellum inconspicuous, half-X-half-XII.

Gonopores in line with the ventral chaetae, sper-

mathecal pores anterior on X. Male pores open

into a transverse groove, lateral to the penial

chaetae in the anterior half of XL

Anterior dorsal bundles with up to 4 hair

chaetae (up to 420 pm long) and up to 5 pectinate

chaetae (up to 130 pm long), latter with upper

teeth thinner than and equal in length to lower,

with indistinct pectinations (Fig. 3A). Ventral

bundles with bifid chaetae of similar size and

form to the dorsals (but without pectinations), 3-5

per bundle anteriorly, fewer posteriorly (Fig. 3B).

Penial chaetae 4-7 per bundle in XI, up to 120 pm

long and slightly curved ectally (Fig. 3C). Penial

chaetae protrude close to the mid-line, on cither

side of the ventral nerve cord medial to the male

pores.

1 cstes anteroventral in X (Fig. 3D). Atria in XI,

bulbous with thick lining tissue and ciliated

lumen, leading to folded protrusible pseudopenes,

which are protruded on one of the paratypes.

Prostate tissue absent. Vasa deferentia thickened.

appearing glandular and often twisted or folded

once or twice between male funnel and atria,

attaching anteromedially to the atria (Fig 3D).

Spennathecal ampullae large and ovoid with

sperm formed into loose masses, clearly separated

from the ducts. Spermathccal ducts constricted a(

one or more points, leading to pores which are

often on a protrusion within small vestibulae,

located anteriorly on X (Fig 3D). Ovaries lying

near the vasa deferentia in XI, female pores not

observed.

Remarks. The absence of prostate tissue and the

thickened vasa deferentia suggest that this new

species belongs in the genus Ainudrilus. Ain¬

udrilus taitamensis Erseus, 1990a, of Hong Kong,

which was only tentatively included in the genus,

is the only other species to have ciliated atria. The

atria and pseudopenes resembles what was called

a two-part atria in A. fultoni and the ectal pail of

the male duct is formed into pscudopenes or elab'

orate ejaculatory ducts in some other species;

(Erseus, 1990a). The thick vasa deferentia are dif¬

ficult to follow in both sectioned and dissected

worms as they are tightly folded with an indistinct

lumen and are obscured by the ovaries. Ainudrilus

taitamensis and Ainudrilus bremlae Erseus, 1997,

both marine/brackish, are the only other Ain¬

udrilus to have hair chaetae. These differ from the

new species in that the former has long tubular

atria and the latter has only two small straight

penial chaetae per bundle and ventral chaetae

with upper teeth much longer than the lower.

Rhizodrilus Smith

Rhizodrilus Smith, 1900: 44.—Baker and Brinkhurst

1981:952.

Monopylephorus Levinson (in part).—Brinkhurst,

1963: 62.—Brinkhurst, 1965a: 148.—Brinkhurst'

1971a: 555.

Type species. Rhizodrilus lacteus Smith, 1900.

Diagnosis. Chaetae all bifid, hairs absent. Vasa

deferentia moderately long, entering atria subapi-

cally. Atria normally tubular with diffuse prostate

tissue, terminating in ejaculatory ducts which exit

either on the inner aspect of large ventrolateral

tolds in the body wall or into a median invagina¬

tion of the ventral body wall (copulatory bursa),

sometimes via protrusible pscudopenes. Sper-

mathecal chaetae and/or penial chaetae modified

in mature specimens, the former associated with

large glands. Some or all of the spermathccal gen¬

italia in IX. Sperm in spermathecae in loose

masses or in oriented bundles. Coeiomocytes

abundant.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

Remarks. Rhizodrilus consists of a few marine

and freshwater species with apparently disparate

distributions (Baker and Brinkhurst, 1981;

Erseus, 1990a). These are united by the presence

in IX of at least some of the spermathecal geni¬

talia. The following species was tentatively

included in Rhizodrilus by Baker and Brinkhurst

(1981).

Rhizodrilus arthingtonae (Jamieson)

Rhyacodrilus arthingtonae Jamieson, 1978.

Rhizodrilus arthingtonae .—Baker and Brinkhurst,

1981: 956.

Holotype. Brown Lake, North Stradbroke Island, Qld

(QMG8881).

Material examined: Qld: Holotype and paratype, local¬

ity as above, 31 Jul 1975, dissected on slides (QM

G8881-2).

Distribution and habitat. Known only from the

type locality, a perched acidic lake. Specimens

collected in organic material in sandy sediment

(Jamieson, 1978).

Diagnosis. Length 66 mm (paratype). Coelomo-

cytes numerous. Anterior chaetal bundles with

3-5 bifid chaetae with the upper teeth slightly

longer than the lower, posteriorly teeth about

equal. Hair chaetae absent. The chaetae are

smaller anteriorly and posteriorly, largest in the

clitellar region. Several clearly bifid penial

chaetae per bundle in XI, 1.5-2 times longer than

somatic chaetae and slightly curved towards the

tip. Vasa deferentia narrow, coiled anteriorly,

passing through the prostate before entering atria

subapically. Atria fusiform to slenderly bulbous,

prostate tissue compact but lobulated, enclosing

the atrium and communicating with the lumen at

numerous points. Slender ejaculatory ducts lead

to male pores on the inner wall of a ventrolateral

fold in the body wall. Accessory genital markings

present, consisting of a pair of conspicuous ellip¬

tical papillae posteromedian to ventral bundles of

IX, each with a short diagonal groove passing

posterolaterally from its centre to 9/10. Sper-

mathecae in X, with indistinct ventrolateral pores

at 9/10. Worm squarish in cross-section.

Remarks. Erseus (1994) rightly questions whether

R. arthingtonae belongs within tins genus, prefer¬

ring to include only those species with large tubu¬

lar atria and a clear involvement of spermathecal

genitalia in IX, rather than X as is normal in the

family. R arthingtonae has only minor involve¬

ment of spermathecal genitalia in IX (just

papillae on IX with grooves leading back to the

spermathecal pores on 9/10). However, Rhizo¬

drilus lacteus Smith, 1900, which has just the

spermathecal chaetae and associated glands in IX

but retains the spermathecae themselves in X is

intermediate between R. arthingtonae and other

Rhizodrilus in this respect and so R. arthingtonae

may represent one extreme of a continuum. Argu¬

ing against this proposition is the fact that the

atria are more upright and less tubular than those

of other Rhizodrilus and Ferraguti et al. (1994)

note the more plesiomorphic spermatozoa! ultra-

structure of R. arthingtonae compared to Rhizo¬

drilus russus Erseus, 1990a and the related

Monopylephorus Levinson, 1884. If R. arthing¬

tonae is excluded from Rhizodrilus , then, short of

creating a new genus, it is difficult to see a more

suitable place for it. Erseus (1994) summarises

the possible relationships between Rhizodrilus

and oilier relevant rhyacodriline genera and, since

we suggest no change to the status quo, these need

not be repeated here. A more satisfactory place¬

ment of R. arthingtonae may have to await further

phylogenetic studies of this and related rhyacodri-

lines.

Breviatria gen. nov.

Type species. Telmatodrilus multiprostatus

Brinkhurst, 1971a, as the first of two species

described in the original account.

Diagnosis. Atria ovoid to erect pear-shaped, with

unciliated lumen, bearing numerous glands of

prostate tissue apically, communicating indepen¬

dently with atrial lumen. Male pores simple on

XI, on inner aspect of protruding ventrolateral

folds of the body wall surrounding a broad

median copulatory chamber. Spermathecal

chaetae large and grooved, penial chaetae numer¬

ous with blunt or bifid tips. Spermathecae nor¬

mally with ventrolateral to dorso-Iateral pores,

located anteriorly on X. Sperm in spermathecae in

organised bundles (not spermatozeugmata) or

loose. Coelomocytes abundant for at least one

species, unknown in the others.

Etymology. Breviatria referring to the short atria,

in contrast to the tubular atria of other genera with

prostate formed into multiple clumps.

Remarks. Of the four species assigned to this new

rhyacodriline genus, three were previously placed

within the Telmatodrilinae and one is new. The

reasons for re-assigning the earlier species were

discussed above (in the Rhyacodrilinae remarks

section). Breviatria is distinguished from other

Rhyacodrilinae by the ovoid to pear-shaped atria

A. M. PINDER AND R. O. BRINKHURST

with prostate tissue formed into discrete glands

with separate (usually stalked) connections to the

ectal part of the atria. Other rhyacodriline genera

with clumped prostate tissue, mostly marine

forms such as Heronidrilus Erseus and Jamieson,

1981 and Heterodrilus Pierantoni. 1902, usually

have the clumps more broadly attached along the

length of elongate tubular atria, although

prostates are more restricted in position in a few

species. The other new genus, Biprostatus,

described below, also has small atria but these

have only two large prostate glands which are

broadly attached to the anterior and posterior

sides of the atria. Other rhyacodrilines with short

atria either have evenly distributed diffuse

prostate tissue (such as most Rhyacodrilus ) or

lack prostate tissue (Ainudrilus and Rhyacodrilus

simplex). Rhizodriltts arthingtonae (discussed

above) also has ovoid atria but the lobes of

prostate tissue are tightly packed over the surface

of the atria and broadly attached rather than

stalked. The new species described below has

numerous coelomocytes but this feature needs to

be re-examined when fresh material of the other

species becomes available.

Breviatria multiprostatus (Brinkhurst)

comb. nov.

Figures 4, 5

Teimatodrilus multiprostatus Brinkhurst, 1971a: 535,

figs 8.27A-F.—Brinkhurst, 1971b: 117, fig. 3C.—

Brinkhurst and Fulton. 1979: 3.—Brinkhurst, 1982: 3,

pi. 7.- Holmquist, 1974: 256, figs 10-11.—Bayly.

1973: 305. -Timms. 1978: 430.—Marshall. 1975: 22,

fig. 3.- Marshall and Winterboum. 1979: 205. The

species description was duplicated in Brinkhurst

(1971b) but the 1971a publication should be used as the

original description.

Holotype. Lake Pedder, Tas. (AMS W4I78).

Material examined. Tas.: Holotype and paratypes,

locality as above, 2-4 Mar 1966 (AMS

W4178-W4182) and I on slide (ROB); Cuvier R„ 10

Oct 1978, 1 on slides (ROB); Collingwood R.. 4 Nov

1978, I on slides (ROB); trickle under the Needles, 11

Nov 1978, 1 on slides (ROB); creek in Frodshams Pass.

12 Nov 1978, I on slides (ROB); Lake Sorell. 2 Feb

1981, 1 sectioned (ROB); Crossing R.. south of Lake

Pedder. southwest Tasmania, 10 Feb 1987, 1 on slide

(QVM:14:3002) and 3 in alcohol (AMP): Lake Sur¬

prise, AMG 8112 441869. 13 Nov 1976, 1 on slide

(QVM: 14:3001) and I in alcohol (AMP). Tentatively

identified immature specimens from various sites in

Tasmania listed by Brinkhurst (1982).

Other records. Tas.: Lake Pedder (Bayly, 1973): South

Esk R. (Brinkhurst and Fulton, 1979); Lakes Sorell and

Crescent (Timms, 1978). New Zealand: Leeston Drain,

near Killinchy, 56 km S of Christchurch (Marshall,

1975; Marshall and Winterboum, 1979); Freshwater R.,

Stewart Island (Cowie et al., 1978).

Distribution and habitat. Recorded from lakes,

streams and rivers in Tasmania and possibly from

New Zealand. The Lake Crescent specimens were

collected from a variety of sediment types,

including sand, detritus and mud (Timms, 1978).

Specimens from Great Lake were collected at

depths of up to 16.5 m (Fulton, 1983b).

Description. Length up to 16 mm. Pharyngeal

glands present on either side of septa 2/3 to 4/5,

gut expanded in VII to form a diverticulum which

completely encircles the gut in VII and protrudes

forward into VI (Fig. 4A).

Somatic chaetae all simple-pointed (Fig. 5A),

up to 8 per bundle anteriorly, fewer posteriorly.

Ventral chaetae missing on penial segment of

mature specimens, or if present then not modified.

Spermalhecal chaetae (Fig. 5B) larger than

somatic chaetae, with extensive musculature,

projecting into an enlarged chactal sac poster¬

iorly on X, into which also empties a separate

elongate gland (Fig. 4C). In one paratype, the

spermathccal chaetal sac opens anteriorly in X,

with the chaetal shaft protruding well anteriad

into IX (Fig. 4B).

Testes anteroventral in X, with sperm funnels

on 10/11 feeding broad vasa deferentia which

wind upwards to enter atria apically (Figs 4B, C).

Atria elongate pear-shaped, may be folded over

dorsally, with unciliated lumen and thick lining

tissue. Atria narrow basally, opening on inner

aspect of folds in the body wall lateral to a median

ventral copulatory chamber. Prostate cells

grouped into numerous glands which narrow

basally and connect to apical third of atria (Figs

4B, C, Fig. 5C). Spermathecae large in X with

short ducts that lead to ventral pores just behind

septa 9/10 (or medially in X in the Lake Sorell

specimen). Sperm loose in ampulla. Ovaries

attached to anterior septa of XI near sperm

funnel. Female funnels large, leading to pores at

intersegmental furrow 11/12 (Figs 4B, C).

Remarks. This is the only Australian tubificid to

have all chaetae simple-pointed. However, many

aquatic Enchytraeidae also have multiple simple-

pointed chaetae in all bundles and immature

forms could be confused with B. multiprostatus.

Of the Tasmanian records, the only identifications

that can be confirmed by re-examination of spec¬

imens are those from Lake Pedder and Lake

Sorell. Specimens from the other sites listed

above are either immature (and so are tentatively

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

Figure 4, Breviatria multiprostatus (Brinkhurst): A, anterior of specimen from Lake Sorell; B, genitalia of paratype

AMS W4I82; C, genitalia of Lake Sorell specimen. Scales: A-C, 0.15 mm.

A. M. PINDER AND R. O. BRINKHURST

Figure 5, Breviatria multiprostatus (Brinkhurst): A, typical somatic chaetae; B, bent spermathecal chaeta of holo-

type; C, atria dissected from holotype. Scales: A, 0.02 mm; B, 0.05 mm; C. 0.2 mm.

identified from the chaetae only) or their identifi¬

cation can no longer be verified because of their

poor condition. It is not known whether identifi¬

cations of specimens from New Zealand were

based on the genitalia of mature specimens or just

the chaetae of immature worms so the identifica¬

tion must be considered tentative. One of the New

Zealand records (Leeston Drain) is unusual in that

the site was enriched with septic tank leachate and

dairy effluent (Marshall and Winterbourn, 1979)

whereas the Australian records are all from non-

enriched sites, including minimally disturbed

alpine streams.

The position of the spermathecal pores and

spermathecal chaetae appears to vary. Paratypes

W4180 and W4182 (Fig. 4B) both have sper¬

mathecal pores anterior in X, but spermathecal

chaetae posterior and anterior (projecting forward

into IX) in X respectively. The Lake Sorcll spec¬

imen has spermathecal chaetae posterior in X but

the spermathecal pore medial on X (Fig. 4C). All

other features of these worms seem similar and

we consider the variation in spermathecal charac¬

ters to be intraspecific for now. The holotype is

dissected rather than sectioned and the original

position of the spermathecae is unclear.

Breviatria pectinatus (Brinkhurst) comb. nov.

Figure 6

Telmatodriluspectinatus Brinkhurst, 1971a: 535. figs ’

8.25D-H.—Brinkhurst, 1971b: 117. fig. 3D— Bayly.

1973: 305.—Holmquist, 1974: 257, fig. 12.—Timms,

1974: 276.—Timms, 1978: 435. The species descrip¬

tion was duplicated in Brinkhurst (1971 b) but the 1971a

publication should be used as the original description.

Holotype. Lake Pedder, Tas. (AMS W4176).

Material examined. Tas.: Holotype and paratypes. Lake

Pedder, 2 Mar 1966, 2 on slides (AMS W4176-7) and

5 on slides (ROB).

Other Australian records. Tas.: Lake Pedder (Bayly,

1973); Lake St Clair (Timms, 1978). Vic.: Lake Tali

Karng (Timms, 1974).

Distribution and habitat. Recorded only from the

above lakes in Victoria and Tasmania. Bayly

(1973) collected this species from Lake Pedder

(in 1972) in sandy sediment but the Lake Tali

Karng specimens were recovered from mud at a

depth of 12 m.

Description. Length unknown, but greater than 10

mm.

Anterior bundles with 9-14 chaetae, upper

teeth longer than the lower and slightly hooked.

Posterior bundles with 5-8 pectinate chaetae with

upper teeth not so long (Fig. 6A). Spermathecal

chaetae single, thin with grooved tips (Fig. 6B),

up to twice as long as somatic ventral chaetae.

Penial chaetae (Fig. 6C) modified with rough,

blunt to bifid tips (absent on paratype W4176),

about 1.5 times longer than somatic ventral

chaetae.

Vasa deferentia broad, coiled once or twice

before entering atria apically (Fig. 6D). Atria

pear-shaped with numerous prostates with short

stalks distributed around the upper half. Stalks

enveloped by the atrial muscle tissue. Individual

peritoneal cells form diffuse cover over basal half

ot atria and some present apically (Fig. 6D). Atria

narrowing to pores on the inner aspect of lateral

folds in the body wall around a shallow median

ventral copulatory chamber. Details of the

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

spermathecae difficult to decipher but possibly

quite folded, with lateral or dorsolateral pores and

containing loose sperm.

Remarks. Pectinate chaetae are more common in

the Tubificinae than the Rhyacodrilinae and the

former usually has pectinations more devel¬

oped anteriorly, rather than posteriorly as in

B. pectinatus.

Figure 6, Breviatria pectinatus (Brinkhurst): A, tips of pectinate dorsal chaetae of holotype; B, spermathecal chaeta

of holotype; C, penial chaetae of holotype; D, male ducts and spermathecal chaetae of paratype AMS W4177.

Scales: A, 0.01 mm; B-C, 0.015 mm; D, 0.05 mm.

A. M. PINDER AND R. O. BRINKHURST

Breviatriapapillatus (Brinkhurst and Fulton)

comb. nov.

Figure 7

Telmatodrilus (Alexandrovia) papillatus Brinkhurst

and Fulton. 1979: 2, figs 9-14.—Fulton, 1983a: 778.—

Fulton, 1983b: 792.

Holotype. Great Lake, Tasmania (QVM: 14:320 and

321).

Material examined. Tas.: Holotype and paratypes, type

locality, 26 Sep 1975, 3 on slides (QVM: 14:320, 321,

326 and 327) and 1 on slide (ROB); Arthurs Lake, 4 Jul

1977, 8 on slide, 2 mature and several immature in alco¬

hol (NMV F88870, F88871 and ROB); Lake Sorrel, no

date, 4 on slides (QVM: 14:322-325).

Distribution and habitat. Recorded only from the

above lakes of the central plateau of Tasmania.

Collected in both sandy and clayey sediments, a t

depths up to 17.5m (Fulton, 1983b).

Description. Length 10-20 mm. C’oelomocytes

not observed. Worms encrusted with fine foreign

material, body wall papillate with several rings 0 f

papillae per segment (Fig. 7A). Prostontium and

segment I (and sometimes 11) may be retracted

into the body. Posterior segments narrow and

Figure 7, Breviatria papillatus (Brinkhurst and Fulton): A, view of body showing copulatory bursa on XI; B, tip of

anterior chaeta; C-D, tips of transitional chaetae; E, tip of posterior hair chaeta; F, male ducts of NMV F88870; G,

spermatheca of NMV F88870. Scales: A, 0.2 mm; B-E, 0.01 mm; F, 0.05 mm; G, 0.025 mm.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

appearing highly contracted in the preserved

specimens.

Anterior chaetae bifid, 5-7 per bundle, with

each tooth broad and flattened (Fig. 7B). Chaetae

changing to a hair-like form, some with vestiges

of teeth (Figs. 7C, D) between VII and X so that

postclitellar bundles have 5-9 curved hair chaetae

(Fig. 7E). Penial chaetae slightly shorter than pre-

elite] lar somatic chaetae, 3-5 per bundle with

bifid tips (Fig. 7F). Spermathecal chaetae not

modified.

Long vasa deferentia connect to short pear-

shaped atria subapically (Fig. 7F). Atria narrow

gradually to enter small median copulatory cham¬

ber (which is everted in some preserved worms)

near the penial chaetae which project from the

protruding lateral walls of the chamber. Three or

more lobes of prostate tissue on atria apically

(Fig. 7F). Spermathecae small and globular

(Fig. 7G) with short ducts leading to lateral

pores anteriorly on X. Sperm not observed in

ampullae.

Remarks. While the chaetae and papillae of B.

papillatus are unusual, the genitalia suggest that

this species is simply an apomorphic member of

Breviatria and to exclude it would leave Brevia-

tria paraphyletic. Specimens from Arthurs Lake,

some of which were dissected and illustrated for

this study (Fig. 7), clearly have spermathecae

with lateral pores whereas spermathecae were not

seen in the type specimens from Great Lake. The

spermathecae of these specimens are very small

(100-150 pm) and do not contain sperm, so may

not be fully developed. Unfortunately, the type

specimens have become too cleared for this fea¬

ture to be checked. The holotype (the only type in

which the genitalia can be seen) has ‘at least 2 or

3 prostate glands’ (Brinkhurst and Fulton, 1979),

but the size of these and their attachment to the

atria is uncertain due to clearing and distortion

during slide mounting. The prostate on one atrium

of the holotype appears to be formed into two or

three large, stalked lobes but on the other atrium

the prostate appears to be a more cohesive mass

with a number of short connections. The prostate

tissue of the Arthur R. specimens is clearly

formed into numerous small glands but connec¬

tions to the atria are obscured. Fresh specimens

from the type locality are required to determine

the unity of this taxon.

Breviatria arvensis sp. nov.

Figure 8

Material examined. Holotype: Arve R. at Arve Rd,

43°11 '20"S 146°46'05"E, Tasmania, 12 Mar 1997, R.

Marchant (NMV) and A. Pinder. dissected on slide

(QVM: 14:3878).

Paratypes: Collection details as for holotype, 1

mature and 1 immature sagitally halved on slide, with

posterior portion of 1 of these in alcohol, and 1 imma¬

ture whole-mounted on a slide (QVM: 14:3879-3881).

Distribution ami habitat. Known only from the

type locality, collected from sand/silt under

cobble at a depth of 10-20 cm.

Etymology: Named arvensis for the type locality.

Description. Length at least 20 mm (none com¬

plete), width of slide mounted worms 0.25-0.3

mm. Prostomium squarish in preserved type

material. Coelomocytes abundant but without

consistent shape, entirely filling coelom in most

of body, including the prostomium, making the

worms appear quite solid when handled.

All chaetae bifid, 55-85 pm long, with upper

teeth longer than lower and nodulus slightly ectal

(Fig. 8A). Up to 8 chaetae per bundle anteriorly,

reduced in number posteriorly. Chaetae of sper¬

mathecal segment (X) present but not modified,

penial chaetae of XI 4 per bundle, straight, with

bifid ectal ends and no nodulus (Fig. 8B),

150-165 pm long, projecting somewhat dia¬

gonally towards the midline to open into median

copulatory chamber (Fig. 8C).

Testes anteroventral in X. Sperm funnels large,

vasa deferentia wide and slightly coiled near fun¬

nel. Point of union of atria and vasa deferentia not

established. Atria ovoid, thin walled, with short

ejaculatory ducts leading to pores on inner aspect

of folds in body wall lateral to a median ventral

copulatory chamber (Fig. 8C). Prostate tissue pre¬

sent as numerous discreet glands, which presum¬

ably connect with atria individually, although this

could not be confirmed. Spermathecae bulbous

with short ducts leading to dorso-lateral pores

anteriorly on X. Spenn in ampullae formed into

numerous bundles. Ovaries anteroventral in XI,

female pores not observed.

Remarks. Although the connections between the

atria and prostate tissue are obscured in the avail¬

able specimens, the numerous separate glands are

similar to those of other Breviatria. The long

straight penial chaetae with bifid tips also resem¬

ble those of other Breviatria. The bifid chaetae of

B. arvensis , with their long upper teeth, differ

from those of all other Breviatria but resemble

those of Biprostatus duplex , described below.

Biprostatus gen. nov.

Diagnosis. Atria small and spherical, bearing

broadly attached prostate tissue formed into

Figure 8, Breviatria arvensis sp. nov.: A, tip of ventral chaeta; B, tips of penial chaetae; C, male genitalia of holo

type. Scales: A-B, 0.01 mm; C, 0.06 mm.

two large glands, one anterior and one pos¬

terior. Atria leading to simple pores on inner

aspects of folds in body wall, lateral to a

median ventral copulatory chamber. Sperma-

thecal chaetae large and grooved, penial

chaetae multiple and bluntly bifid. Sperma-

thecal pores lateral, sperm in ampullae in loose

bundles. Coelomocytes abundant in anterior seg¬

ments in at least one species, to be confirmed for

the rest.

Type species: Telmatodrilus bifidus Brinkhurst

and Fulton, 1979.

Remarks. The type species of Biprostatus was

originally placed within the telmatodriline genus

Telmatodrilus. but is transferred to this new rhya-

codriline genus for reasons already discussed

above (see remarks for Rhyacodrilinae). This

genus is separable from most others in the Rhya¬

codrilinae by virtue of the small atria with large

bilobate prostate glands. Although some Hetero-

drilus, such as H. inermis and H. rapidensis, also

have short atria with large paired prostate glands,

these glands appear to be stalked and the chaetae

of Heterodrilus (1 or 2 per bundle, at least some

of which are trifid) are diagnostic (Erseus, 1981).

TUBIFIC1D OLIGOCHAETES FROM AUSTRALIA

Biprostatus bifidus (Brinkhurst and Fulton)

comb. nov.

Figure 9

Telmatodrilus ? (Telmatodrilus ?) bifidus Brinkhurst

and Fulton, 1979: 3, figs 15-19.

Holotype. Great Lake, Tas. (QVM: 14:328).

Material examined. Tas.: Holotype and paratypes,

Great Lake, 1975, 10 on slides (QVM:14:328-339);

Arthurs Lake, 4 Apr and 22 May 1977, 4 on slides

(ROB).

Distribution and habitat. Recorded only from the

above lakes of the central plateau of Tasmania.

Collected in both sandy and clayey sediments, at

depths up to 17.5m (Fulton, 1983b).

Diagnosis. Length up to 25 mm. Anterior bundles

with up to 13 bifid chactae with upper teeth

longer than the broad lower. (Fig. 9A) Number

of chaetae diminishing from V1I1-X with

upper teeth becoming shorter than the lower after

the clitellar region (Fig. 9B). Chaetae of some

species have groove-like markings (orna¬

mentations) ectally. Spermathecal chaetae

straight, single, with grooved tips (Fig. 9C),

varying in width and length, from shorter than to

3 times longer than somatic chaetae. Penial

chaetae bifid (Fig. 9D), up to 7 per bundle, twice

as long and thick as somatic ventral chaetae.

Atria small, elongate ovoid with short ejaculatory

ducts (Fig. 9E). Each atrium with a pair of

broadly attached prostate glands, one anterior,

which envelops the vas deferens, the other

posterior, occupying much of the coelom (Fig.

9E). Male pores and penial chaetae on the lateral

walls of a large median copulatory chamber.

Spermathecae ovoid, with short duct-like exten¬

sion entally and lateral pores. Sperm in ampullae

formed into loose bundles. Coelomocytes not

recorded.

Figure 9, Biprostatus bifidus (Brinkhurst and Fulton): A, tip of anterior chaeta; B, tips of posterior chaetae; C, tips

of spermathecal chaetae of Arthurs Lake specimen; D, tips of penial chaetae of holotype; E, genitalia of

QVM:14:338. Scales: A-B, 0.015 mm; C, 0.015 mm; D, 0.01 mm; E, 0.12 mm.

A. M. PINDER AND R. O. BRINKHURST

Biprostatus duplex sp. nov.

Figures 10, 11

Telmatodrilus sp. 2.—Pinder and Brinkhurst, 1994.

Material examined. Holotype: Que R.. just below Que

R. Mine dam. Tasmania, AMG 8014 914956, 27 Apr

1987, Wayne Fulton and Brett Mawbey (IFC), trans¬

verse serially sectioned specimen (with the posterior

whole-mounted) on 15 slides (QVM: 14:4207).

Paratypes: Collection details as for holotype, 5 seri¬

ally sectioned and 3 mounted whole or dissected on

slides, (QVM: 14:4208, 4210-4215, 4218): Que R. at

Murchison Highway, Tasmania (AMG 8014 901963).

27 Apr 1987, W. Fulton and B. Mawbey (IFC), 2 seri¬

ally sectioned and 2 whole-mounted on slides

(QVM: 14:4209, 4216. 4217, 4223).

Other material. Tas: Southwell R.. above llellyer

mine, AMG 8014 945960, 20 Dec 1988, S. Chilcott

(IFC), 1 on slide, 5 in alcohol (AMP): Douglas Creek,

AMG 8114 206692, 16-17 Feb 1990, S. Chilcott and L.

Mellefont (IFC), I on slide (AMP); Creek at Charlies

Hill, AMG 8111 297938, 19 Feb 1990. B. Mawbey

(IFC). 2 in alcohol (AMP).

Distribution and habitat. The above sites are all

rivers in western Tasmania.

Etymology: Named duplex for the double prostate

gland.

Description. Length up to 20 mm. Prostomium

short, bluntly conical. Coelomocytes large,

abundant in a few' preclitcllar segments.

Anterior chaetae 3-7 per bundle (Figs 10A, C),

fewer posteriorly, nodulus slightly distal, upper

teeth much longer than lower. Posterior chaetae

2-4 per bundle, nodulus clearly distal, upper teeth

also much longer than low'er. Chaetae shortest in

11 (75 pm) and posteriorly, otherwise about 110

pm. Spermathecal chaetae single in X and/or IX,

140-160 pm long and grooved ectally (Figs 10D,

E), associated with large glands (Fig. 10G), pro¬

truding from the body wall in line with somatic

ventral chaetae. Penial chaetae 4-7 per bundle,

straight with notched ectal ends (Fig. 10F), no

nodulus, 120-160 pm long. Penial chaetae lie

medial to atria and project somewhat diagonally

toward midline to open into median depression of

the body wall.

Sperm funnels large, vasa deferentia thin,

slightly coiled at first, then running posteriad to

enter the small globular atria subapically. Atria

with prostate tissue formed into large anterior and

posterior lobes (Fig. 1 IB). Short ejaculatory ducts

lead to pores on inner aspect of folds in body wall

lateral to a median copulatory chamber (Figs

11 A, B). Thick glandular tissue present ventrally

on either side of 10/11, more so in X (Figs 10G,

1 IB). Spermathecae oval (Fig. 10G), with short

thick-walled ducts and pores antero-laterally in X.

Spenn in ampullae formed into bundles.

Remarks. This species is similar to B. biftdus in

many respects. The genitalia, bifid or notched

penial chaetae and anterior somatic chaetae are

similar in the two species, although the latter are

less numerous in B. duplex. They differ in that the

upper teeth of the posterior somatic chaetae of B.

duplex are longer than the lower (like Breviatria

arvensis) whereas the upper teeth are shorter than

the lower on the posterior chaetae of B. biftdus.

The tips of the spermathecal chaetae also appears

to differ.

Bothrioneurum Stoic

Bothrioneurum Stoic, 1886: 647.—Michaelsen,

1900: 54.—Stephenson, 1930: 752.—Brinkhurst,

1971a: 539.

Type species. Bothrioneurum vejdovskyanum

Stoic. 1886.

Diagnosis. All chaetae bifid, hair chaetae absent.

Prostomium with a middorsal sensory pit. Vasa

deferentia short to long. Atria tubular and covered

with diffuse prostate gland cells except for ectal

portion, leading to protrusible pseudopenes bear¬

ing diverticula (paratria) with accessory glands.

Spenn stored within external spermatophores,

attached to body wall. Spennathecae absent.

Penial chaetae may be modified. Coelomocytes

large and abundant.

Bothrioneurum vejdovskyanum Stoic

Bothrioneurum vejdovskyanum Stoic, 1886: 647.—

Michaelsen, 1900: 54.—Brinkhurst, 1971a: 540, figs

8.29A-D.—Brinkhurst, 1982: 1.

Material examined. NSW: North Katoomba sewage

treatment plant constructed wetlands, 10 Aug 1994,

tentative identification of immature womis (AWT).

Other Australian records. NSW: Barrington R.,

Gloucester (Brinkhurst, 1982). Tas.: Arve R.

(Brinkhurst, 1982).

Distribution and habitat. Cosmopolitan

(Brinkhurst, 1971a) but uncommon in Australia.

Diagnosis. Length 28-35 mm. Anterior bundles

with 4-6 bifid chaetae with upper teeth longer

than lower, posteriorly fewer with teeth equally

long. Ventral bundles of XI with 4 slightly

hooked, club-headed penial chaetae arranged in a

fan. Vasa deferentia long, joining tubular atria

apically. Atria covered with prostate gland,

except for tenninal portion, and leading to pro¬

trusible pseudopenes which bear paratria with

accessory glands. Spennathecae absent. Body

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

Figure 10, Biprostatus duplex sp. nov.: A, ventral chaeta of II; B, ventral chaeta of other anterior segments; C, two

posterior chaetae; D, enlarged tip of spermathecal chaeta of QVM: 14:4214; E, whole spermathecal chaeta of

QVM:14:4214; F, tips of penial chaetae of QVM:14:4214; G, transverse section through segment X, composite

from several sections of holotype. Scales: A-C, E and F, 0.01 mm; G, 0.05 mm.

A. M. PINDER AND R. O. BRINKHURST

Figure 11, Biprostatus duplex sp. nov.: A, transverse section through segment XI, composite from several sections

of paratype QVM: 14:4223; B, reconstruction of genitalia from several specimens. Scales: A-B, 0.05 mm.

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

wall may appear spotted or papillate, but smooth.

Prostomium with dorsal sensory pit, often diffi¬

cult to detect but visible as a small indentation

when a slide mounted specimen viewed laterally.

Remarks. The only mature specimens examined

are from Barrington R. and these matched the

description of the species, except that the pro-

stomial pit was not observed. All other identifi¬

cations are of immature specimens based on

the presence of the prostomial pit and the form of

the somatic chaetac. An electron micrograph

of the prostomial pit is provided by Brinkhurst

and Gelder (1991).

Branchiura Beddard

Branchiura Beddard, 1892.—Michaelsen, 1900:

40.—Stephenson, 1930: 749.—Brinkhurst, 1971a: 562.

Type species. Branchiura sowerbyi Beddard,

1892.

Diagnosis. Hair chaetae present. Vasa deferentia

short, joining atria medially. Atria cylindrical,

covered in prostate cells. A glandular diverticula

(paratria) attached to atria where the latter join

large eversiblc pseudopenes. Coelomocytes not

conspicuous. Dorsal and ventral gill filaments

present posteriorly.

Branchiura sowerbyi Beddard

Branchiura sowerbyi Beddard, 1892: 325, pi. XIX,

figs 1-15.—Michaelsen, 1900:40.—Stephenson, 1930:

750.—Brinkhurst, 1971a: 563, figs 8.36D-F.—

Brinkhurst, 1971b: 114, fig. 2H.—Brinkhurst, 1982:

1.—Hogg and Norris, 1991:515.—Timms, 1981: 188.

Kawamuria japotiica Stephenson, 1917: 89, figs 1-5.

Holotype. Royal Botanical Gardens, London (Typus

amissus).

Material examined. NSW: Parramatta R., 17 Oct 1981

(AMS W197365); Wingecarribee R. at Greenstead, 14

Nov 1991, 2 in alcohol (AWT); Murray R. at Dora

Dora, 24 Mar 1994 (MDFRC); Megarritys Creek,

Femsworth Avenue, 3 Feb 1993, I in alcohol (AWT);

Rushy Billabong, 16 Jun 1981, 1 on slide (MDFRC);

Ryans Billabong, 27 Jul and 6 Dec 1982, 1 on slide, I

in alcohol (MDFRC). Qld: Leichhardt R., Coolullah

H.S., 115 km NW Cloncuiry, 25 Apr 1988. 1 on slide

(SAM E2814). Vic.: junction of Thomson and MacAl-

ister R.s, 31 Jun 1979, 1 in alcohol (NMV F77640);

Mitta Mitta R„ 1 Feb 1975, 2 Dec 1980 and 3 Mar

1982, 5 on slides, 3 in alcohol (NMV F77633-8). WA:

Herdsman Lake, 2 Nov 1989, I on slide (MUR); Wun-

gong Brook, Jan 1998, 1 two-tailed worm in alcohol

(AMP).

Other Australian records. ACT: Murrumbidgee R.

(Hogg and Norris, 1991). Qld: Brookfield, Goldcreek

(Brinkhurst, 1971a). Vic.: Thomson R. (Brinkhurst,

1982); Lake Purrumbete (Timms, 1981). WA: Lake

Claremont and Lake Monger, Perth (Brinkhurst,

1971b).

Distribution and habitat. Cosmopolitan

(Brinkhurst, 1971a), widespread in Australia,

occurring in a wide range of environments.

Diagnosis. Length 30-185 mm. Dorsal anterior

chaetal bundles with 1-4 (or more) short hair

chaetae and 5-12 (or more) chaetae that vary

from simple-pointed to bifid with short upper

teeth (which may be replicated). Posteriorly hairs

fewer and shorter and non-hair chaetae with less

replication of upper teeth. Ventral bundles with

6-11 bifid chaetae with upper teeth shorter than

lower, even simple-pointed anteriorly. Gills and

genital anatomy as for genus.

Other tubificid taxa

Timms (1978) records the tubificine Peloscolex

sp. (and ‘Peloscoiex’, undoubtedly a misprint)

from Lake St Clair, based on identifications by K.

V. Naidu. These refer to some form of papillate

tubificid, though presumably differing in chaetal

morphology from Breviatria papillatus. These

should not be considered definitive generic iden¬

tifications since body wall papillae are character¬

istic of, but not diagnostic for, Peloscolex Leidy,

1850, now revised as Spirosperma Eisen, 1879,

and other genera (Brinkhurst, 1979; Holmquist,

1978, 1979). We have seen several papillate

immature worms from Victoria (including speci¬

mens from the Plenty, Glenelg, Curdies and La

Trobc Rivers) and New South Wales (Georges

R.) that have chaetae very similar to some Spiros-

perma, but mature specimens are required for

identification.

The species coded Telmatodrilus sp. 1 in Pin-

der and Brinkhurst (1994) cannot yet be described

from the few poorly preserved specimens. This

species, which may be a Biprostatus, has anterior

chaetae bifid (like those of B. duplex ), posterior

chaetae simple pointed, and has modified penial

and spermathecal chaetae. Specimens have been

collected from Que and Southwell Rivers in

northwest Tasmania.

Numerous other undescribed tubificids, mostly

from Tasmania and Western Australia, have been

accumulated by the senior author since the prepa¬

ration of this manuscript and no doubt others

await discovery.

Zoogeography

Discussion of tubificid biogeography in the

region is limited by the low number of specimens

A. M. PINDER AND R. O. BRINKHURST

that have been examined front Australia, New

Zealand and Asia. No area of Australia has been

thoroughly surveyed, although projects are under¬

way to improve coverage of particular areas such

as the southwest of Western Australia, Tasmania

and South Australia. Nonetheless, some patterns

can be discerned.

Twenty-nine species of Tubificidae, in 12 tubi-

ficine and rhyacodriline genera, have been

recorded from Australian inland waters so far

(Table 1). Of these, 16 species are restricted to

Australia and two more are known only from

Australia and New Zealand.

The nonendemic component consists of 11

common species within widespread genera (i.e.

the tubificine genera Tubifex, Limnodrilus, Aulo-

drilus and Potamothrix and the rhyacodrilines

Rhyacodrilus, Branchiura and Bothrioneurum).

Several of these species also occur in New

Zealand (Brinkhurst and Fulton, 1979, 1982;

Marshall, 1975; Timms, 1982). Of the cosmopoli¬

tan species, only Aulodrilus pigueti, Aulodrilus

plurisela and Branchiura sowerbyi have been

recorded from northern Australia (the Queensland

records of Limnodrilus and Tubifex are from the

southeast of the state), but this may reflect the low

number of worms examined from this region. So

far, records of Aulodrilus pigueti have been

restricted to northern Australia, but other cos¬

mopolitan species are widespread in southern

Australia and seem to occur in a wide range of

habitats. The endemic species appear to have

more restricted distributions.

One of the three endemic genera, Antipodrilus,

is widespread, although most of the species are

not. Antipodrilus davidis, with the new range

extensions into Western Australia and South Aus¬

tralia, is the most widely distributed and is now

known from across southern Australia and New

Zealand. By contrast, A. mage/ensis appears to be

Table 1. Species list of Tubificidae known from Australia. * = known only from Australia, # = known

only from Australia and New Zealand, #? = New Zealand identification dubious, others cosmopolitan.

NSW includes Australian Capital Territory, n = north, sw = southwest.

Subfamily Genus Species

Qld

Australian state

WA(n) WA (sw) SA

NSW

Vic.

Tas.

Tubificinae Tubifex tubifex

•

Limnodrilus hoffmeisteri

•

Limnodrilus udekemianus

•

Limnodrilus clapareianus

•

Potamothrix bavaricus

•

Antipodrilus magelensis*

•

Antipodrilus davidistt

•

Antipodrilus timmsi*

•

Antipodrilus multiseta*

•

Antipodrilus plectilus*

•

Aulodrilus plurisela

•

Aulodrilus pigueti

•

Aulodrilus limnobius

•

Rhyacodrilinae Rhyacodrilus coccineus

•?

•

Rhyacodrilus megaprostatus*

•

Rhyacodrilus bijidusii

•

Ainudrilus billabungus*

•

Ainudrilus stagnalis*

•

Ainudrilus nhamus*

•

Ainudrilus fultoni*

•

Rhizodrilus arthingtonae*

•

Breviatrius multiprostatus #?

•

Breviatrius pectinatus*

•

Breviatrius papillalus*

•

Breviatrius arvensis*

•

Biprostatus biftdus*

•

Biprostatus duplex*

•

Bothrioneurum vejdovskyanum

•

Branchiura sowerbyi

•

TUBIFICID OLIGOCHAETES FROM AUSTRALIA

a species of the north and interior, A. timmsi is

known only from the south-east mainland while

A. plectilus and A. multiseta have not been

recorded outside a few sites in Tasmania. All

species show a preference for lentic habitats.

Both species of the new genus Biprostatus are

known only from Tasmania. The other new

genus, Breviatria , has four species, three of which

are apparently restricted to Tasmania. One of

these, Breviatria multiprostatus , may also occur

in New Zealand (Cowie et al. 1978; Marshall and

Winterbourn, 1979) but the identifications are

uncertain. The fourth, Breviatria pectinatus , is

known from a few lakes in Tasmania and

Victoria.

Other endemic rhyacodrilines belong to genera

with disjunct global distributions. Rhizodrilus

mostly consists of marine species with restricted

distributions, often limited to single records from

the type localities (Baker and Brinkhurst, 1981;

Erseus, 1990a). The only Australian species, Rhi¬

zodrilus arthingtonae, is known only from one

lake on North Stradbroke Island, Queensland

(Jamieson, 1978). Ainudrilus mostly consists of

coastal Pacific or Caribbean marine species

(Erseus, 1990ab; 1997), except for the four fresh¬

water Australian species. Of these, A. stagnalis

and A. billabongus have been recorded only from

the Northern Territory, A. nharna only from

southwest Western Australia and A. fultoni only

from Tasmania.

Of the 18 species restricted to Australia and

New Zealand, ten occur in Tasmania and eight are

known only from that state (including most

species of the new rhyacodrilinc genera). Simi¬

larly, about half of the 26 species of Australian

Phreodrilidae are restricted to Tasmania (Pinder

and Brinkhurst, 1997). By contrast, only three

tubifields ( Rhvacodrilus megaprostatus , Rhyaco-

drilus bifidus and Antipodrilus timmsi ) and one

phreodrilid ( Phreodriloides notabilis Benham,

1907) are known only from southeast mainland

Australia. This is despite the examination of

worms from twice as many localities (and

locality-dates) from Victoria and New South-

Wales as from Tasmania.

Of the tubificids known only from Tasmania,

three (A. plectilus. A. multiseta and B. bifidus) are

known only from Great Lake and Arthurs Lake on

the eastern edge of the central plateau, and one ( B.

papillatus) is known only from these lakes and

nearby Lake Sorell. While not wishing to place

too much emphasis on this, because oligochaete

records from other lakes are limited, it is interest¬

ing to note that two phreodrilids are also known

only from these lakes (Pinder and Brinkhurst,

1997). In addition, numerous other invertebrates

and fish are apparently restricted to these lakes,

though mostly to Great Lake (Frankenberg, 1974;

Fulton, 1983ab; Invertebrate Advisory Commit¬

tee, 1994; Timms, 1985; Williams, 1974). These

lakes pre-date the late pleistocene glaciations that

led to the creation of most other lakes in Tas¬

mania and lie outside earlier glacial influence

(Davies, 1974). Sutherland et al. (1973) and

Sutherland (1980) suggest that lacustrine environ¬

ments may have existed in the Great Lake basin

since the late Oligocene when basalt flows

blocked the upper Ouse R.. The age of Great Lake

has been seen as a factor contributing to local

endemicity in its fauna (Timms, 1985). Other

ancient lakes, such as Lake Baikal (Russia), Lake

Ohrid (Europe) and Lake Tanganyikae (Africa)

also have numerous endemic oligochaetes,

whereas this is rare in more recent lakes (Martin,

1996). The few Tasmanian lakes of recent (Pleis¬

tocene/Holocene) origin from which oligochaetes

have been identified appear to have only species

that are widespread in the state. These include

pre- and post-impoundment Lake Pedder as

reported by Lake (1998), a few glacial lakes

(Dobson, Dove and St. Clair) surveyed by Timms

(1978) and a dune barrage lake in the north-east

(Blackman’s Lagoon, sampled by the first

author). However, records from many other Tas¬

manian lakes are required to confirm the apparent

endemism of the oligochaeta fauna of Great Lake,

Arthurs Lake and Lake Sorell.

Insufficient localities have been examined from

other parts of Australia to generalise about

regional diversity. The Phreodrilidae appear to be

quite diverse in the southwest of Western Aus¬

tralia (Pinder and Brinkhurst, 1997 and unpub¬

lished data) but only one endemic tubificid is

known from there so far. Three tubificids (Ain¬

udrilus stagnalis, Ainudrilus billabongus and

Antipodrilus magelensis) are known only front

inland and northern Australia so far, but so few

localities have been examined in these regions

that it is difficult to speculate what further work

will reveal.

The Australian Tubificidae seem to have a dif¬

ferent biogeographic history to the apparently

Gondwanan Phreodrilidae (Pinder and Brink-

hurst. 1997). Antipodrilus appears most similar to

genera that are widespread in the northern hemi¬

sphere but absent in Africa and South America

(Brinkhurst, 1991). The new rhyacodriline gen¬

era, Breviatria and Biprostatus, are of uncertain

affinity as the phylogenetic relationships are

unclear in this possibly paraphyletic subfamily.

Other endemic rhyacodrilines are members of a

A. M. PINDER AND R. O. BRINKHURST

cosmopolitan freshwater genus (Rhyacodrilus

spp.) or appear to be allied to marine forms

(Ainudrilus spp.) or are only tentatively assigned

to a widespread genus whose monophyly is

uncertain (Rhizodrilus arthingtonae). As almost

nothing is known of the oligochaete fauna of

southeast Asia and New Zealand, little comment

can be made about affinities within the

Australasian region.

Acknowledgements

We express our gratitude to all those who have

provided specimens for this study. The work has

been funded mostly by grants from the Australian

Biological Resources Study. Richard Marchant of

Museum Victoria made available additional fund¬

ing and has been extremely generous with all

aspects of support needed to complete this study.

Histological work was funded by the Queen Vic¬

toria Museum and Art Gallery’s Plomley Founda¬

tion and serial sections were prepared by Gordon

Thomson of Murdoch University. Dr M. Fcrraguti

provided access to an unpublished manuscript.

Brenton Knott and Peter Davies kindly shared

their knowledge of Tasmanian lakes. Andrew

Boulton, Diana Jones and Stuart Halse have

allowed the first author to use their laboratories

and facilities during the last few years.

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Memoirs of Museum Victoria 58(1): 77-88 (2000)

NEW AUSTRALIAN SPECIES OF OECETIS ALLIED TO O. COMPLEXA

KIMMINS (TRICHOPTERA: LEPTOCERIDAE)

Alice Wells

Australian Biological Resources Study, Environment Australia, Canberra, PO Box 787, ACT 2601, Australia

(alice.wells@ea.gov.au)

Abstract

Well, A., 2000. New Australian species of Oecelis allied to O. complexa Kimmins

(Trichoptera; Leptoceridae). Memoirs of Museum Victoria 58(1): 77-88.

Seven new species of long-horned caddisfly together with Oecelis complexa Kimmins form

a discrete set in the Australian Oecelis fauna, here called the complexa- group, based on wings

with a long footstalk on Fork I, male inferior appendages four-lobed, and phallus with paired

spiny parameres. Characteristics of this group appear to conflict with the placement of Oecetis

complexa in the most recent subgeneric and species group classification of world Oecetis.

Introduction

Long-homed caddisflies of the cosmopolitan

genus, Oecetis McLachlan, 1877 (Trichoptera:

Leptoceridae: Leptocerinae), are diverse and

often abundant in lotic and lentic waters through¬

out Australia. Neboiss (1986) listed 19 Australian

species in the Atlas of Trichoptera of the SIV

Pacific — Australian Region and, in another

work, described a further six (Neboiss, 1989).

These last species, with several species from New

Guinea and Indonesia, he placed in a * reticu-

lata- group’, distinguished by males having a

dorsal ‘shield’ (usually with reticulate cuticular

sculpture) overhanging the terminal abdominal

segments. Many additional undescribed Aus¬

tralian species have been represented in

museum collections for several years. The pre¬

sent work, the first part in a review of the

genus in Australia, deals with Oecetis

complexa Kimmins and seven new closely

allied species, here called the complexa-

group. The remaining 18 described species

and around 40 new species are to be reviewed

in subsequent works.

Treatment of the Australian Oecetis fauna in

isolation from the world fauna is insular. A clas¬

sification of world Oecetis in an unpublished

thesis by Chen (1992), based on a phylogenetic

analysis, divided the genus into four subgencra,

all further divided into species groups. According

to Chen, three of the four subgenera arc repre¬

sented in Australia, each by one species group.

Neboiss’ (1989) 'reticulata'- group species share

the derived features of the larger worldwide

group to which Chen assigned them, as do also a

large group of Australian species, among them

the very common and widespread O. pechana

Mosely, which has wings with hair short and Fork

1 sessile, and in the male, forewing bearing

patches of androconia (scales), and phallus with a

single internal spiny paramere. Chen’s assign¬

ment of the remaining Australian species to one

subgenus is questionable. Oecetis complexa,

which has males with paired spiny parameres, he

grouped on the basis of putative sharing of the

derived condition in which the male paramere

spines are absent (and, plesiomorphically, the

phallus symmetrical), with 12 other Australian

species (all lacking parameres) and an assortment

of Oriental, Palaearctic and Neotropical species.

Within this subgenus, O. complexa was assigned

to a species group sharing the feature RPu 2

divided at about one-tenth the length of the

forewing (i.e.. Fork I very short), and within the

group, clustered with O. parka Mosely and

O. inscripta Kimmins, on the basis of sharing the

derived feature of ‘inferior appendages with

basodorsal arms prominent.’ Since the basis of

subgcneric placement is rejected here, for the pre¬

sent a new complexa- group is recognised in the

Australian fauna, defined by wings with short

hair on veins, the forewing (Fig. 25) with Fork 1

short, its the footstalk about as long as the fork; in

males, the inferior appendages comprising four

lobes, and the phallus with paired spiny, often

complex and asymmetrical, parameres (Figs

26-33).

Material and methods. Specimens were prepared

for study following methods used by Wells

(1990) for micro-caddis flies (Flydroptilidae).

ALICE WELLS

Although females have been sorted into vials

with males of some species, many are so similar

that until verified by breeding or rearing, the

associations are considered only tentative. Thus,

females are not described here.

Depositories are abbreviated as follows: AN1C,

the Australian National Insect Collection, Can¬

berra, Australian Capital Territory; BMNH, the

Natural History Museum, London, England;

OSS, Office of the Supervising Scientist (now

ER1SS, Environmental Research Institute of the

Supervising Scientist), Northern Territory; NMV,

Museum Victoria, Melbourne, Victoria; NTM,

Northern Territory Museum and Art Galleries,

Darwin, Northern Territory; and QM, Queens¬

land Museum, Brisbane, Queensland. ‘ARR’

and ‘ARRS’ are used for Alligator Rivers

Region, Northern Territory and Alligator Rivers

Region Survey (conducted by the OSS), respec¬

tively. ‘WTH’ numbers are Wet Tropics

Heritage numbers from the study by Walker et al.

(1993).

Key to males of the Oecetis complexa -group

1. Preanal appendages fused with tergum IX (e.g. Figs 7, 8).2

Preanal appendages free (Figs 17, 20, 23).6

2. Tergum X produced distally to form paired horn-like sclerotised spines

twisted ventrally around the phallus (Figs 14,15). O. uptoni sp. nov.

Tergum X not greatly produced...3

3. Tergum X, in dorsal view, shallowly concave apically; in ventral view,

mesal margins of inferior appendages obliquely slanted to form deep V

(Fig. 3). O. complexa

Tergum X with a more or less V-shaped excision (Figs 5,8), although apices

of lobes may be convergent.4

4. Apices of distal lobes of tergum X convergent in dorsal view (Fig. 8); in ven¬

tral view, basodorsal lobe of inferior appendages distally obliquely angled to

shallow U-shaped mesal concavity (Fig. 9)... O. obliqua sp. nov.

Apices of distal lobes of tergum X divergent in dorsal view (Figs 5, 11); in

ventral view, basodorsal lobes of inferior appendages separated by a deep

U-shaped concavity (Figs 6, 12)...5

5. Parameres terminating in a pair of equal length darkly sclerotised spines

(Figs 10-12, 28); lobes formed by bisection of tergum X obliquely truncate

apically (Fig. 11)..... O. adelaidica sp. nov.

Parameres with the second distal spine subapical (Figs 4, 27); lobes formed

by bisection of tergum X tapered to apices (Fig. 5)..0. paracomplexa sp. nov.

6. Inferior appendages in ventral view with basoventral lobe forming a pair of

parallel ‘pillars’ (Fig. 24); in lateral view, main body and basodorsal lobe

about equal length and thickness (Fig. 22)..... O. parallela sp. nov.

Inferior appendages in ventral view with basoventral lobe not pillar-like

(Figs 18, 21); in lateral view, main body far broader than the finger-like

basodorsal lobe (Figs 16, 19).7

7. In lateral view, main body of inferior appendages about equal width through¬

out length, truncate apically, somewhat clasper-like in ventral view (Figs 16,

18). ,.0. blythi sp. nov,

In lateral view, main body of inferior appendages broad and irregular in

shape, forming stout lobes in ventral view (Figs 19,21).. ..O. glebula sp. nov.

Oecetis complexa Kimmins

Figures 1-3, 26, 34

Oecetis complexa Kimmins in Mosely and Kimmins,

1953: 284.

Material examined. Queensland: holotype male, Mur-

willumbah. New South Wales (BMNH): 2 males, 4

females. Camp Mtn, 31 Mar 1967, N. Dobrotworsky

(NMV); male, 6 females, Bullimba Creek, nr Brisbane,

Site R, Kinmax St, riffle, 23 Oct 1971 (NMV); male,

Girraween Natl Pk, nr Wyberba, 10 Oct 1973, A.

Neboiss (NMV); male, female. Middle Claudie River,

Iron Ra., 2-9 Oct 1974, M.S. Moulds (NMV); male.

Iron Range. 16 Oct 1974, M.S. Moulds (NMV); male,

female, 16 km W of Ravenshoe, 2 Jan 1975, M.S.

Moulds (NMV WTH-1360); male, Gordon Creek, Iron

Ra.,16 Oct 1975, M.S. Moulds (NMV); 2 males, 4

females, Alice River, Hervey Range River, 25 km W of

Townsville, 9 May 1979, A. Wells (NMV WTH-1361);

male. Jamboree Heights, Brisbane, 6 Oct 1979, G.

Daniels (NMV); 2 males, Mothar Mtn, 12 km SE of

OECETIS COMPLEXA AND ALLIED SPECIES

Figures 1-9, Oecetis species, male genitalia, lateral, dorsal and ventral views: 1-3, O. complexa Kimmins; 4-6, O.

paracomple.xa sp. nov.; 7-9, O. obliqua sp. nov.

Gympie, 29 Oct 1980. A. Neboiss (NMV); male,

female, Tinaroo Dam, Nov 1982, T. Hinger (NMV):

male, female, Booloumba Creek, 8 km SW of Kenil¬

worth, 26°39'S 152°39'E, 12 Dec 1984, G. The-

ischinger (NMV): male, female, Booloumba Creek, 8

km SW of Kenilworth, 26°39’S 152°39’E, 12 Jan 1986,

G. Theischinger (NMV); male, female. Emu Creek,

State Forest nr Bcnarken, 26°53’S 152°08’E, 15 Jan

1986, G. Theischinger (NMV); 2 males. Crows Nest

Falls, N of Toowoomba, 27°14'S 152°07'E, 18 Jan

1986, G. Theischinger (NMV); 3 males, 12°44’S

145°16'E, Claudie River, Iron Range Natl Pk, 25 km

NW Lockhart River, 10 Nov 1988, K. Walker (NMV);

males, females, 17°08’S 145°44'E, Mulgrave River, 8

km NW Gordonvale, 15 Nov 1988, K. Walker (NMV

WTI1 1359). New South Wales: male, Blandford. 8

Oct 1976, M.S, Moulds (NMV): male, Clarence River,

at Yates Crossing, 26 Oct 1981, Wells and Carter

(NMV); male, 3 females, Swan Crossing nrComboyne,

13 Feb 1999, G. Theischinger (ANIC). Victoria: male,

Morrison. Moorabool River, 25 Mar 1953, A. Neboiss

(NMV); 3 males, Greendale, 6 Jan 1956, Neboiss

(NMV); male, Warrandyte, Yarra River, 1 Feb 1959,

Neboiss (NMV); male, Delatite, 10 Dec 1962, Neboiss

(NMV); male, female, Thurra River, Cape Everard, 22

Mar 1970. A. Neboiss (NMV): male, Tyers River, Site

22, 24 Feb 1974 (L.R.E.S.) (NMV); males, females,

Otway Ra., East Branch Barwon River outflow from

ALICE WELLS

Figures 10-18, Oecelis species, male genitalia, lateral, dorsal and ventral views: 10-12, O. adelaidica sp. nov.;

13-15, O. uptoni sp. nov.; 16-18, O. blythi sp. nov.

OECETIS COMPLEXA AND ALLIED SPECIES

Figures 19-24, Oecelis species, male genitalia, lateral, dorsal and ventral views: 19-21, O. glebula sp. nov.; 22-24,

O parallela sp. nov.

Lake Elizabeth, nr forest, 1 Apr 1975, J. Aldenhoven

(NMV); males, females, Gellibrand River, E of Gelli-

brand, 26 Jan 1982, A. Neboiss and R. StClair (NMV);

males, females, Gellibrand River, at Lower Gellibrand,

22 Feb 1982, K. Walker (NMV). South Australia: 2

males, 2 females. Spring Creek, Wilmington, 22 Oct

1975, A. Wells (NMV). Western Australia: male.

King Edward River, 14°04’S 126°I2’E, 3 Sep 1996, I.

Edwards (NMV).

Diagnosis. The male genitalia of O. complexa

share with O. paracomplexa sp. nov., O. obliqua

sp. nov., O. uptoni sp. nov. and O. adelaidica sp.

nov., the derived states of preanal appendages

fused with tergum X, and slender elongate lateral

lobes on the inferior appendages. (). complexa is

distinguished from these other species by other

features of the male genitalia: the inferior

appendages are separated ventrally by a deep, V-

shaped cleft (Fig. 3) in contrast to the U-shaped

separation of the other species and the basodorsal

lobe is broad when viewed laterally (Fig. 1); on

tergum X, the proximal part is somewhat rectan¬

gular, the distal lobes separated by a broad con¬

cavity (Fig. 2) and extended distally to form a pair

of spiny structures; on the phallus, the parameres

have a laterobasal spine and, more distally, a

small black dorsal spine proximal to the darkly

sclerotised acute apical spine.

Distribution (Fig. 34). Widespread in eastern

Australia, with an anomalous record from each of

South Australia and northern Western Australia.

Remarks. New figures of the male genitalia (Figs

1-3, 26) are supplied to aid comparisons.

ALICE WELLS

Fig. 25, Oecetis parallela sp. nov., fore- and hind wings.

Oecetis paracomplexa sp. nov.

Figures 4-6, 27, 35

Material examined. Holotype. Male, North Queensland,

Middle Claudie River, 29 Jun 1982, Schneider and

Daniels (NMV T-17393).

Paratypes. Queensland: male, same data as for holo¬

type (NMV); male, Lockerbie area. Cape York, 13-27

Apr 1973, S.R. Monteith (AN1C); male, Dulhunty

River, at Telegraph Crossing, 10 Feb 1992, D.

Cartwright and A. Wells (QM).

Other material examined. Western Australia: male,

22°23’S 117°56’E GPS, 37 km NNE of Tom Price. 6

Oct 1995, D.C.F. Rentz and P.J.M. Greenslade (ANIC).

Diagnosis. O. paracomplexa sp. nov. shares with

O. complexa, O. obliqua, O. itptoni and O. ade-

laidica , the derived states of preanal appendages

fused with tergum X, and slender elongate lateral

lobes on the inferior appendages. In general geni-

talic form it most closely resembles O. complexa ,

but differs in that the inferior appendages have a

U-shaped basal separation ventrally, the baso-

dorsal lobe narrowly digitiform in lateral view,

and tergum X in dorsal view deeply divided to

form 2 divergent triangular lobes.

Description. Genitalia, Figs 4—6, 27. Segment IX

narrow middorsally and ventrally, broadly

rounded laterally; preanal appendages slightly-

tapered apically, fused to tergum X. Tergum X

wide at base, bifid almost throughout length,

apices of lobes distally divergent. Inferior

appendages in ventral view with main body

slender, basoventral lobe short and obliquely

truncate, in lateral view basodorsal lobe and a

small median lobe digitiform. Phallus (Fig. 27)

short, smoothly curved downwards, relatively

simple with the fused parameres and phallobase

forming a short, broad, sclerotised cover, distally

divided into 2 subequal spines.

Etymology>. Named for its resemblance to O.

complexa', Latin— para — like.

Distribution (Fig. 35). Far northern Queensland

and the Hamersley Range, Western Australia.

Oecetis obliqua sp. nov.

Figures 7-9, 28, 36

Material examined. Holotype. Male, Archer Creek, nr

Midstream Falls, 5 Apr 1997, G. Theischinger and

F. Mueller (ANIC).

Paratypes, Queensland: male, same data as for holo¬

type; male. Iron Range, Cape York Peninsula, 2 -9 Jun

1971, E.F. Riek (ANIC); 2 males. Gregory River cross¬

ing nr Goodwood, 24.viii.1977, K.L. Lambkin (NMV

WTH-1393); 2 males. State Forest, 24 km along Golds-

borough Rd, nr Gordonvale, 27 Dec 1980, M.S. and

B.J. Moulds (NMV WTH-1391); male. Upper Annan

River nr Shipton's Flat, S of Cooktown, 1 Jan 1981,

M.S. and B.J. Moulds (NMV WTH-1392); 2 males,

Jourama Falls, 19 Mar 1997, G. Theischinger and F.

Mueller (ANIC).

Other material examined. Queensland: male, Iron

Range, Cape York Peninsula, 2-9 Jun 1971, E.F. Riek

(ANIC); male, 15°4US 145°12’E, Annan R, 3 km

WbyS Black Mountain, 27 Sep 1980, J.C. Cardale

(ANIC); 2 males, female, I5°14’S 145°07’E, 7 km N

Hope Vale Mission, 4 Oct 1980, J.C. Cardale (ANIC);

3 males, 15°16*S 144°59’E, 14 km WbyN Hope Vale

Mission, 8-10 Oct 1980, J.C. Cardale (ANIC); 3 males,

3 females, 15°47’S 145°17’E, Moses Creek, 4 km

NbyE Mt Finnigan, 14 Oct 1980, J.C. Cardale (ANIC);

8 females. State Forest. 24 km along Goldsborough Rd,

nr Gordonvale, 27 Dec 1980. M.S. and B.J. Moulds

(NMV WTH-1391).

Diagnosis. Oecetis obliqua shares with (). com¬

plexa. O. paracomplexa. O. adelaidica and O.

uptoni, the derived states of preanal appendages

fused with tergum X. and slender elongate lateral

lobes on the inferior appendages, but is distin¬

guished by the short, spiny convergent distal

lobes on segment X and the broadly oblique dis¬

tal margin of the basoventral lobes of the inferior

appendages in ventral view,

Description. Genitalia. Figs 7-9, 28. Segment IX

broadest ventrolaterally; preanal appendages

fused with tergum X. Tergum X bifid, the short

acute distal lobes being separated basally by a V-

shaped cleft, their apices converging. Inferior

appendages in ventral view with basoventral

lobes broad-based and obliquely truncate

OECETIS COMPLEXA AND ALLIED SPECIES

Figures 26-33, Oecetis species, phallus: 26, O. complexa Kimmins; 27, O. paracomplexa sp. nov.; 28, O. obliqua

sp. nov.; 29, O. adelaidica sp. nov.; 30, O. uptoni sp. nov.; 31, O. blythi sp. nov.; 32, O. glebula sp. nov.; 33,

O. parallela sp. nov.

ALICE WELLS

apicomesally, in lateral view main body narrow,

about same width as dorsobasal lobe, but longer.

Phallus (Fig. 28) with parameres forming a pair of

stout, sclerotised lateral spines, the base of the

phallus is irregular, making interpretation diffi¬

cult, but there appears to be no clear phallobase

such as occurs in O. glebula sp. nov. or O. blythi

sp. nov.

Etymology. Descriptive of the obliquely angled

apicomesal margin of the inferior appendages;

Latin — obliquus — slanting.

Distribution (Fig. 38). Northern Queensland.

Oecetis adelaidica sp. nov.

Figures 10-12, 29, 37

Material examined. Holotype. Male, Northern Terri¬

tory. Adelaide River. 15 km E of Stuart Highway, 15

Aug 1979, J. Blyth (NMV T—17394).

Paratype: Northern Territory: male, same data as

for holotype.

Diagnosis. Oecetis adelaidica shares with O.

complexa, O. paracomplexa, O. obliqua and O.

uptoni, the derived states of preanal appendages

fused with tergum X, and slender elongate lateral

lobes on the inferior appendages. It is unique in

having the parameres on the phallus terminating

in a pair of more or less equal length darkly scle¬

rotised spines.

Description. Genitalia, Figs 10-12. 29. Abdomi¬

nal segment IX subquadrate in ventral view, the

preanal appendages fused with tergum X. Tergum

X deeply cleft, forming 2 slightly divergent,

almost pillar-shaped lobes, their apices obliquely

truncate. Inferior appendages in ventral view

divided by a deep U-shaped concavity, the

basoventral lobe apically truncate, the main body

slender, curved; a small setatc digitiform inner

lobe visible in dorsal view probably represents the

basodorsal lobe. Phallus (Fig. 29) relatively nar¬

row, cursed, fused parameres and phallobase

forming a dorsal sheath, on each side divided api¬

cally into a pair of equal sclerotised spines,

basally a small lateral spur, its tip sclerotised.

Etymologv. Named for the Adelaide River, the

type locality.

Distribution (Fig. 37). Adelaide River, Northern

Territory.

Oecetis uptoni sp. nov.

Figures 13-15, 30, 38

Material examined. Holotype. Male, Northern Terri¬

tory, 12°42’S I30°58’E, Berry Springs, 31 Mar 1992,

A. Wells and J. Webber (NTM).

Paratypes. Northern Territory: 2 males, same data

as for holotype (NTM); male, ARR Magela Creek at

Rum pipeline, 12 Mar 1991, P. Dostine (NTM); male,

ARR Magela Creek at Rum pipeline, 18/19 Mar 1991,

P. Dostine (NTM); 2 males, 12°27’S 131°03’E,

Howard Springs, 25 Mar 1991, de Jong, van Achterberg

and Wells (NMV); 2 males. Holmes Jungle, 7 Apr

1991. Ilorak, Upton and Wells (ANIC): 2 males,

Baroalba Springs, 12°48’S 132°49 - E, 16 Jan 1992,

Wells, Webber and Bickle (NTM); male, same locality,

4 Mar 1992, Wells (NTM); 2 males, Berry Springs,

12°42’S 130°58’E, 31 Mar 1992. Wells and Webber

(NTM); male, I2°42 , S 132°57’E, Kakadu Natl Pk,

Magela Creek, OSS Site/009, 24 Apr 1992, A. Wells

and J. Webber (NTM).

Other material examined. Northern Territory:

female, same data as for holotype (NTM); female.

Holmes Jungle, 7 Apr 1991. Horak, Upton and Wells

(ANIC); female, 12°42’S I32°57’E, Kakadu Natl Pk,

Magela Creek, OSS Site/009, 15 Aug 1991. A, Wells

and J. Webber (NTM); female, Berry Springs. I2°42’S

130°58’E, 31 Mar 1992. Wells and Webber (NTM); 2

males. N2 ARR. Magela Creek at Rum Pipe, 30/31 Mar

1992. P. Dostine (NTM); 4 males, same loc. and col¬

lector, 6/7 Apr 1992 (NTM); male. Magela Creek,

23/24 Mar 1992, P. Dostine (NTM).

Diagnosis. Oecetis uptoni shares with O. com¬

plexa, O. paracomplexa. O. obliqua and O.

adelaidica, the derived states of preanal

appendages fused with tergum X, and inferior

appendages with lateral lobes slender and elon¬

gate. It most closely resembles O. complexa in

having tergum X modified distally to form spines,

but the spines are more elaborate and curve under

the phallus. Among all complexa- group species, it

can be recognised immediately by the sharply

down-turned paratnere spines on the phallus.

Description. Genitalia, Figs 13-15, 30. Segment

IX more uniform in length than other complexa-

group species; preanal appendages small, club-

shaped, free. Tergum X broad-based, deeply

divided in distal two-thirds, forming spines that

curl under the phallus. Inferior appendages broad-

based, basoventral lobes almost rounded

anteromesally, main body forming slender lobes

distally, with a short dorsobasal lobe visible in

lateral view. Phallus (Fig. 30) membranous and

curving ventrally, parameres short sclerotised,

each in form of a sharp down-turned spur with an

acute dorsal spine.

Etymology. Named for Murray Upton, in appreci¬

ation of his support during my several years in the

Northern Territory.

Distribution (Fig. 38). Northern Northern

Territory.

Remarks. Oecetis uptoni was not among the

numerous species of Oecetis collected in a

OECET1S COMPLEXA AND ALLIED SPECIES

1988-1989 survey of the Trichoptera of the Alli¬

gator Rivers Region (Wells, 1991), and despite

much subsequent collecting in the ARR, espe¬

cially on Magela Creek, only very few specimens

have been taken. The real or apparent rarity of this

species illustrates a curious phenomenon noticed

during regular sampling at a Magela Creek site

over the years 1991-1993. Several species were

taken once or twice only over 14 months of stan¬

dardised weekly light trap-sampling, while other

species were collected regularly almost through¬

out the period. All specimens of O. uptoni were

taken in March and April, generally the end of the

wet season in this seasonal monsoon area. This

species itself may be strongly seasonal. Alterna¬

tively, our sampling successes may reflect recruit¬

ment from upstream (?escarpment) populations,

and failure to establish as the water flow drops

and the stream dries to intermittent pools at the

beginning of the dry season.

Oecetis blythi sp. nov.

Figures 16-18, 31, 39

Material examined. Holotype. Male, New South Wales,

Severn River, 29°28’S I5I°29’E, 23 Oct 1981, Wells

and Carter (NMV T 17395).

Paratypes. New South Wales: male, same data as

holotype (NMV): male, Clarence River at Yates Cross¬

ing, 26 Oct 1981, Wells and Carter (NMV). Queens¬

land: 4 males, Tinaroo Dam (nr Kairi Creek). 22 Jun

1971, E.F. Riek (ANIC); 6 males. Upper Ross River,

below weir. SW Townsville, 8 May 1979, A. Wells

(NMV); male. Emu Creek State Forest nr Benarkin,

26°53’S 152°08’E, 15 Jan 1986, G. Theisehinger

(NMV); male. Crows Nest Falls, N Toowoomba,

27°I4’S 152°07’E, 18.1.1986,0. Theisehinger (NMV).

Other material examined. Queensland: female, same

data as holotype (NMV); male, 36 km S of Miriam

Vale, 25 May 1971, E.F. Riek (ANIC); 3 females, Tina¬

roo Dam (nr Kairi Creek), 22 Jun 1971, E.F. Riek

(ANIC); male. South Pine River, 8 km W of Samford,

21 Oct 1980, A. Neboiss (NMV); 10 females, Clarence

River at Yates Crossing, 26 Oct 1981, Wells and Carter

(NMV); male, Blackdown Tableland Natl Pk, Nov

1982, T. Hinger (NMV); 3 females, Emu Creek State

Forest nr Benarkin. 26°53'S 152°08 , E, 15 Jan 1986, G.

Theisehinger (NMV); male, 3 females, Emu Creek,

Suite Forest nr Benarkin, 26°53’S 152°08'E, 15 Jan

1988, G. Theisehinger (NMV); male, 3 females, Tina¬

roo Pines Caravan Park, 10 Jun 1972, N. McFarland

(NMV); 4 males. 13 females, Alice River, Hervey

Range Road, 25 km W Townsville, 8 May 1979, A.

Wells (NMV); males, females, Obi Obi Creek, 8 km

SW Mapleton, 23 Oct 1980, A. Neboiss (NMV). Vic¬

toria: 3 males. Swan Lake. 30 km NW of Portland, 27

Feb 1976, P.A. Meyer (NMV); male, Wyperfeld Natl

Pk, Lake Werribean, 6 Apr 1977, J. Blyth (NMV); 3

males, 4 females. Lake Albacutya, 16 Jan 1980, J. Blyth

(NMV). South Australia: male, female. North

Adelaide, 28 Nov 1975, M. Davies (NMV).

Diagnosis. In common with O. glebula and O.

parallela, O. blythi has the preanal appendages

free and lateral lobes of the inferior appendages

stout. The latter structures are somewhat interme¬

diate in size between the form of the other two

species, and the ventral lobes of the inferior

appendages are subtriangular in ventral view,

rather than rounded as in O. glebula or subrectan-

gular as in O. parallela Oecetis blythi can be

distinguished readily in ventral view by the pair

of slender processes representing the basodorsal

lobe, on the inner side of the inferior appendages,

lying almost parallel to the phallus.

Description. Genitalia, Figs 16-18, 31. Segment

IX short middorsally, relatively long laterally and

ventrally; preanal appendages elongate. Tergum

X reduced to a short rounded lobe. Inferior

appendages short and rounded mesoventrally;

main body stout, short, the inner apical angle

spur-like; in lateral view basodorsal lobe setate,

digitiform, in ventral view, lying alongside the

phallus. Phallus (Fig. 31) elongate with parameres

forming a complicated set of spines, including 1

dorsal pair which almost equal in length and are

membranous and free for most of the length of the

phallus; basally a slender curved spine; dorsally a

bract-like phallobase.

Etymology. Named for John Blyth whose collect¬

ing efforts added so much interesting material to

the NMV collection.

Distribution (Fig. 39). Disjunct: mainly

Townsville area of north-eastern Queensland,

through north-eastern New South Wales to west¬

ern Victoria; a single record from North Adelaide,

South Australia (probably from the Torrens

River).

Oecetis glebula sp. nov.

Figures 19-21, 32, 40

Material examined. Holotype. Male, Western Australia,

Ellendale, Greenough River, E of Geraldton, 11 Sep

1974, K. Carnaby (NMV T 17396).

Paratypes. Western Australia, 2 males, same data as

holotype (NMV); 3 males, Milistream Crossing Pool,

21 Oct 1970, J.C. Cardale (ANIC); 5 males, 10 females,

Lockyer Gorge, Harding River, Pilbara, 19 Oct 1979,

J. Blyth (NMV); male, female, Fortescue Falls,

Hamersley Ra. Natl Pk, 27 Oct 1979, J. Blyth (NMV).

Other material examined. Western Australia: 2

females, Milistream Crossing Pool, 21 Oct 1970, J.C.

Cardale (ANIC); male, N of Carnarvon, De Grey

Station Road, 29 Jun 1972, N. McFarland (NMV);

ALICE WELLS

male, Wittenoom Gorge, Hamersley Ra., 20 Feb 1977,

M.S. and B.J. Moulds (NMV); male, Hamersley Ra., 20

Feb 1977, M.S. and B.J. Moulds (NMV); 6 males, 13

females. Midstream, Fortescue River, S of Roeboume,

17 Nov 1978, M.S. and B.J. Moulds (NMV); male,

female, Wooramel River. Gascoyne Junction-Mullewa

Rd. 11 Sep 1979, J. Blyth (NMV);I0 females, Lockyer

Gorge, Harding River, Pilbara, 19 Oct 1979, J. Blyth

(NMV); female, Fortescue Falls, Hamersley Ra. Natl

Pk, 27 Oct 1979, J. Blyth (NMV); 2 males, female,

21°37’S 117°06’E, Midstream Natl Pk, 24 Apr 1992,

P.S. Cranston (ANIC).

Diagnosis. In common with O. blythi and O. par¬

alleled O. glebtila has the prcanal appendages free

and lateral lobes of the inferior appendages stout.

It is clearly distinguished from O. blythi and O.

parallela by the more rounded basoventral lobes

of the inferior appendages and pair of parameres

of unequal length lying dorsally along the length

of the phallus.

Description. Genitalia, Figs 19-21, 32. Segment

IX short laterally and dorsally; preanal

appendages free, elongate. Tergum X reduced to

a short, rounded lobe. Inferior appendages shorter

than in O. blythi , main body and basodorsal lobe

slender in lateral view; in ventral view inferior

appendages have a lumpy appearance, main body

and basoventral lobe stout, basodorsal lobe

diverges from the phallus. Phallus (Fig. 32)

slender medially, overlying it the parameres in

the form of a long, loose, membranous spine on

right, a shorter, hooked spine on left; phallobase a

bract-like sheath.

Etymology. Descriptive of the general shape of

the male genitalia; Latin — gleba — lumpy.

Distribution (Fig. 40). North-west Western Aus¬

tralia, from Geraldton, about 400 km north of

Perth, to north of the Pilbara region.

Oecetis parallela sp. nov.

Figs 22-24, 32,41

Material examined. Holotype. Male, North Queensland.

Upper Freshwater Creek, Whitfield Range nr Cairns, 3

Apr 1975, M.S. Moulds (NMV WTH-I412, T-17397).

Paratypes. Queensland: 4 males, same data as for

holotype (NMV); male. Mossman Gorge, 16 Jun 1971,

E.F. Rick (ANIC: WTH-1413); 3 males. Little Mul-

grave R„ 28 Jun 1971, E.F. Riek (ANIC: WTH-1411);

2 males. Cap Creek, Mt Finlayson Range, S of Cook-

town, 23 Nov 1974, M.S. Moulds (NMV WTH-1414);

male. Kearney Falls. Goldsborough Valley, 26-27 Mar

1977, G. Theischinger and L. Mueller (ANIC); 5 males.

State Forest, 24 km along Goldsborough Rd nr Gordon-

vale, 27 Dec 1980, M.S. and B.J. Moulds (NMV

WTH-1410).

Other material examined. Queensland: 5 females,

same data as for holotype (NMV); male, Kuranda, 15

Jun 1971, E.F. Riek (ANIC); female, Mossman Gorge,

16 Jun 1971. E.F. Riek (ANIC: WTH-1413); 10

females. Little Mulgrave R., 28 Jun 1971, E.F. Riek

(ANIC: WTH-1411); 3 males, Lock-Davics Creek Rd,

Lamb Ra., Mareeba District, 10 Nov 1974, M. Moulds

(NMV WTH-1395); female. Cap Creek, Mt Finlayson

Range, S of Cooktown. 23 Nov 1974. M.S. Moulds

(NMV WTH-1414); 2 males. Cog Creek, Mt Finlayson

Ra., S of Cooktown, 25 Nov 1974, M.S. Moulds (NMV

WTH-1416); 3 males, 4 females. Upper Freshwater

Creek, Whitfield Range nr Cairns, 15 Dec 1974, M.S.

Moulds (NMV WTH-1409); male. Forty Mile Scrub.

65 km SW of Mt Garnet. 19 Dec 1974, M.S. Moulds

(NMV); male, 2 females, 16 km W of Ravenshoe, 2 Jan

1975, M. Moulds (NMV WTH-1394); male, Tully

Falls, S of Ravenshoe, II Jan 1977, M.S. and B.J.

Moulds (NMV WTH-1415); 3 females, Keamev Falls,

Goldsborough Valley, 26-27 Mar 1977, G. The¬

ischinger and L. Mueller (ANIC); male, Currunda

Creek, trib. of Freshwater Creek, Cairns District, 30

Apr 1979, A. Wells (NMV); male. Birthday Creek

Falls, via Paluma, 11 Apr 1980. I.D. Naumann and J.C.

Cardale (ANIC); male, I5“47'S I45°I4 E, Shiptons

Flat, 17-19 Oct 1980, J.C. Cardale (ANIC); 2 females.

State Forest, 24 km along Goldsborough Rd nr Gordon-

vale, 27 Dec 1980, M.S. and B.J. Moulds (NMV

WTH-1410); male. 17°02'S I45°37'E, 3 km NbyE Mt

Tip Tree, 20 Oct 1980, J.C. Cardale (ANIC).

Diagnosis. In common with O. blythi and O. gle-

bula, O. parallela has the preanal appendages free

and lateral lobes of the inferior appendages stout.

It is readily distinguished from both by the almost

straight-sided more elongate basodorsal lobes of

the inferior appendages in ventral view, and

narrower lateral lobes in lateral view.

Description. Genitalia, Figs 22-24, 32. Segment

IX short middorsally, otherwise relatively long;

preanal appendages elongate, slender. Tergum X

reduced to a short truncate lobe. Inferior

appendages with all lobes about the same length:

basoventral lobe produced more than in other

species, almost straight-sided, apically truncate,

main body narrower, rounded apically, baso-

dorsal lobe slender. Phallus (Fig. 32) long and

slender; parameres in form of a set of unequal

sclerotised spines ventral to the sheathing

phallobase.

Etymology’. Descriptive of the almost parallel

alignment, in ventral view, of the lobes of the

inferior appendages.

Distribution (Fig. 41). Far north-eastern

Queensland, particularly in the Atherton Table¬

land area.

OECETIS COMPLEXA AND ALLIED SPECIES

Oecetis blythi

Oecetis glebula

Figures 34-41, collecting sites for Oecetis complexa-gtoup species: 34, O. complexa Kimmins; 35, O. paracom-

plexa sp. nov.; 36, O. obliqua sp. nov.; 37, O. adelaidica sp. nov.; 38, O. uptoni sp. nov.; 39, O. blythi sp. nov.; 40,

O. glebula sp. nov.; 41,0, parallela sp. nov.

ALICE WELLS

Discussion

Within the Australian Oecetis fauna, species of

the complexa -group are distinct and possibly so in

the world fauna too. It is apparent that these Aus¬

tralian taxa are not accommodated in Chen’s

(1992) scheme and thus further assessment of the

group in the broader context is needed.

Species of the complexa -group are found more

or less peripherally on the Australian continent

(Figs 34 41), although as yet they are unknown

from south-western Western Australia or from

Tasmania. Several species appear to be quite

widespread but most have been collected in the

more northerly parts of Australia. The curiously

disjunct distributions seen here in O. complexa,

O. paracomplexa and O. blylhi are a recurrent

feature among species in the Australian Oecetis

fauna. The veracity of the identifications has been

checked carefully. Possible explanations are gaps

in sampling (real, but unlikely to be the full expla¬

nation given the intensity of collecting Australia

wide), high vagility, or loss of diversity in parts of

the continent as a result of aridification and/or

recent agricultural or pastoral practices. An alter¬

native explanation may lie in differences in the

biology of the species.

Acknowledgements

My thanks go to Arturs Neboiss who made avail¬

able to me the material from Museum Victoria

and checked through a first draft of the

manuscript, and to the editor, for critical

comments that led to improvements.

References

Chen, Y.E., 1992. Revision of the Oecetis (Trichoptera:

Leptoceridae) of the World. Unpublished PhD

Thesis, Clemson University, South Carolina, USA

Ilickin, N.E., 1967. Caddis larvae. Lan’ae of the British

Trichoptera. Associated University Presses, lnc. :

Cranbury, New Jersey.

McLachlan, R., 1877. A monographic revision and syn,

opsis of the Trichoptera of the European fauna

Vol. 6: 281 348. J. van Voorst: London.

Mosely, M.E. and Kimntins, D.E., 1953. Thu

Trichoptera (caddisjties) of Australia and Nev,

Zealand. British Museum (Natural History);

London. 550 pp.

Neboiss, A., 1986. Atlas of the Trichoptera of the

SW Pacific Australian Region. Dr W. Junk

Publishers: Dordrecht.

Neboiss, A., 1989. The Oecetis reticulata species-group

from the south-west Pacific area (Trichoptera:

Leptoceridae). Bijdragen tot de Dierkunde 59(4)'

191-202.

Ross, H.H., 1944. The caddis flies, or Trichoptera of

Illinois. Bulletin of the Illinois Natural History

Survey 23: 1-326.

Schmid, F„ 1987. Considerations diverses sur quelques

genres Leptocerins (Trichoptera, Leptoceridae)

Bulletin de / 'Institut Royal des Sciences Naturellex

de Belgique. Entomologie Supplement 57: 1147.

Walker, K.. Neboiss, A.. Dean. J. and Cartwright, D„

1993. A preliminary investigation of the caddis-

/lies (Trichoptera: fnsecta) of the Queensland

Wet Tropics World Heritage Area. Museum of

Victoria, Department of Entomology: Abbotsford,

Victoria 177 pp.

Wells, A.. 1990. The microcaddisflies (Trichoptera:

Hydroptilidae) of north Sulawesi. Invertebrate

Taxonomy 3: 363—406.

Wells, A., 1991. A guide to the caddisflies (Tri¬

choptera) of the Alligator Rivers region, Northern

Territory. Office of the Supervising Scientist for

the Alligator Rivers Region, Open File Record No.

84.

Wiggins, G.B., 1977. Larvae of the North American

caddisfly genera (Trichoptera). University of

Toronto Press: Toronto and Buffalo.

Memoirs of Museum Victoria 58(1): 89-124 (2000)

THAUMATELSONINE STENOTHOIDS (CRUSTACEA: AMPHIPODA): PART 1

Traudl Krapp-Schickel

Forschungsinstitut u. Museum A. Koenig, Adenauerallee 150, D-53113 Bonn, Germany

(traudl.krapp@uni-bonn.de)

Abstract

Krapp-Schickel, T., 2000. Thaumatelsonine stenothoids (Crustacea: Amphipoda): Part 1.

Memoirs of Museum Victoria 58(1): 89-124.

The Thaumatelsonidae were separated from Stenothoidae by Walker (1906) and reduced to

a subfamily of Stenothoidae by Barnard (1972). The history of the subfamily is reviewed and

its included genera placed in two divisions. All genera and species of the less specialised divi¬

sion are revised here. Four new genera arc erected, Raukumura, Ptychotelsotl, Pycnopyge and

Yarra. Raumahara waroona and Yurra unguiserm are described as new species, Raumahara

rongo Barnard placed in the newly erected genus Raukumara. All belong to this thau¬

matelsonine group but it is shown that similar structures in Pycnopyge are homoplasies which

developed independently. This genus is removed from Thaumatelsoninae.

Contents

Abstract .89

Introduction .89

Taxonomy.90

Key to thaumatelsonine species with spoon-shaped telson.90

Raumahara Barnard .91

Raukumara gen. nov.104

Pycnopyge gen. nov.104

Ptychotelson gen. nov.106

Chucullba Barnard .109

Pseudothauniatelson Schellcnberg .112

Thaumatelsonella Rauschert and Andres .113

Yarra gen. nov.117

Conclusions .123

Acknowledgments.123

References .123

Introduction

While Walker (1906) was studying the

amphipods collected during the Discovery

Antarctic expedition, he noted a totally unusual

telson-shape and created a new genus for which

he coined the generic name Thaumatelson (from

Greek “thauma", meaning “wonderful”). Gur-

janova (1938) erected Parathaumatelson for

Metopella nasica Stephensen, 1927. At the same

time she created the family Thaumatelsonidae,

comprising her genus, Thaumatelson Walker,

1906, Prothaumatelson Schellcnberg, 1931 and

Pseudothauniatelson Schellenberg, 1931, all

having a very unusual urosome.

Barnard (1964: 71) was clearly uneasy with the

generic division within the Thaumatelsonidae

(and indeed the Stenothoidae s.l.) and later

(1972b: 158) opined .. the Stenothoidae being

unnaturally systematised... at present signify not

phyletic lines but grades of evolution ..

Barnard (1972a: 318) erected Raumahara and in

1972b noted (p. 160) “... this genus bridges the

differences between Stenothoidae and Thauma¬

telsonidae to such an extent that the two families

become qualitatively inseparable. The concept of

a subfamily ‘Thaumatelsoninae’ remains useful.”

Barnard had studied this mainly Australian group

while having at his disposal very few specimens

TRAUDL KRAPP-SCHICKEL

(often single representatives) from Australian

collections. His species have never been reported

since.

Comparison of new material, studied at

Museum Victoria (Melbourne), the Australian

Museum (Sydney) and the Smithsonian Insti¬

tution (Washington) revealed some errors in

descriptions and drawings of this group. New

taxa could be erected, others removed. This paper

deals with only one group of thaumatelsonines,

those that share a proximally thickened, three-

dimensional ly inserted and spoon-shaped telson

(Table 1). Others will be discussed in later contri¬

butions. Similarities and differences between the

groups are discussed in Conclusions at the end of

this contribution.

The habitus was observed in glycerine, under

stereo- as well as phase contrast microscope;

dissected parts were mounted in Faure’s medium.

The following abbreviations are used (in alpha¬

betic order): AM, Australian Museum, Sydney,

Australia; BMNH, Natural History Museum,

London, United Kingdom; CTK, personal collec¬

tion of Traudl Krapp, Bonn, Germany; MVR,

Museo C'ivico di Storia Naturale, Verona, Italy;

MNB, Museum fur Naturkunde dcr Humboldt-

Universitat, Berlin, Germany; NHRS, Naturhis-

toriska Riksmuseum. Stockholm, Sweden; NMV,

Museum Victoria, Melbourne, Australia; NZOI,

New Zealand Oceanographic Instilut (now

National Institute for Water and Atmosphere),

Wellington, New Zealand; SAM, South African

Museum, Cape Town, South Africa; USNM, US

Natural History Museum, Smithsonian Insti¬

tution, Washington, USA; WAM, Western

Australian Museum, Perth, Australia; ZMH,

Zoologisches Museum der Universitat, Hamburg,

Germany.

Table 1. Species treated in this paper, with current generic placement.

Original combination

Current generic placement

Chucullha alia Barnard. 1974

Chucullba

Chucullba warea Barnard, 1974

Chucullba

Pseudolhaumatelson patagonicum Schellenberg, 1931

Pseudolhaumatelson

?Raumahara virdurorum Krapp-Schickcl and Andres, 1998

Ptychotelson

Prothaumatelson carinaturn Shoemaker, 1955

Pycnopyge

Raumahara dertoo Barnard, 1972a

Raumahara

Raumahara judithae Moore, 1981

Raumahara

Raumahara noko Barnard, 1974

Raumahara

Raumahara waroona sp. nov.

Raumahara

Raumahara rongo Barnard, 1972b

Raukumara

Pseudolhaumatelson cyproides Nicholls, 1938

Thaumatelsonella

Thaumatelsonella kingelepha Rauschert and Andres, 1991

Thaumatelsonella

Yarra unguiserra gen. et sp. nov.

Yarra

Taxonomy

Key to thaumatclsonine species with spoon-shaped telson

1. Gnathopod 2 chelate.2

Gnathopod 2 subchelatc....7

2. Gnathopod 1 subchelate.3

Gnathopod 1 chelate. Raumahara..A

3. Antenna 1 article 2 swollen. Ptychotelson virdurorum

Antenna 1 article 2 slender, ordinary. Ruukumara rongo

4 Uropod 3 ramus article 2 well developed; antenna 2 article 3 subquadrate

. Raumahara noko

Uropod 3 ramus article 2 reduced; antenna 2 article 3 ovoid.5

5. Urosomite 1 dorsal hump reaching 50% of telson. Raumahara dertoo

Urosomite 1 dorsal hump reaching <25% telson.6

THAUMATELS0N1NE STENOTHOID AMPHIPODA: PART 1

6. Uropod 1 rami unequal, uropod 3 ramus article 1 with 1 seta shorter than

uropod 3 tip; gnathopod 2 propodus proximally bulbous.

. Raumahara judithae

Uropod 1 rami subcqual, uropod 3 ramus article 1 with long seta sitting upon

rounded tip of uropod 3; gnathopod 2 propodus proximally slender.

. Raumahara waroona

7. Antenna I articles 1 and 2 swollen but not nasiform; gnathopods 1 and 2

rectangularly lengthened, palms similarly oblique. Chucullba warea

Antenna 1 article 1 nasiform, lengthened, considerably or totally overlapping

article 2; gnathopods not as above.8

8. Gnathopods 1 and 2 propodus with parallel margins, palms much shorter

than posterior margin, palmar angle on gnathopod 2 more strongly

developed than on gnathopod 1. Chucullba alia

Gnathopods 1 and 2 propodus margins not parallel, palms not as above.9

9. Antenna 1 article 2 strongly nasiform, geniculate; gnathopods 1 and 2 dactyli

strongly serrate... Yarra unguiserra

Antenna 1 article 1 strongly nasiform; gnathopod 1,2 dactyli smooth.10

10. Uropod 2 rami clearly unequal. Pseudothaumatelsonpatagonicum

Uropod 2 rami scarcely different.11

11. Gnathopod 1 palmar corner angular; basis of gnathopod 1 with many short

setae; uropod 3 peduncle shorter than ramus article 1; ramus article 2 three

times as long as wide. Thaumatelsonella cyproides

Gnathopod I palmar corner rounded; basis of gnathopod I with long, dense

setae; uropod 3 peduncle as long as ramus article 1 and as article 2; ramus

article 2 about twice as long as wide. Thaumatelsonella kingelepha

Raumahara Barnard

Raumahara Barnard, 1972a: 318. — Barnard, 1972b:

160. — Barnard and Karaman, 1991: 397. — Moore,

1981: 959. — Krapp-Schickel and Andres, 1998: 52.

Type species. Raumahara dertoo Barnard, 1972

(original designation).

Diagnosis. General habitus bivalve-like (fig. 1).

Antenna 1 peduncular article 2 ordinary to

swollen, with dorsal process more or less over¬

lapping article 3; accessory flagellum of 1 article.

Antenna 2 article 3 rectangularly shortened, ovoid

or bulbous. Mandibular palp reduced to 1 short

article, or lacking. Maxilla I palp biarticulate.

Maxilla 2 plates ordinary (tandem), or outer plate

riding up on inner one. Maxillipcd outer plate pre¬

sent, different, short. Gnathopods 1 and 2

propodochclate-forcipate. Pereopods 3-7 basis

rectilinear, dactylus on inner and outer surface

with a row of stiff, short setae, distally ending in

a second tip. Pereopod 4 elongate, usually same

as pereopod 3. Urosomite 1 dorsally thickened to

a fold vaulting over urosomite 2 and reaching or

overlapping telson insertion. Transparent septae

(or wings) on urosomite 3. Uropod 3 ramus

clearly of 2 articles, or article 2 reduced or lack¬

ing. Telson horizontal, spoon-shaped, dorsally

excavated; posterior end of urosomite 1 process

possibly fitting in this excavation; proximally

thickened, three-dimensionally coalesced with

urosomite 3.

Table 2 tabulates differences between the four

species now placed in this genus.

Raumahara dertoo Barnard

Figures 1, 18

Raumahara dertoo Barnard, 1972a: 318, figs

193-194. — Barnard, 1974: 113, fig. 69, 70.

Type material. Molotype. Western Australia, Cape Nat-

uraliste, intertidal, common seaweeds, WAM (female,

1.55 mm).

Material examined. Paratypes. 6 specimens (USNM

230426).

Non-type. Western Australia. Walpole, ZMH

K38211 (1 individual mounted on slide). Albany, ZMH

K38203 (6 specimens in alcohol), ZMH K38204 (6

specimens in alcohol), ZMH K38205 (1 specimen in

alcohol). Walpole, ZMH K.3821I (1 specimen in

alcohol).

Diagnosis. Antennae 1 and 2 subequal. Antenna I

peduncular article 2 ordinary or weakly swollen;

accessory flagellum of 1 article. Antenna 2 article

3 ovoid. Ocular lobes short, acute. Mandibular

palp lacking. Gnathopod 1 propodochelate;

gnathopod 2 propodochelate, dactylus about one-

third propodus. Urosomite 1 with thick fold vault¬

ing over urosome and half of telson (fig. 1).

TRAUDL KRAPP-SCHICKEL

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THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

Figure 1. a, general habitus of Raumahara or Ptychotelson in normal position, where most parts of legs and

antennae are hidden by the enlarged coxae; on anterior end thickened peduncle antenna 1, on posterior end

thickened urosome plugging or protecting the filter groove.

b, Raumahara dertoo Barnard, urosome of 1.3 mm juvenile, showing the transparent “wings” connecting telson and

uropod 3.

c, Raumahara dertoo Barnard, urosome of 1.5 mm adult; wings not visible as telson adjacent to uropod 3. Scale =

0.1 mm.

TRAUDL KRAPP-SCHICKEL

Uropod 3 ramus with long, visible seta subapi-

cally on article 1; article 2 may be prolongation

on article 1, surpassing insertion of seta (fig. 1).

Telson spoon-like, three-dimensional, proximally

thickened and coalesced with urosomite 3 ven-

trally, depth in proximal half about two-thirds

total length; directed horizontally.

Length 1.0-1.55 mm.

Distribution. Cape Naturaliste to Albany, south

Western Australia; intertidal, common seaweeds.

Biology. Female ovigerous from September to

November.

Remarks. Barnard’s assignment of this species to

Stenothoidae was probably based on the lack of a

nasiform process. Antenna 1 article 2 is sub¬

quadrate to weakly swollen; antenna 2 article 3 is

not rectangular-slender but ovoid (drawn but not

in diagnosis of Barnard, 1972a); and uropod 3

ramus consists apparently of 1 article (although

article 2 may be the minute, thin prolongation

next to a long and striking seta inserted sub-

apically on article 1). Barnard’s drawings of the

urosome never show the ventral margin of the

vaulting process of urosomite 1 which is a very

thick fold for fortifying the whole urosome dor-

sally, and probably also plugs the body pos-

teroventrally if necessary. There are some (glan¬

dular?) ducts of unknown function on the dorsal

margin of this process (fig. 1). The telson can be

in contact with uropod 3 but can also be in an

upright position when transparent septae on the

dorsal margins of urosomite 3 and uropod 3

peduncle become visible (fig. 1).

Raumahara noko Barnard

Figures 2, 3

Raumahara noko Barnard, 1974: 113-117, figs 71,

72.

Type material. Holotype. South Australia. Pearson

Island, in algae, 18-24 m. SAM C3535 (female, 1.6

mm, in alcohol).

Material examined. Western Australia. Walpole, ZMH

K39213, K.39214 (1 adult, 1.6 mm. on 2 slides).

Albany, ZMH K38204 (2). Ceduna, ZMH K38206 (1).

South Australia. Port Lincoln, ZMH K.38207 (7). Kan¬

garoo Island, Hartmann, ZMH K38783 (3). Tasmania.

The Gardens. St Helens, from tide pools. G.C.B. Poore,

Feb 1980, NMV J3278. (1 juvenile on slide).

Diagnosis. Antennae I and 2 subequal. Antenna 1

peduncular article 2 weakly swollen, with age

increasingly vaulting over article 3 proximally;

accessory flagellum of 2 articles. Antenna 2

article 3 exceptionally short, subquadrate to

trapezoid. Ocular lobes strong, subacute.

Mandibular palp reduced to 1 short but clearly

visible article. Gnathopod 1 propodochelate.

Gnathopod 2 propodochelate, slender, dactylus

about 30% propodus length. Urosomite 1 with

fold vaulting over urosomite 2 and one third of

telson, less voluminous than in type species.

Uropod 3 ramus with 2 robust articles. Telson

like a very flat spoon, basally thickened, three-

dimcnsionally coalesced with urosomite 3.

directed and inserted horizontally.

Length 1.0-2.0 mm.

Redescription. Antenna 1 subequal to shorter

antenna 2. Antenna I subcqual to head + pere-

onites 1+2 (dorsal length); peduncular articles 1

and 3 without process, article 2 dorsomedially

slightly vaulting over article 3, extension depend¬

ing on age, reaching as far as one third article 3,

process bearing a duct and 1 long seta; ratio of

lengths of articles 2 : 1 < 0.5; article 3 longer than

article 2 ventrally, subequal dorsally; accessory

flagellum short. Inarticulate; article 2 much nar¬

rower than basal article and with 1 long terminal

seta; flagellum longer than peduncle, of 7 articles,

aesthetascs present. Antenna 2 peduncular article

3 shortened, subquadratc, article 4 shorter than 5:

flagellum subequal to peduncle, of 7 articles.

Upper lip ventrally notched, lobes rounded.

Mandibular incisor well developed, with broad

cusps; lacinia mobilis on left mandible broad,

with major incisions, on right mandible with dis¬

tal part finely serrated; raker spine row devel¬

oped, 4 strong spines; molar cusp with 3 robust

curved spines; palp clearly developed, of 1 short

article; 1-2 setae apically (cf. fig. 2 in juvenile

with 1 seta, fig. 3 in adult with 2 setae; cf. also

Barnard, 1974: fig. 71 with 1 seta, on other draw¬

ings of mandible no palp shown). Lower lip inner

lobes coalesced, mandibular lobes well devel¬

oped. Maxilla 1 inner plate with 1 subapical seta;

outer plate mediodistally setose, apically armed

with 4 serrate robust spines, 1 stout simple spine

and 1 thin, short and stiff one; palp biarticulate,

extending beyond outer plate, with spines apically

and medioapically. Maxilla 2 plates tandem-

grouped; outer plate scarcely extending beyond

inner one, apically with 3 setae (cf. fig. 3 here

with Barnard, 1974: fig. 71 showing 4 setae), I on

outer margin; inner plate with 2 long apical setae,

on inner margin 1 shorter, stiff one mediodistally

Maxillipcd inner plate reaching half length ol

ischium, with 1 seta and 2 spines apically; outer

plate reaching 50% along merus ( = palp article

1); with 2 setae apically, 2 medially; palp of 4

articles; length of merus and propodus subequal:

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

Figure 2. Raumahara noko Barnard, a, habitus 1.1 mm, b, anterior body of neonatus. c, sketch of mouthparts in situ.

Antennae 1, 2 at scale = 0.1 mm. Mandible at scale = 0.05 mm.

TRAUDL KRAPP-SCHICKEL

Figure 3. Raumahara noko Barnard. Gnathopods 1,2 at scale x = 0.1 mm; urosome (from 4 different specimens and

in different shape of fixation; attention to the “wings” connecting uropod 3 with telson) at scale y = 0.1 mm; uppd

lip and maxilla 2 at scale z = 0.05 mm.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

article 3 (propodus) much narrower than article 2;

carpus, propodus distally and dactylus with inner

margin furry.

Gnathopod 1 shorter than gnathopod 2. Coxa 1

reduced, subquadrate, covered by coxa 2; basis

with translucid lobe; merus posterodistally

rounded, about as long as triangular carpus, which

is very narrow between merus and propodus dis¬

tally; propodus with parallel margins anter- and

posteriorly, with 1 strong seta anteromedially and

2 weak ones distally; posterior margin slightly

concave, length : breadth >2:1; propodochclate,

palm armed with short setae; palmar comer with

a pair of robust spines, tip of propodus upwards

curved; dactylus as long as palm, inner margin

smooth, beset with few spines. Gnathopod 2

propodochelate-forcipate. Coxa 2 expanded, ante¬

rior margin regularly convex, posterior one nearly

straight, ventrally straight to rounded; posterior

margin with single spines; ischium slim, longer

than merus; carpus subtriangular, lobe with dense,

short setae and longer, forward-curved spines;

propodus slender, length ratio of dactylus : total

propodus = 33-40% (in juvenile propodus rela¬

tively shorter, thus dactylus more than one third

propodus); terminally a hooked cusp and notch;

palm with stiff setae; dactylus also terminally

hooked, matching well with the propodal notch;

inner margin with similar setae; outer margin with

1 long seta.

Pereopods 3 and 4 slender, subequal, but carpus

in pereopod 4 longer; dactylus about half pro¬

podus length, hind margin on inner and outer sur¬

face with dense, stiff setae, at distal end of this

row a second tip, thus dactylus seemingly bifid.

Coxa 3 more than twice as deep as broad; pos¬

terodistally some spines. Coxa 4 dominant, wider

than deep, getting wider with age; front margin

slightly convex, ventral margin straight, posterior

margin strongly convex; depth anteriorly and pos¬

teriorly subequal. Pereopods 5-7 slender, ratios of

corresponding articles only slightly different;

dactylus with prominent setal comb on anterior

margin inside and outside. Coxae 5-7 short; coxa

5 trapezoid, weakly produced posterodistally;

coxa 6 oblong, roundly produced posteriorly;

coxa 7 blunt.

Gills and oostegites on pereopods 2-5, not

found on pereopods 6 or 7.

Uropods 1-3 extending less far posteriorly in

succession. All rami (and distal regions of pedun¬

cles) with very short setae dorsally. Uropod 1

peduncle subequal or longer than subequal rami;

peduncle with 3 spines; inner ramus inserted

much less distally; outer ramus (extending the

further and more distally inserted) with 1 strong

medial spine. Uropod 2 peduncle as long as

longer ramus, with 1 strong spine distally, 1 medi¬

ally; inner ramus very little shorter and inserted

more proximally; without spination. Uropod 3

peduncle 4- ramus about as long as peduncle of

uropod 2; peduncle distally with 1 spine, subequal

to biarticulate ramus; ramus article 1 > article 2,

article 1 distally with 1 spine, article 2 nail-like,

well developed. Large, transparent septum on

urosomite 3 dorsally; on uropod 3 peduncle

dorsally also a thin, transparent “wing”.

Telson not reaching end of uropod 3 ramus;

basis situated under the vaulted urosomite I and

fused with urosomite 3 medially (fig. 3); distally

tapering, apex rounded, spoon-like, dorsolaterally

and basally with sclerotic clasps; with 1-2 pairs

of setae. Telson not always in contact with body

as urosomite 3 has 2 thin plates in the gap (figs 2,

3), and uropod 3 and telson may be retracted (and

protected?) between them (cf. Barnard, 1974: fig.

71, uropod 3). (This may be what Barnard (1974:

113) meant by “urosomite 3 very distinct.”)

Colour. Antenna 1 articles 1 and 2, eyes, and

broad dorsal stripe on pereonites 1-7 brick-red,

mesosome and urosome orange, gnathopods 1

and 2 and pereopods 3-7 yellow-green, coxae

yellow.

Distribution. South coast of Australia from

extreme west to Tasmania. Littoral and sub¬

littoral, 18-24 m (Barnard, 1974) including tide

pool.

Biology. Eggs in December to January.

Remarks. Having a well developed uropod 3

article 2, this species is the most plesiomorphic

within its genus. Also, the flat spoon-like telson,

rectangular antenna 2 article 3 and mandibular

palp, though very reduced, confirm this impres¬

sion. The other species, R. dertoo, R.judithae and

R. waroona have reduced or no second article on

uropod 3 ramus, an ovoid to bulbous antenna 2

article 3 and no mandibular palp. The process on

antenna 1 article 2 shows a (glandular?) duct and

a long sensory seta; the process on urosomite 1 has

some (glandular?) ducts dorsally.

This species was described from a single

female of 1.6 mm and has not been reported since.

As Barnard’s description is very short a detailed

redescription is given.

Raumahara judithae Moore

Figures 4, 5

Raumahara judithae Moore, 1981: 959, figs 14, 15.

Type material: Holotype. Tasmania, Tinderbox, from

algae, 3-M m, BMNH 1980:187 (female, 1.5 mm).

98 TRAUDL KRAPP-SCHICKEL

Figure 4. Raumahara judithae Moore. Habitus female, 1.8 mm; antenna 1,2, maxilliped at scale x = 0.1 mm; max¬

illae 1, 2 at scale y = 0.1 mm; mouthpart-complex (mandible, upper and lower lip) at scale z = 0.1 mm.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

Figure 5. Raumahara judilhae Moore. Gnathopods 1, 2, urosome at scale = 0.1 mm; pereopods 3, 4 at scale = 0.2

mm; uropod 3 in free enlargement.

100

TRAUDL KRAPP-SCHICKEL

Material examined. Western Australia, Albany, ZMH

K38785 (2). South Australia, Kangaroo Island, ZMH

K38210 (3).

Diagnosis. Antenna 1 peduncular article 2 nasi-

form, swollen; accessory flagellum with 2

articles. Antenna 2 article 3 bulbous. Mandibular

palp lacking. Gnathopod 1 propodochelate.

Gnathopod 2 propodochelate, propodus strongly

bulging anteroproximally and less slender than in

other species; dactylus about 40-50% of propo¬

dus length. Urosomite 1 with fold vaulting over

urosomite 2 and insertion of telson; uropod 3

ramus with 2 articles, article 2 minute, vanishing,

spiniform, articulation between article 1 and arti¬

cle 2 vanishing; telson boat-like, three-dimen¬

sional, proximally coalesced with urosomite 3,

inserted horizontally.

Length 1.2-2.0 mm.

Distribution. Tasmania; Kangaroo Island (South

Australia), Albany (Western Australia); 3-4 m

depth.

Biology. Female ovigerous in November and

December.

Remarks. Gnathopod 1 propodus has 1 or 2

medial (plus 1 distal) spines anteriorly. Gnatho¬

pod 2 appears to be more powerful than in other

species, the bulbous basis of propodus showing

strong bundles of muscles. Because of the

“wings” or septae on urosomite 3 the insertion of

uropod 3 is partly hidden. Uropod 3 ramus article

2 is reduced to a tiny, spine-like peak on article 1,

and the articulation is scarcely visible. But, as

mentioned by Barnard (1974: 113), article 2

shows always (as in article 1) a seta at its insertion

and this indicates the region of the barely observ¬

able articulation. Coxa 4 is trapezoid, anteriorly

deeper than posteriorly, only little overlapping

coxa 3.

The urosome is specialised compared to

other members of this genus being the only

species with the rami of uropod 1 clearly differ¬

ent in length, while in uropod 2 the rami are

as in all species slightly unequal. Uropod 3 is

reduced, directly in line with the lengthened uro¬

somite 3 as a robust, solid short cone with spine¬

like article 2 nearly fused to article 1. Spination is

extremely poor; uropod peduncles have only one

spine each, rami are smooth, and the telson has

0-2 setae.

The species was described from a single

specimen and has not been reported again until

now.

Raumahara waroona sp. nov.

Figures 6, 7

Material examined. Holotype. Western Australia.

Groyne, Cape Peron (32°16'S, 115°41 'E), R. Spring-

thoipe, AM P54514 (1 slide, sex?, 1.2 mm).

Paratype. Type locality, AM P54515 (1 specimen in

alcohol, sex?, 1.2 mm).

Additional material. Type locality. AM P54516 (2

specimens in alcohol); AM P54517-P54520 (4 slides).

Diagnosis. Antenna 1 peduncular article 2 with

strong process dorsally. Antenna 2 peduncular

article 3 suboval to subrectangular. Mandibular

palp lacking. Maxilla 1 palp of 2 articles. Maxilla

2 inner plate reduced. Maxilliped very slender,

outer plate short. Gnathopod 1 propodochelate.

Gnathopod 2 propodochelate-forcipate, dactylus

about 33-^40% length of propodus. Urosomite 1

very elongate, with large hump overlapping; uro¬

somite 2 reduced to small triangle: urosomite 3

rectangular, fused with broad basis of telson.

Uropods 1 and 2 with subequal rami, subequal to

peduncle, without spines or setae; uropod 3 ramus

clearly of only 1 article, semicircularly rounded,

not reaching ramus insertion of uropod 2, article 2

absent; with 1 apical, centrally inserted, long seta.

Hump of urosomite 1 reaching half way along

telson. Telson spoon-like, deeply excavated.

Length 1.0-1.2 mm.

Description. Body smooth, very similar to most

species of Raumahara. Head about as long as

pereonite 1. Rostrum inconspicuous. Eyes of

stenothoid shape, normal. Lateral cephalic lobes

shortly produced, rounded. Midcephalic keel and

epistome developed. Pereonite 4 about as long as

pereonites 1-3 together. Urosomites I and 2 free;

urosomite 3 broadly coalesced with telson. Uro¬

somite 1 with large dorsal hump vaulting over

urosomite 2 and insertion of telson, hump show¬

ing similar structure as nasiform process on

antenna 1 article 2.

Antenna 1 shorter than antenna 2, as long as

head + pereonites 1-3 together (dorsal lengths).

Antenna 1 peduncle articles I and 3 without pro¬

cess, article 2 nasiform dorsomedially, with very

dense aesthetascs; length ratio of articles 1 : 2 : 3,

about 5 : 3 : 2 on dorsally visible margin, but

articles strongly overlapping; accessory flagellum

short, broad, uniarticulate, 2 terminal setae; flag¬

ellum longer than peduncle, of 7 articles, with

many long aesthetascs. Antenna 2 peduncle

article 3 suboval to subrectangular, article 3 as

long as article 4, article 4 shorter than article 5;

flagellum shorter than peduncle, of 7 articles.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

101

Figure 6. Raumahara waroona sp. nov. Antennae 1, 2, gnathopods 1, 2 at scale 0 = 0.1 mm; maxilla 1, 2, mandible,

upper lip, maxilliped at scale y = 0.1 mm; maxilla 2 in free enlargement.

Figure 7. Raumahara waroona sp. nov. Pereopods 3, 4, 5 at scale x = 0.1 mm; urosome from 3 specimens and in

different fixation, at scale y = 0.1 mm; uropod 3 in free enlargement.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

103

Upper lip ventrally notched, lobes rounded.

Mandible incisor on both sides well developed,

left and right mandible not very different; raker

spine row developed, of 3 (-4?) spines; molar

cusp with 3 robust spines; palp absent. Lower lip

inner lobes coalesced, mandibular lobe present.

Maxilla 1 inner plate subquadrangular, truncate,

with 1 subapieal seta; outer plate with 1 medio-

distal long seta, apically armed with 4 serrate,

robust spines, 1 stout simple spine and I thin,

short and stiff one; palp biarticulate, ratio of

lengths, 2 : 5, extending much beyond outer plate,

with few spines and teeth apically and medioapi-

cally. Maxilla 2 inner plate reduced, short, and

outer plate curved over inner plate (always, or

artefact in slide?); outer plale apically with 3 long

setae, 1 on outer margin; inner plate with 1 long

apical seta, 1 shorter on inner margin medio-

distally. Maxilliped inner plate reaching half

length of ischium, with 1 seta and I spine api¬

cally; outer plate reaching one third of merus ( =

palp article 1); with 1 seta subapically, 1 medi¬

ally; palp of 4 articles; article 3 (propodus) nar¬

row like article 2; propodus distally and dactylus

inner margin furry.

Gnathopod 1 shorter than gnathopod 2; coxa 1

reduced, subquadrate, covered by coxa 2; merus

posteredistally rounded, about as long as trian¬

gular carpus, which is slightly produced between

merus and propodus; propodus rectangular, ante¬

rior margin slightly convex, with 1 seta; posterior

slightly concave; length : breadth about 2:1;

propodochelate, palm armed with small distal and

medial spines; palmar comer curved upwards;

dactylus about 40% of propodus length. Gnatho¬

pod 2 propodochelate-foreipate; coxa 2 expanded,

longer than basis, anterior margin regularly

convex, posterior one nearly straight, ventrally

rounded; posterior and posteroventral margin

with single, short spines; ischium longer than

merus; carpus subtriangular, lobe with dense,

short setae, distoapically some longer ones;

propodus slender, length ratio of dactylus: total

propodus = 33-40%; terminally a hooked cusp

and notch, but no defining spines; dactylus also

terminally hooked, matching well with the propo-

dal notch.

Pereopods 3 and 4 slender, subequal; dactylus

about half propodus length, hind margin on inner

and outer surface with dense, stiff setae; coxa 3

more than twice as deep as wide. Coxa 4 domi¬

nant, wider than deep, front margin slightly

convex, ventral margin straight or very slightly

concave, posterior margin strongly convex, ante¬

riorly deeper than posteriorly. Pereopods 5-7

slender, ratios of corresponding articles only

slightly different; dactylus with prominent setal

comb on anterior margin inside and outside;

coxae 5-7 short; coxa 5 trapeziform, produced

posterodistally; coxa 6 oblong, roundly produced

posteriorly; coxa 7 blunt.

Uropod 2 extending less far posteriorly than

uropod 1, uropod 3 very short. Uropod 1 pedun¬

cle subequal to subequal rami, without spines,

setae or pectination. Uropod 2 peduncle as long as

subequal rami, inner one inserted more proxi-

mally; no spination or pectination like uropod 1.

Uropod 3 ramus consisting of 1 clear article,

semicircular, not reaching ramus insertion of uro¬

pod 2, article 2 absent; with 1 apical long seta,

centrally inserted. Urosomite I very long, large

hump vaulting over urosomite 2 and shortly over

telson insertion; urosomite 2 reduced to small tri¬

angle; urosomite 3 rectangular, fused with broad

basis of telson.

Telson spoon-like, shorter than uropod 1 rami,

about half length of hump of urosomite 1, hori¬

zontally inserted, deeply excavated and with char¬

acteristic semicircular hump proximoventrally;

insertion of telson three-dimensionally thickened

and fused with urosomites 3 and 2.

Distribution. Cape Peron, Western Australia.

Etymology. This material was found on Cape

Peron, south of Perth. A nearby locality with an

aboriginal name is Waroona, which with its

double “o” reminded me of R. tier too. the closest

species.

Remarks. This species is very similar to R.

judithae. While in R. noko uropod 3 is composed

of peduncle and a biarticulate ramus, in R. dertoo

and R. judithae article 2 is vanishing; in R.

waroona sp. nov. no traces of article 2 are to be

found, article 1 is as long as wide and bears api¬

cally one long seta. Thus this new species resem¬

bles the genus Chucullba in having uropod 3

reduced to only one article, probably the pedun¬

cle, with the — steadily present — apical long

seta. Also, maxilla 2 of the present species is sim¬

ilar to Chucullba but the latter has fewer setae.

Finally, reduced body length is common to both.

While basic Raumahara females are ovigerous at

2 mm, the new species, R. waroona , reaches only

1.2 nun and the specialised species of Chucullba

are also rarely longer than 1 mm. However, the

morphology of the gnathopods and the nasi-

form swollen antenna 1 article 2 (instead of arti¬

cle 1) places this new species undoubtedly in

Raumahara.

104

TRAUDL KRAPP-SCHICKEL

Raukumara gen. nov.

Type species. Raumahara rongo Barnard, 1972b.

Diagnosis. Antenna 1 ordinary, peduncle slender.

Mandibular palp lacking. Maxilla 2 plates ordi¬

nary, i.e. in tandem, but outer plate unusually

slender, with 2 long setae apically. Coxa 4

trapezoid, enlarged. Gnathopod 1 subchelate-

rectipalmate. Gnathopod 2 propodochelate-

forcipate, strong. Dactylus of pereopods unusu¬

ally short, inside and outside with comb-like row

of setules; propodus to merus feeble, ischium

very long. Urosomites free, no dorsal process, but

urosomites 2 and 3 triangular, dorsally much

shorter than ventrally; urosomite 1 longer than

urosomites 2+3 together. Uropod 3 ramus of 2

articles. Tclson flat, but spoon-like and excavated

dorsally, horizontally inserted, basically fused

with urosomite 3.

Etymology’. For the Raukumara Range on Auck¬

land Island, New Zealand, alluding also to

Raumahara which the new genus resembles

(feminine).

Remarks. Barnard (1972b: 318) wrote “The reten¬

tion of R. rongo in Raumahara is provisional and

based on the probability that other species will be

discovered that will clarify the generic partition of

the thaumatelsonin stenothoids.”

Characters shared with species of Raumahara

are: antenna 2 peduncular article 3 ovoid, as in R.

dertoo; shape of gnathopod 2 with dactylus c.

40% of propodus as in R. judilhae; a comb of

setules on inner and outer sides of pereopodal

dactyli; the pereopodal dactyli bifid as in all

species of Raumahara ; urosomite 3 and peduncle

uropod 3 forming transparent “wings” dor¬

sally; and the telson basically coalesced with

urosomite 3.

Differences from specialised species of

Raumahara and Ptychotelson gen. nov. are:

antenna 1 peduncular article 2 ordinary, slender

(even in hyperadults), and urosomite 1 ordinary,

without dorsal prolongation, and not longer than

urosomites 2+3 together.

One character shared with Ptychotelson is the

subchelate-rectipalmate first gnathopod.

The characters unique to Raukumara are: pere¬

opods 3-7 dactyli extremely short, their propodus

to merus elongate, feeble, and ischium very long,

3-4 times as long as wide (versus twice as long as

wide in other genera).

This new genus differs from the others in the

following main characters: from Raumahara and

Ptychotelson gen. nov. in the shape of antenna 1

peduncular article 2; ratio of gnathopod 1 to

gnathopod 2; shape of pereopods 3-7; length of

urosomite 1 and spination of uropod 1 peduncle.

It differs from Raumahara in having the gna¬

thopod 1 subchelate-rectipalmate and from

Ptychotelson in having no mandibular palp.

Removal of this species from the (now

monophyletic genus) Raumahara seems justified.

Raukumara rongo (Barnard) comb. nov.

Figure 8

Raumahara rongo Barnard, 1972b: 160-162, figs

91,92.

Type material. Holotype. New Zealand, Kaikoura.

NZOI 102 (female, 2.1 mm).

Material examined. Type locality. USNM 2304273 (3

ovigerous females, 1.8-2.0 mm; 2 juveniles in alcohol).

USNM 230427 (1 female, 1.9 mm, on slide).

Diagnosis. Antenna 2 peduncular article 3 ovoid.

Coxa 2 anterior margin more parallel to posterior

one, less concave than in species of Raumahara.

Gnathopod 2 propodochelate, dactylus more than

one third (about 40%) propodus, proximally

thickened; propodus gnathopod 1 less than half as

long as that of gnathopod 2. Coxa 4 trapezoidal,

anteriorly deeper than posteriorly. Urosomite 3

and uropod 3 peduncle with transparent “wing”

dorsally. Uropod 3 ramus with spine-like article

2, about as long as article 1. Telson like a flat

spoon.

Length 1.5-2.0 mm.

Distribution. New Zealand (Kaikoura, Gisborne,

Whangaparaoa Peninsula, Leigh). Caulerpa

brownii, 5-6 m; in crevices in Durvillaea-zone,

barnacles encrusted with sponge.

Biology. Female ovigerous in January.

Remarks. The “missing seta on maxilla 1 IP"

(Barnard 1972b: fig. 92c) is present as in other

species. The telson is horizontally flat but not

stenothoid-like, as it is dorsally excavated like a

flat canoe or spoon, and — more importantly —

proximally thickened and coalesced with uro¬

somite 3. The “numerous setules” of Barnard’s

description are totally lacking within his type

series but are probably crystals derived during

preservation.

Pycnopyge gen. nov.

Type species. Prothaumatelson carinatum Shoe¬

maker, 1955.

Diagnosis. Antennae 1 and 2 ordinary; accessory

flagellum of 1 rudimentary article. Mandibular

palp lacking. Maxilla 1 palp long, no articulation

THAUMATELSON1NE STENOTHOID AMPHIPODA: PART 1

105

Figure 8. Raukumara rongo (Barnard). Maxilla 1 (left), antenna 1,2, gnathopod 1, pcreopod 4 distally, urosome of

2 specimens, telson at scale 0 = 0.1 mm; maxilla 1 (right) at scale y = 0.1 mm; mandible at scale y = 0.05 mm;

gnathopod 1 ’, gnathopod 2 at scale x = 0.2 mm.

106

TRAUDL KRAPP-SCHICKEL

concemable, thus of I article. Maxilla 2 ordin¬

ary. Maxilliped outer plate vanishing, inner plate

fused. Gnathopod 1 subchelate. Gnathopod 2

propodochelate-forcipate. Pereonite 4 very large.

Urosomite 1 with dorsal fold, but not overlapping

urosomite 2; thus uropod 2 lies over not after uro-

pod 3. Urosomites 2 and 3 coalesced so that uro¬

pod 3 lies over, not after, uropod 3. Uropod 3

ramus article 2 well developed. Telson Hat,

horizontal, not coalesced with urosomite 3, not

spoon-like or dorsally excavated.

Etymology. The Greek word pyge, pygaion means

rump, back, while pyknos (Greek) signifies a con¬

densed, thickened area. Together they describe

the special habitus, the dorsal fold on urosomite 1

not overlaping urosomite 2, and the fusion of uro¬

somites 2 and 3 (neuter).

Remarks. Shoemaker (1955) described, inade¬

quately, a species from Alaska as Prothaumatel-

son carinatum. Barnard (1972a) removed it from

that genus, assigning it provisionally to Rauma-

hara, although he lacked information about

important details.

Characters shared with Raumahara are: uro¬

somite 1 with a dorsal fold (but only protecting

urosomite 1 and not vaulting over urosomite 2 as

in Raumahara ); propodochelate gnathopod 2.

Differences from Raumahara : antennae 1 and 2

peduncles ordinary, flagellae reduced, very short

(versus article 1 nasiform or article 2 swollen to

nasiform in Raumahara): maxilla 1 palp with¬

out discemable articulation, thus probably 1

articulate (v. 2 articles): maxilla 1 outer plate very

broad and blunt, subquadrate (v. slim): maxilliped

outer plate vanishing (v. present); gnathopod

1 propodus subchelate, without palmar corner

(v. chelate-subchelate-rectipalmate), carpus and

merus strongly lobate (v. ordinary); pereo-

pods 3-7 dactyli ordinary, very long, longer

than half propodus, smooth (v. with a charac¬

teristic double row of setae inside and outside);

oostegites large (v. small); uropod I peduncle

clearly longer than rami (v. about equal); uro¬

somite 1 with thickened fold dorsally, but not

overlapping the rest of urosome (v. vaulting

over urosomites 2 and 3 and part of telson); uro¬

somites 2 and 3 coalesced (v. free); telson

ordinary (v. coalesced with urosomite 3,

thickened proximally, spoon-like excavated

dorsally).

Removal from (the exclusively Australian)

Raumahara is necessary and a separate genus

justified.

Pycnopyge carinatum (Shoemaker) comb. nov.

Figures 9, 10

Prothaumatelson carinatum Shoemaker, 1955:

33-34, fig. 13.

Raumahara carinata. — Barnard, 1972a: 318.

Material examined. Type specimens. Off Point Barrow,

Alaska, USNM 181883 (1 slide). USNM 195166 (4

specimens in alcohol).

Diagnosis. Antennae 1 and 2 ordinary, both with

very short flagellum; accessory flagellum rudi¬

mentary article. Mandibular palp lacking.

Gnathopod 1 subchelate, no palmar center; car¬

pus strongly and merus weakly lobed. Gnathopod

2 propodochelate, dactylus about 40% of propo¬

dus length. Pereopods 3-7 dactyli very long,

longer than half propodus, smooth. Urosomite 1

dorsally thickened as a fold, not overlapping.

Urosomites 2 and 3 coalesced. Uropod 1 peduncle

exceeding rami. Uropod 3 ramus of 2 articles,

W'ell developed. Uropod 3 exceeding uropod 2

peduncle. Telson flat, horizontal, not coalesced

with urosomite 3.

Length 1.8-3.0 mm.

Distribution. Arctic Ocean, Alaska, off Point

Barrow (USA). 217 ft (66 m), perforated rocks,

stones.

Remarks. Though having chelate gnathopod 2,

this species does not belong to the (southern

Australian-Antarctic) thaumatelsonine steno-

thoids, as the telson is not fused with urosomite 3,

nor spoon-like or dorsally excavated, and the palp

of maxilla 1 has only one article. Fortification of

the urosome is created in another way, and chela¬

tion of gnathopod 2 is probably a homoplasous

development in Arctic regions.

Ptychotelson gen. nov.

Type species. 1 Raumahara virdurorum Krapp-

Schickel and Andres, 1998.

Diagnosis. Antenna 1 peduncular article 2

swollen. Mandibular palp of ordinary length,

articulation doubtful, seemingly articles coa¬

lesced; maxilla 2 ordinary. Coxa 4 trapeze¬

shaped, enlarged. Gnathopod 1 subchelate-

rectipalmate, gnathopod 2 propodochelate-

forcipate. Urosome segments free, urosomite 1

with dorsal fold vaulting over urosome, > uro¬

somite 2 + 3; uropod 3 ramus 2 articles well

developed. Telson spoon-like excavated dorsally,

horizontally inserted, proximally coalesced with

urosomite 3.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

107

Figure 9. Pycnopyge carinatum (Shoemaker). Antennae 1, 2, gnathopods 1, 2 (detail) at scale = 0.1 mm; gnathopod

2’ (complete) at scale = 0.2 mm.

108

TRAUDL KRAPP-SCHICKEL

Figure 10. Pycnopyge carinatum (Shoemaker). Maxilla 1 at scale x = 0.1 mm; mandible, maxilla 2, maxillipedat

scale y = 0.1 mm; pereopod 7, urosome at scale z = 0.1 mm.

Etymology. Greek ptyx, -ychos means fold,

referring to the three-dimensional, boat- or

spoon- shaped tclson and process of urosomitc 1

overlapping the urosome like a fold or pleat.

Remarks. Krapp-Schickel and Andres (1998)

were doubtful of the generic placement of

IRaumahara virdurorum and recognised that a

thorough revision was needed to understand its

relationship to other species with subchelate-rec-

tipalmatc gnathopod 1 or long mandibular palp.

Now, after a detailed study of type material, it can

be shown that the two species with similar

gnathopod 1 are not closely related. Prothauma-

telson nasutum has a vertical telson but very sim¬

ilar mandibular palp; Raukumara rongo has a less

specialized urosome, a non-swollen ordinary

antenna 1 peduncle and lacks a mandibular palp.

With the closely related species of Raumahara.

R. dertoo. R. noko, R. judithae and R. waroona,

the new taxon shares specialised characters of

“swollen" antenna 1 peduncular article 2 (in R.

dertoo only weakly), propodochelate gnathopod

2, a row of setae inside and outside dactyl i of

pereopods 3-7, special construction of the uro¬

some with its combination of vaulting urosomite

1 and excavated telson. Pfychotelson is less spe¬

cialised in the mandibular palp being of ordinary

length (probably 1-articulate) and the gnathopod

1 subchelate-rectipalmate.

This new genus differs from its nearest

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

109

relatives in the following main characters: from

Raukumara in antenna 1 peduncular article 3

swollen and produced (versus slender, ordinary);

mandibular palp long (v. absent); urosomite I

with a thick fold fortifying the urosome (v. uro¬

somite 1 lengthened but not thickened or folded);

and from Raumahara in long mandibular palp

(v. reduced to a short stump or absent); and

gnathopod 1 subchelate- rectipalmate (v. chelate).

Ptychotelson virdurorum

(Krapp-Schickel and Andres) comb. nov.

Figures 11,12

1 Raumahara virdurorum Krapp-Schickel and

Andres, 1998: 56-58, figs 1, 2.

Material examined. Holotype. South Australia. Steat-

ford Bay, Port Lincoln, ZMH K38388 (ovigerous

female, 1.6 mm, I slide). Type locality, ZMH K38389

(41 specimens in alcohol). Victoria. Point Lonsdale,

ZMH K308390 (4); Cape Otway, NMV J39258,

J39264, J39265 (3 slides); Port Phillip Bay, CTK (1

slide).

Diagnosis. Antenna 1 peduncular article 2 dor-

sally prolonged and swollen (sometimes article 1

also a little thickened). Antenna 2 peduncular

article 3 subquadrate. Mandibular palp well

developed, but no articulation visible. Gnathopod

1 subchelate-rectipalmate. Gnathopod 2 propodo-

chelate-forcipate, dactylus c. one third propodus.

Urosomite I with thick fold vaulting over

urosome and insertion of telson. Uropod 3

ramus with 2 articles. Telson spoon-like, three-

dimensional, proximally coalesced with

urosomite 3, inserted horizontally.

Length 1.0-1.6 mm.

Distribution. South Australia to Victoria; inter¬

tidal, algae (mostly Rhodophyceae including

corallines and epiphytes).

Biology. Female ovigerous in December.

Remarks. Drawings of young and adult specimens

are presented to show how ratios on limbs and

body segments change with age. This allometry

can also be noticed in R. judithae and R. waroona.

Chucullha Barnard

Chuculiba Barnard, 1974: 108.

Type species. Chuculiba alia Barnard, 1974

(original designation).

Diagnosis. Antenna I peduncular article I weakly

swollen or with strong nasiform process; acces¬

sory flagellum of 1 very short article. Antenna

2 peduncular article 3 subrectangular or

trapezoidal. Mandibular palp lacking, mandible

body vestigial. Maxilla 1 palp biarticulate.

Maxilla 2 outer plate riding upon reduced inner

one, plates partly coalesced. Maxilliped outer

plate vanishing. Gnathopods 1 and 2 propodus

very similar, subchelate, rectangularly length¬

ened, palm shorter than posterior margin which is

parallel to anterior one. Pereopods 3-7 similar,

basis rectilinear, ischium elongate, length much

greater than twice breadth (cf. Raukumara)-,

dactyli smooth or with comb of setae. Pereonite 4

elongate, longer than head + pereonites 1-3.

Urosomites 2 and 3 coalesced basally. Uropods 1

and 2, dorsal margins of peduncle and rami pecti¬

nate, inner ramus shortened. Uropod 3 reduced to

I short article (probably peduncle) plus long seta.

Telson spoon-like, three-dimensional, proximally

coalesced with urosomite 3, inserted horizontally,

deeply excavated.

Remarks. Strong processes on antenna 1 also exist

in Raumahara, but on article 2 and not, as here,

on article 1. The maxilla 2 has riding plates as in

Raumahara waroona. Gnathopods 1 and 2 have

propodi alike and with lengthened parallel

margins as is found in Parathaumatelson. The

characteristic comb on dactyli of pereopods 3-7,

inside and outside, always present in Raumahara,

is present in one species of Chuculiba and absent

in the other. Uropodal rami are reduced to a dif¬

ferent degree also in Raumahara. However, in the

latter urosomites 2 and 3 are separated, though

dorsally narrowed and ventrally much wider, thus

giving a triangular shape in lateral view while in

Chuculiba articulation between urosomites 2 and

3 is scarcely visible. C. alia is more specialised

than C. warea in antenna 1, maxilla 1, nathopods,

pereopodal dactyli (only one seta instead of a

comb), length of uropods 1 and 2 inner ramus,

and length of uropod 3.

Antenna 2 peduncular article 4 is always

shorter than article 5 in Raumahara, Pycnopyge.

Raukumara and Chuculiba , whereas it is always

longer in Yarra, Pseudothaumatelson and Thau-

matelsonella. As the members of the first group

live in shallow algae and the latter predominantly

in deeper sands, this difference could be

correlated with feeding behaviour.

Chuculiba alia Barnard

Chuculiba alia Barnard, 1974: 108-110, figs 65, 66.

Type material. Holotype. Western Australia. Middleton

Beach, Albany, intertidal, sandy rocks, coralline algae.

WAM 134-71 (female, 1.41 mm).

Material examined. Western Australia (USNM,

141159; 141160 (4 specimens in alcohol); AM P43271

110

TRAUDL KRAPP-SCHICKEL

Figure 11. Ptychotelson virdumrum (Krapp-Schickel and Andres), a, habitus of adult female, 1.6 mm. b, middle of

body of juvenile, 1.1 mm. c, sketch of mouthparts in situ, d, complete mandible and distal pail seen from other side;

other figures, 4 enlargements of different views of mandible at scale = 0.025 mm.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

111

Figure 12. Ptychotelson virdurorum (Krapp-Schickel and Andres). Antennae 1, 2, gnathopods 1, 2, pereopod 3 or

4 distally, urosome at scale =0.1 mm; maxilla 1,2, maxilliped, antenna 1 ’ at scale = 0.05 mm; maxilla 1 outer plate

enlarged.

112

TRAUDL KRAPP-SCHICKEL

(1 specimen in alcohol). Western Australia. CapcNatu-

raliste, ZMH K38213; Margaret River, ZMH K38215,

K38216; Walpole-Albany, ZMH K38217, K38219,

K38220, K38221. South Australia. Ceduna-Adelaide,

ZMH K38217, K38224. Victoria. Melbourne, Geelong,

ZMH K38218d. (In all c. 100 specimens in alcohol).

Diagnosis. Midcephalic keel weak. Antenna 1

peduncular article! strongly nasiform, largely to

totally overlapping article 2. Maxilla 1 inner plate

lacking seta. Maxilla 2 in riding position but sep¬

arate lobi still distinguishable. Gnathopods 1 and

2 similar, rectangularly lengthened, with parallel

margins, but gnathopod 1 palm oblique and

gnathopod 2 palm nearly subchelate- rectipal-

mate. Pereopods 3-7 dactyli smooth. Urosomite 1

with thick fold vaulting over urosome and inser¬

tion of telson. Uropod 3 consisting of 1 minute

article and short, tiny seta.

Length 1.1-1.4 mm.

Distribution. Western Australia to Victoria.

Intertidal, sandy rocks and coralline algae.

Biology. Female ovigerous at end of September.

Chucullba warea Barnard

Chucuilba warea Barnard, 1974: 110-112, figs. 67,

68 .

Type material. Western Australia. Cape Naturaliste,

intertidal, algae and rocks, WAM 135-71 (female, 1.19

mm).

Material examined. Type locality, USNM 230428,

USNM 141161 (2 specimens in alcohol). Cape Natural¬

iste, ZMH K382I4 (2). South Australia, Ceduna, ZMH

K38223 (2).

Diagnosis. Antenna 1 peduncular article 1 and

article 2 (weakly) swollen. Maxilla 1 inner plate

with seta. Maxilla 2 in riding position, lobi fused.

Gnathopods land 2 similar, both palms oblique.

Pereopods 3-7 dactylus with comb of short setae

as in Raumahara. Urosomite 1 with thick fold

vaulting over urosome and insertion of telson.

Uropod 3 ramus with 1 minute article (probably

peduncle) and long seta apically.

Length 1.2 mm.

Distribution. Cape Naturaliste, Western Aus¬

tralia; Ceduna, South Australia. Intertidal, algae

and rocks.

Biology’. Female ovigerous in September.

Pseudothaumatelson Schellenberg

Pseudothuumatelson Schellenberg, 1931:110.

Type species. Pseudothaumatelson patagonicum

Schellenberg, 1931 (original designation).

Diagnosis. Antenna 1 peduncular article 1

nasiform, accessory flagellum of 1 article.

Mandibular palp present, stout, without articula¬

tion, as long or longer than incisor. Maxilla 1 palp

of 2 articles. Maxilla 2 ordinary. Gnathopods 1

and 2 subchelate, shape similar, size different.

Pereopods 3-7 bases rectilinear, dactyli smooth.

Pereonite 4 elongate. Urosomites 1-3 free;

urosomite 1 with long process vaulting over uro¬

somite 2 and most of telson. Telson medium,

marginally sclerotized, horizontally inserted,

excavated, margins thickened.

Pseudothaumatelson patagonicum Schellenberg

Pseudothaumatelson patagonicum Schellenberg,

1931: 110-112, fig. 60. — Rauschert and Andres, 1991:

228, fig. 1.

Pseudothaumatelson cyproides. — Bellan-Santini

and Ledoyer, 1986: 425-427. fig. 29. — Branch et al„

1991: 12 [not Nicholls, 1938],

Material examined. Syntypes of Pseudothaumatelson

patagonicum. Falkland Is, NHRS 738 (slide), 3603 (I

ovigerous female in alcohol). Falkland Is, MNB 22802

(1 iuvenile in alcohol, partly damaged).

Other material. Prince Edward Is, MVR

Cr3389-Cr3397 (slides).

The two other specimens deposited by Bellan-Santini

and Ledoyer (1986: 425) in SAM could unfortunately

not be found in South Africa or in Marseille.

Diagnosis. Antenna 1 peduncular article 1 nasi¬

form. Mandibular palp long, obtuse, without

articulation, 2 apical and 1 subapical setae.

Gnathopods l and 2 subchelate, propodi

gnathopods 2 : 1 subequal; on both bases anteri¬

orly few long setae. Gnathopod 1 palmar comer

rounded, basis with few setae anteriorly. Gnatho¬

pod 2 palmar comer excavated, with distal hump

and 2 strong spines. Urosomite 1 with vaulting

process. Uropod 1 rami a little different, longer

ramus subequal to peduncle. Uropod 2 rami

clearly different, both shorter than peduncle.

Uropod 3 peduncle shorter than subequal rami.

Telson dorsally shallow excavated, fused with

urosomite 3.

Length 1.8-2.5 mm.

Redescription. After Schellenberg (1931),

Rauschert and Andres (1991), material of Bellan-

Santini and Ledoyer (1986) and complemented

from material examined [square brackets].

Eyes large. Antenna 1 peduncular article 1

nasifonn, accessory flagellum [of 1 article].

Mandibular palp as long as incisor, of 1 article ,

with 2 apical and 1 subapical setae. Gnathopod 1

subchelate, much smaller than gnathopod 2, palm

oblique, about same length as posterior margin of

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

113

propodus, palmar comer rounded, with spines;

basis with a few long setae anteriorly. Gnathopod

2 subchelate (rudirnentarily carpochelate, as car¬

pus acutely prolonged and propodus proximally

narrowed), palm with “pearls” (Bellan-Santini

and Ledoyer lacking), distally concave, defined

by hump with 2 strong spines; anterior margin

smooth or with 1 or 2 long setae; basis with few

long setae on anterior margin. Pereonite 4

elongate, as long as pereonites 1-3. Pereo-

pods 5—7 basis rectilinear, with about 12 short

setae. Urosomites 13 free. Urosomite 1 with pro¬

cess vaulting over urosomites 2 and 3 and over¬

reaching most of telson. Uropod 1 with long,

spinose peduncle, longer than barely unequal

rami [Rauschert and Andres (fig. 1) show the

rami much shorter than peduncle but this is mis¬

leading as rami in type material are broken];

uropod 2 peduncle long, spinose, rami unequal,

the longer shorter than peduncle. Uropod 3

peduncle robust, article 1 about twice as long

as large, article 2 in type material broken [as

long as article 1, pointed]. Telson medium,

half length of urosomite 1 dorsally, margin¬

ally sclerotisized, horizontally inserted, shallow

excavated, coalesced with urosomite 3, with 2

setae.

Distribution. Falkland Islands; Prince Edward

and Marion Islands (subantarctic), 95-197 m.

Biology. Female ovigerous in November.

Remarks. This species was first described from

two individuals, reexamined by Rauschert and

Andres (1991). Since the description of

Pseudothaumatelson cyproides by Nicholls

(1938) it has always been doubtful that two

species existed. Barnard and Karaman (1991:

697) stressed the “strong dissimilarity” in the

sizes of gnathopods 1 and 2 (less obvious in

shapes) but there are many other similarities

between the two nominal species. However, their

localities are not close: Schellenberg's P.

patagonicum comes from the Falkland Islands

(subantarctic, 197 m), while Nicholls found

his P. cyproides in Adelie Bay, Antarctica (c. 50

m). Three additional specimens, discovered at

more than 95 m depth from subantarctic

Marion and Prince Edward Islands (Bellan-

Santini and Ledoyer, 1986; Branch et al., 1991)

were attributed to P. cyproides without

discussion. These probably refer to Pseudothau¬

matelson patagonicum.

Bellan-Santini and Ledoyer (1986) figured two

different 1-articulate mandibular palps; one fits

with the description of P. patagonicum , the other

one is sitting under the mandible body but has a

corresponding length. Their figure of maxilla 1 is

misleading (Rauschert and Andres, 1991: 227)

but reorientation of the slide (MVR CR3389)

could confirm a normally shaped maxilla 1 with

2-articulate palp.

Distribution. Falkland Islands; Prince Edward

and Marion Islands, 95-197 m.

Thaumatelsonella Rauschert and Andres

Thaumatelsonella Rauschert and Andres 1991: 230.

Type species. Thaumatelsonella kingelepha

Rauschert and Andres, 1991 (original

designation).

Diagnosis. Antenna 1 peduncular article 1 nasi-

form, accessory flagellum of 1 article. Mandibu¬

lar palp present, of 3 clearly articulated articles,

article 3 slim. Maxilla 1 palp of 2 articles. Maxilla

2 ordinary. Gnathopods land 2 subchelate, shapes

similar, sizes different. Pereopods 3-7 bases rec¬

tilinear, dactyli smooth. Pereonite 4 elongate.

Urosomites 1-3 free. Urosomite 1 with long pro¬

cess vaulting over urosomite 2 and most of telson.

Telson medium, horizontally inserted, dorsally

excavated.

Thaumatelsonella kingelepha

Rauschert and Andres

Thaumatelsonella kingelepha Rauschert and Andres,

1991: 230-235, fig. 1,2.

Material examined. Holotype. King George Island,

Antarctica, sublittoral, sponges and bryozoans, 50 m,

MNB 27271 (female 2.5 nun). Paratypes. Collected

with holotype, MNB 4582-4586 (male 2.1 mm, 2

females 2.4 and 2.6 mm).

Additional material: Near Elephant Islands, Antarc¬

tica, fine sand, 260m, ZMH K33480 (male, 1.9 mm),

ZMH K33481 (male, 2.1 mm).

Diagnosis. Antenna 1 peduncular article 1 nasi-

fornt. Mandibular palp of 3 articles. Gnathopods

1 and 2 subchelate, ratio of lengths of propodus

gnathopod 2 : 1 less than 2; bases gnathopods 1

and 2 with long, dense setae. Gnathopod 1 palmar

comer rounded, carpus distally widened. Gnatho¬

pod 2 palmar corner rounded (female) or with

small hump and little excavation (male), with 1-2

setae on anterior margin; carpus with triangular

process, densely beset with short setae. Urosomite

1 with process vaulting over urosomites 2 and 3

and two-thirds of telson. Uropod 1 rami scarcely

different, shorter than peduncle. Uropod 2 pedun¬

cle spinose, rami scarcely different, little shorter

than peduncle. Uropod 3 strong, peduncle as long

114

TRAUDL KRAPP-SCHICKEL

as each article of ramus, each about twice as long

as wide. Telson folded, excavated, fused with

urosomite 3.

Length 2.1-2.6 mm.

Distribution. King George Is and Elephant Is,

sublittoral, on Porifera, Bryozoa, sandy bottom.

50-260 m.

Biology>. Female ovigerous in February.

Thuumatelsonella cyproides

(Nicholls) comb. nov.

Figures 13, 14

Pseudothaumatelson cyproides Nicholls, 1938:

53-55, fig. 28 [non Bellan-Santini and Lcdoyer, 1986:

425^127, fig. 29. —non Branch et al„ 1991: 12].

Type material. Syntypes. Commonwealth Bay,

Antarctica, AM PI 8730 (1 male, 1 female, 2 mm).

Material examined: Cape Bird, AM P43242 (1

specimen, slide).

Diagnosis, Antenna 1 peduncular article 1 nasi-

form. Mandibular palp of 3 articles. Gnathopods

1 and 2 subchelate, propodus of gnathopod 2 less

than twice as long as gnathopod I; basis of

gnathopod 1 with many short setae; basis of

gnathopod 2 with many long setae. Gnathopod 1

palmar comer angular. Gnathopod 2 palm

rounded (female), distally scarcely excavated,

proximally straight (male), without hump. Uro¬

somite I vaulting over urosomites 2 and 3 and

most of telson. Uropod 1 rami subequal, shorter

than peduncle. Uropod 2 rami scarcely different,

longer ramus subequal to peduncle. Uropod 3

strong, robust peduncle shorter than ramus article

1, shorter than article 2; peduncle less than twice

as long as wide, ramus more than twice as long as

wide, ramus article 2 more than 3 times as long

as wide. Telson dorsally excavated, fused with

urosomite 3.

Length: 2.0-2.1 mm.

Redescription. Body smooth. Mead shorter than or

equal to pereonites 1+2. Rostrum inconspicuous.

Eyes of stenothoid shape, normal. Lateral

cephalic lobes rounded. Pereonite 4 about as long

as pereonites 1-3 together. Urosomites 1 and 2

free. Urosomite 3 broadly coalesced with telson.

Urosomite 1 with large dorsal hump vaulting over

urosomite 2 and most of telson.

Antenna 1 shorter than antenna 2, as long as

head plus pereonites 1-3 (dorsal lengths

together); peduncular articles 2 and 3 without pro¬

cess, article 1 elongate, nasiform dorsomedially;

ratio of lengths of articles 1 : 2 : 3 about 7:4:3

(visible margin dorsally, but article 1 strongly

overlapping); accessory flagellum short, broad,

uniarticulate, with 3 terminal setae; flagellum

subequal to peduncle, of 7 articles, with many

long aesthetascs. Antenna 2 with article 4 longer

than article 5; flagellum subequal to peduncle, of

10 articles.

Mandibular incisors not different, narrow and

weak; with 2 broad raker spines; small molar cusp

with 2 robust spines; mandibular palp well devel¬

oped, of 3 articles, distal one much narrower.

Lower lip inner lobes coalesced, outer ones regu¬

larly rounded. Maxilla 1 inner plate subquadran-

gular, truncate, with 1 apical seta; outer plate with

4 senate, robust spines, 1 stout simple spine and

1 thin, short and stiff spine; palp biarticulate, ratio

of lengths of articles 2:5, extending much beyond

outer plate, with spines apically and l seta sub-

apicaliy. Maxilla 2 ordinary, outer plate and inner

plate with long setae apically. Maxilliped inner

plate reaching two-thirds along ischium, with I

seta and 1 spine apically; outer plate reaching half

along merus (palp article 1); with 1 seta apically,

3 medially; palp of 4 articles; palp article 3

(propodus) narrower than article 2; propodus

distal margin and dactylus inner margin setose.

Gnathopod 1 subchelate, much shorter than

gnathopod 2. Coxa 1 reduced, subquadrate,

covered by coxa 2; basis anteriorly with many

dense, short setae; merus setose, posterodistally

rounded, about as long as triangular carpus, which

is slightly produced between merus and propodus;

propodus triangular, anterior margin slightly

convex, with 2-3 setae; posterior margin shorter

than palm, palmar corner pronounced, with 2

robust spines; ratio length : breadth about 3 : 2,

palm armed with long distal and medial spines

and setae; dactylus about half of propodus length.

Gnathopod 2 subchelate. Coxa 2 expanded,

longer than basis, anterior margin convex with

slight comer, posterior margin straight, ventrally

rounded; posteroventral margin with single, short

spine: basis anteriorly densely beset with many

long setae; ischium much shorter than merus.

which is posterodistally pointed; carpus subtrian-

gular, posterodistal comer reaching proximal

third of propodus, moderately pointed, with

dense, short setae, distoapically some longer

ones; propodus widened, hind margin regularly

rounded, with 3 long setae, palm defined by 2

strong spines and a palmar comer of >150°, no

defining hump or excavation; propodus on

anterior margin 1 long seta; ratio of lengths of

dactylus : propodus = 33-40%, dactylus on inner

margin with many setae.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

115

Figure 13. Thaumotelsonella Nicholls. Habitus ovigerous female, 2.1 mm; antenna 1’ at scale x = 0.1 mm; maxil¬

lae 1, 2, lower lip, mandible, maxilliped at scale y = 0.1 mm; antennae 1, 2, epimeral plates 1,2 at scale y = 0.05

mm.

116

TRAUDL KRAPP-SCHICKEL

Figure 14. Thaumotelsonella cyproides Nicholls. Gnathopods 1, 2 (left), urosome at scale x = 0.2 mm; gnathopods

1 2’ at scale x = 0.1 mm; pereopods 3, 4, 5, 7 at scale x = 0.4 mm.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

117

Pereopods 3 and 4 slender, subequal, bases

anteriorly with dense, short setae, dactyli about

half propodus length. Coxa 3 ratio of length to

width 3. Coxa 4 dominant, wider than long, front

margin straight, ventral margin slightly convex,

posterior margin rounded, anteriorly not deeper

than posteriorly. Pereopods 5-7 slender, ratio of

corresponding articles only slightly different, on

all bases anteriorly and posteriorly many short

setae.

Uropod 2 extending less posteriorly than uro-

pod 1. Uropod 3 extending as far as uropod 1.

Uropod 1 peduncle longer than subequal rami,

with many spines; rami lacking spines or setae.

Uropod 2 peduncle shorter than subequal rami,

without spination or pectination on rami, similar

to uropods 1 and 3. Uropod 3 peduncle sub¬

quadrate, with 1 apical spine, ramus clearly of 2

articles, robust; article 1 with 2 apical spines,

slender, about twice as long as broad; article 2

narrow, sharply pointed, smooth. Urosomite 1

very long, with large hump vaulting over uro¬

somite 2 and most of telson. Urosomite 2 reduced,

articulation not oblique as in P. patagonicum , but

vertical to dorsal prolongation of urosomite 1.

Urosomite 3 rectangular, subquadrate, fused with

broad basis of telson.

Telson spoon-like, shorter than uropod 1 rami,

about two-thirds length of urosomite 1 hump, hor¬

izontally inserted, excavated; insertion of telson

three-dimensionally thickened and fused with

urosomite 3 but not urosomite 2.

Colour. Red eyes, orange back.

Distribution. Cape Bird, Ross Sea (77°S) and

Commonwealth Bay, Terre Adelie Coast (67°S),

Antarctica, 46 m (25 fm) -130 m.

Biology. Female ovigerous in November.

Remarks. Nicholls described this species from

two specimens. His figures and text are poor and

the mouthparts unknown. The species was men¬

tioned again by Bellan-Santini and Ledoyer

(1986) and Branch et al. (1991) both referring to

the same material. The validity of this species was

often doubted (cf. P. patagonicum) but important

details were missing.

At the Australian Museum 1 was lucky to find

another specimen of this species with material of

Metopoides from Cape Bird, Antarctica. This

specimen and its geographical locality correspond

with Nicholls’ description of P. cyproides. Its

mandibular palp corresponds with that of Thau-

matelsonella and its generic position is changed

accordingly.

The differences between Thaumatelsonella

kingelepha and T. cyproides are exceedingly

small, and concern antenna 1 article 1, coxa 3,

palmar comers of gnathopods 1 and 2, and seta-

tion of bases and proportions of uropod 3 articles.

It is quite probable that these are due to body size

as the described material of T. kingelepha is 2.5

mm while that of T. cyproides is 2.1 mm. It has

been shown for Raumahara that allometry is pre¬

sent to a high degree. But before making a defi¬

nite decision on the specific distinctness of these

two taxa, one from the subantarctic and the other

from Antarctica, I await additional material.

The essential difference between Pseudothau-

matelson and Thaumatelsonella is in the structure

of the mandibular palp. Jerry Barnard discussed

“several examples of species with obvious

gnathopodal relationships now separated from

each other generically on the basis of metameric

differences in mandibular and maxillary palps

..(see Introduction). The small size of

mandibular palp article 3 in Thaumatelsonella

suggests only small steps from a 3-articulate palp

to a palp with length of article 1 + article 2, but

vanishing articulation between them. Having both

these conditions in one genus would be analogous

to having a rudimentary and absent palp as is seen

in Raumahara.

In this complex the condition of mandibular

palp of “1 article” includes species with a short

stump and other with a palp of nearly full length

but without articulation. Future descriptions

should indicate length of the palp relative to the

incisor. In Raumahara (and in the Stenothoe

monoculoides- species complex) the arrangement

of maxilia 2 plates varies from in tandem to a rid¬

ing position, probably in response to different

feeding strategies. But in Raumahara and

Stenothoe more species arc known from greater

numbers of specimens. The division of Pseu-

dothaumatelson and Thaumatelsonella is retained

until a more detailed analysis of more species

reveals the significance of a robust, long

mandibular palp with coalesced articles versus the

ordinary, 3-articulate palp or a short stump of one

article.

Yarra gen. nov.

Type species. Yarra unguiserra sp. nov.

Diagnosis. Antennae 1 and 2 geniculate. Antenna

1 peduncular article 2 a nasiform process; acces¬

sory flagellum lacking. Mandibular palp lacking.

Maxilla 1 palp of 2 articles. Maxilla 2 plates aber¬

rant, in riding position. Maxilliped inner plate

with apical knob, outer plate absent. Gnathopods

118

TRAUDL KRAPP-SCHICKEL

1 and 2 subchelate, different in size, similar in

shape. Urosomite 1 with dorsal fold, vaulting over

urosomites 2 and 3 and part of telson. Uropods 1

and 2 smooth, rami unequal. Uropod 3 robust,

ramus of 2 articles. Urosomites free. Telson hori¬

zontally inserted, spoon-like, dorsally excavated,

marginally scleritic.

Etymology. Following Jerry Barnard’s habit of

using local names, the genus is named for the

scenic Yarra River which meanders behind the

Abbotsford Annexe, Museum Victora, in whose

collections this exciting animal was found.

Yarra unguiserra sp. nov.

Figures 15-18

Material examined. Holotype. Australia, eastern Bass

Strait, 15.5 km from Point Ricardo, 37°53.I8'S,

148°28.96'E, 45 m, sand-shell, 26 Sep 1990, Smith-

Mclntyre grab (stn MSL-EG 44), NMV J45739 (sex?. 2

mm, slide).

Paratype. Australia, eastern Bass Strait, 7.3 km SSW

of Cape Conran. 37°52.67'S, 148°42.06 E, 48 m, sand-

shell, 28 Sep 1990, Smith-Mcintyre grab (stn MSL-EG

60), NMV J22638 (1 specimen in alcohol, 1.2 mm).

Additional material. Type locality. NMV J45737 (1

in alcohol). Eastern Bass Strait, 24 km NNE of Eddys-

tone Point. 40°43.9"S, 140° 32.5 E, 56 m, muddy sand,

14 Nov 1981. epibenthic sled, RV Tangaroa, NMV

J3767 (1 specimen. 1.2 mm, slide). Eastern Bass Strait,

15.5 km SW of Point Ricardo. 37°53.14S,

148°28.94'E, 45 m, medium sand, 4 Jun 1991, Smith-

Mclntyre grab (stn MSL-EG 81), NMV J45738 (1 spec¬

imen in alcohol, sex?. 1.2 mm).

Diagnosis. Antenna 1 peduncle forming a knee in

article 2. thus forcing article 3 to insert subrec-

tangularly. Antenna 2 knee between peduncular

articles 4 and 5. Maxilla 2 plates riding, each with

1 apical spine. Gnathopods 1 and 2 palmar comer

well defined, palm longer than posterior margin,

slightly concave, serrated; dactyli characteristi¬

cally serrated inside. Gnathopod 1 dactylus with 2

sharp teeth. Gnathopod 2 dactylus with 3 huge

sharp teeth, in the gaps a deep incision and a long

seta inserted. Uropods 1 and 2 smooth, peduncle

strong, rami unequal, dorsally pectinate. Uropod

3 robust, of 2 articles. Urosomites 1-3 free,

covered with external “skin", linking the dorsal

fold of urosomite 1 with uropod I insertion.

Telson horizontally inserted, spoon-like, dorsally

excavated, margins scleritic.

Length 1-2 mm.

Etymology• The specific name (unguis (L.), nail, +

serra (L.), saw) describes the morphology of the

palm and dactylus of the gnathopods (noun in

apposition).

Description. Body smooth, stout, coxae 2-4 dom¬

inating, coxae 3 and 4 incised on anteroventral

comer. Head high, narrow. Rostrum inconspicu¬

ous. Eyes normal. Lateral cephalic lobes rounded.

Pereonite 4 about same width as head + pere-

onites 1-3. Urosomites free. Urosomite 1 with

large dorsal hump vaulting over urosomites 2 and

3 and great part of telson. Urosomites 2 and 3 not

arranged horizontally, but vertically. Urosomite 3

broadly coalesced with telson.

Antenna 1 shorter than antenna 2, geniculate;

peduncular articles 1 > 2 > 3; article 1 sub¬

quadrate, 1.5 times as long as wide; article 2 dor¬

sally with nasiform process, ventrally with right

angled margin, forcing geniculate insertion of

article 3; accessory flagellum lacking; flagellum

shorter than peduncle article 1, of 5 articles, with

long aesthetascs. Antenna 2 geniculate between

peduncular articles 4 and 5; article 3 subquadrate;

article 4 on outer side of knee longer than article

5 as apical margin oblique; flagellum shorter than

peduncle, of 5 articles.

Upper lip lobes asymmetrically rounded.

Mandibular incisors on both sides well devel¬

oped: raker spines huge, bifurcate; molar cusp

well visible; palp absent. Lower lip not found.

Maxilla 1 inner plate small, w'ithout spines or

setae: outer plate with 6 robust, acute spines,

arranged not bilaterally but in a crown, not very

different from each other; palp biarticulate, length

ratio 1: 4, extending much beyond outer plate,

with 1 stout spine and 1 acute, setose spine. Max¬

illa 2 plates subequal in size but sitting in riding

position, with 0 or 1 apical small spines. Maxil-

liped inner plate scarcely reaching half way along

ischium, with 1 seta subapically, 1 knob apically;

outer plate absent; palp of 4 articles, article 3

(propodus) narrow, beset with many setae as on

dactylus.

Gnathopod 1 much smaller than gnathopod 2;

coxa 1 reduced, anteroventral ly with 1 seta; basis

medioposteriorly with 2 long, smooth setae;

merus and ischium with many short setae;

ischium not reaching propodus; propodus hind

margin about same length as slightly concave,

serrate, oblique palm; palmar comer well defined;

outer margin with group of apical, short setae, and

subapical long ones, medially with small and

short ones; dactylus with deep serration on inner

margin. Coxa 2 regularly rounded anteriorly,

much widened, with short setae along anterior

margin; basis three-quarters length of coxa 2,

with 2 long, smooth setae posteriorly; ischium to

proximal propodus posterior margin beset with

many short setae; merus and carpus each with 1

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

119

Figure 15. Yarra unguiserra gen. and sp. nov. Habitus of 1.2 mm specimen; mouthparts at scale = 0.1 mm, except

middle sketch of mandible in free scale.

120

TRAUDL KRAPP-SCHICKEL

Figure 16. Yarra unguiserra gen. sp. nov.. Antennae 1, 2, gnathopods 1, 2, pereopod 3 at scale = 0.1 mm.

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

121

Figure 17. Yana unguiserra gen. nov., sp. nov. Coxae 3, 4, pereopods 4, 5, 6, 7 at scale x = 0.05; urosome com¬

plete and epimeral plate 3 at scale y = 0.1 mm, telson + urosomite 3 + uropod 3 detail at scale y = 0.05 mm.

122

TRAUDL KRAPP-SCHICKEL

Figure 18. SEM pictures, a, b, Raumahara dertoo Barnard, urosome with shallow spoon-shaped telson. Yarn

unguiserra gen. and sp. nov. c, gnathopod 2; d. urosome with telson about as deep as wide.

long seta distoposteriorly; propodus length of

palm twice hind margin; palm strongly senate,

with 10-12 teeth, deeply incised; palmar corner

beset with 2 setae; propodus anterior margin

smooth, anterodistally with 2 short setae; dactylus

with “triple tip”, created by strong and expanded

dactylus, twice deeply incised on inner margin,

having one long seta in each notch.

Percopods 3 and 4 strikingly smooth and

simple, only scarce and very short setae on distal

end of some articles; basis, like gnathopod, poste¬

riorly with 2 setae. Pereopod 3 (in juvenile) arti¬

cles similar to pereopod 4; in adult (see figures of

habitus or detail) carpus swollen, while articles of

pereopod 4 are similarly slender. Pereopods 5—7

slender, ratios of corresponding articles similar;

length diminishing from 5 to 7.

Uropods 1-3 extending less posteriorly in suc¬

cession. Uropod I totally unarmed; peduncle

subequal to longer ramus, ratio of lengths of

rami 2 : 3; both rami dorsodistally finely pecti¬

nate. Uropod 2 similar in shape to uropod 1 but

shorter and stouter. Uropod 3 robust, with smooth

peduncle, articles of ramus much shorter and

narrower, of subequal length, article 2 dorsal ly

pectinate.

Telson spoon-like, horizontally inserted,

dorsally excavated, with 1-2 plumose setae on

distal third; more than half length fused with

urosomite 3.

Distribution. Bass Strait, Australia, 45^18 m

depth, medium sand or shell-sand.

Remarks. In Barnard and Karaman’s (1991: 684)

key to genera of Stenothoidae this species

keys out to Pseudothaumatelson in spite of hav¬

ing the antenna I article 2 strongly nasi form

(instead of article 1) and, with article 3, genicu¬

late. However, the mandibular palp is totally lack¬

ing and maxilla 2 is riding, both supposedly apo-

morphic character states shared only with

Chucullba. The most striking differentiating

characters are the much more specialized uro¬

some in Chucullba , antenna 1 article 2 thickened

(like Raumalwra, Ruukumara, Ptychotelson )

instead of article I ( Chucullba ) and the very

aberrant, deeply serrate dactylus in gnathopods I

and 2 of Yana (instead of smooth margin in all

other genera of this group). This surprising

species, placed here in a separate genus, may use

a pair of long setae on the bases of gnathopods 1

and 2 and pereopods 3 and 4 as or in addition to

THAUMATELSONINE STENOTHOID AMPHIPODA: PART 1

123

oostegites. No females with typical oostegites

were found. Coxa 4 is very broad with nearly rect¬

angular margins, like coxae 3+4 together in

Cyproideidae.

Conclusions

Having studied these — until now all called thau-

matelsonine — genera, I feel certain that Gur-

janova’s idea of separating classical stenothoid

genera from those ones with a “wonderful telson”

is still valid. In spite of Barnard’s fears (1972a:

318) there are no problems in defining a vertically

inserted, hugely elevated, rudder-like telson such

as in Antatelson, Ausatelson and Thaumatelson.

This cladc (which will be treated in Part 2 of this

series) is highly aberrant, all living in the south of

Australia (Bass Strait) and Antarctica, and all are

reported to have a similar ecology.

Less striking are those genera which Barnard

saw as bridging stenothoids and thaumatelsonids

and which are better known now. Their common

character is a telson with sclerified margins, basi¬

cally coalesced with urosomite 3. This telson can

be adjacent to the urosomc (see R. rongo Barnard,

1972b: fig. 92: R. noko Barnard, 1974: fig. 71,

and figures herein). It may also be separate from

the urosome or upright (see Rawmhara judithae

Moore, 1981: fig. 14; Pseudothaumatelson

cyproides Nicholls, 1938: fig. 28, and figures

herein), thus appearing variously boat-, shovel- or

spoon-like. In the habitus drawings of Pseit-

dothaumatelson cyproides (Nicholls, 1938) and

Thaumatelsonella kingelepha (Rauschert and

Andres, 1991), or in figures given in this

paper, the vaulted prolongation of urosomite 1 is

shown to fit in the excavation of the telson, thus

strengthening the unity of the urosome.

While the bulge on urosomite 1 is differently

developed, the three-dimensional ly inserted,

proximally thickened, folded, marginally stiff¬

ened and partly coalesced telson is always

present, thus offering a well defined synapo-

morphy. This paper illustrates different types of

telson within stenothoid genera:

1. a flat, horizontally inserted, 2-dimensional,

free telson (basic Stenothoidae);

2. a proximally thickened, three-dimensionally

inserted, basically coalesced, marginally stiffened

and dorsally excavate telson, which is described

by different authors as boat- or spoon-like (treated

here); and

3. a vertically , 2-dimensionally inserted,

hugely enlarged telson (treated in Part 2 of

thaumatelsonine species).

Acknowledgements

Hans Georg Andres (Zoological Museum, Ham¬

burg) kindly offered me Australian stenothoids to

sort (work supported by the university). This ini¬

tial study demanded a personal visit to Australia

for additional collecting and examination of

museum collections. Gary Poore (Museum

Victoria, Melbourne) who invited me to work in

the Crustacea Laboratory and John Moverley who

offered lodging, enabled my long and fruitful stay

to take place without financial support. Lively

written and spoken discussions with Sandro Ruffo

(Museo di Storia Naturale, Verona) and Wim

Vader (University and Museum, Tromso)

throughout all the stages of this paper brought

many additional stimuli. As a result of a similar

zoological passion, my husband Franz Krapp

accepted my long absence from home. Full

hearted thanks to all of them.

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Memoirs of Museum Victoria 58(1): 125-136 (2000)

REDIAGNOSIS OF THE ENDEMIC SOUTHERN AUSTRALIAN GENUS

PARASTACILLA HALE, 1924 (CRUSTACEA: ISOPODA: ARCTURIDAE) WITH

DESCRIPTIONS OF TWO NEW SPECIES

Rachael A. King

Crustacea Laboratory, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

and

Department of Zoology, University of Melbourne, Vic. 3010, Australia

(rking@museum.vic.gov.au)

Abstract

King, R.A., 2000. Rcdiagnosis of the endemic southern Australian genus Paraslacilla Hale,

1924 (Crustacea: Isopoda: Arcturidae) with descriptions of two new species. Memoirs of

Museum Victoria 58(1): 125-136.

The arcturid genus Parastacilla Hale, 1924 is rediagnosed and a key to its four species

presented. All species, P. truculenta Hale, 1924; P. bakeri Hale, 1924; P. torus sp. nov. and

P. tingara sp. nov. have limited distribution in southern Australia.

Introduction

The arcturid isopod genus Parastacilla was

erected by Hale (1924) to include two endemic

South Australian species P. truculenta and

P. bakeri. Hale distinguished the new genus on

the basis of the robust antennae, stout uniarticu-

late antenna 2 flagellum, flattened anterior

pereopods and unsegmented pleon.

Hale believed that Parastacilla resembled

Astacilla Cordiner to which he assigned some

southern Australian species; these now belong to

Neastacilla Tattersall. The species of Parasta¬

cilla share many morphological similarities with

species of Neastacilla from southern Australia:

elongation of pereonite 4, an extremely genicu¬

late body between pereonites 4 and 5 and similar

sexual morphologies with curved, short appendix

masculina and simple penial plate.

Pereopods 2 to 4 of Parastacilla are uniquely

flattened and sparsely setose. The pereopods of

Neastacilla species are elongate with closely

spaced, long setae. In Parastacilla pereopod 2

has a small dactylus and unguis and pereopods 3

and 4 lack a dactylus. Generally, in species of

Neastacilla pereopods 2 to 4 have a dactylus

and unguis. In Parastacilla the body is

stout and cylindrical while species of Neastacilla

are elongate and slender. Antenna 2 of

Parastacilla is stout with a flagellum of two

articles; the distal article is very small and

bears a claw. Antenna 2 of Neastacilla is

slender with two or three flagellar articles

of similar lengths. The fusion of the head and

pereonite I, a characteristic of the arcturid

family, is extended anteriorly and incised in

species of Parastacilla ; this extension is evident

only to a small extent in some species of

Neastacilla.

The genus was partially redescribed from the

types by Hale (1946) but has not been studied

since. Two new species have been discovered in

collections at Museum Victoria, Melbourne, and

the description of these species has uncovered

errors in Hale’s original generic diagnosis. The

types of both P. bakeri and P. truculenta have

been examined to rediagnose the genus and its

species. The new species described herein and a

survey of museum collections has extended the

known distribution of the genus from South Aus¬

tralia to Victoria, Tasmania and Western Aus¬

tralia.

Abbreviations are: NMV, Museum Victoria,

Melbourne, Australia; SAM, South Australian

Museum, Adelaide, Australia.

Arcturidae Bate and Westwood, 1868

Parastacilla Hale, 1924

Parastacilla Hale, 1924: 209-212.—Hale, 1946:

187-188.

Type species. Parastacilla truculenta Hale, 1924

(by original designation).

125

126

RACHAEL A. KING

Diagnosis. Body cylindrical and strongly genicu¬

late. Lateral margins of head and pereonite 1

extended anteriorly and separated by a lateral

incision. Antenna 2 stout; flagellum of 2 articles,

second article very small with claw and spines

medially along length. Maxillipedal palp article 4

narrower than article 3. Pereopod 1 within margin

of head; with dactylus barely longer than wide,

without unguis, with setae on the medial and

oblique distal margin. Pereopods 2 to 4 flattened

and compact, with paired sparse and widely

spaced setae; setae on ischium and carpus shorter

than the article. Pereopod 2 with very small

dactylus present on mesial surface, with unguis

(sometimes with secondary unguis). Pereopods 3

and 4 dactylus absent. Pereopods 5 to 7 progres¬

sively shorter; dactylus with primary and slightly

smaller secondary unguis (sometimes fused);

pereopod 5 basis twice length of pereopod 6

basis. Pereonite 4 up to 9 times as long as pere¬

onite 3; males with pereonite 4 as long as in

female. Pleon with dorsolateral wings. Uropodal

inner ramus with 2 setae of subequal length. Male

pleopod 1 exopod laterally notched, with 2 prox¬

imal setae of unequal length on posterior face.

Male pleopod 2 with appendix masculina short,

reaching end of endopod, curved, with ridge on

posterior face, apex simple.

Composition. Parastacilla truculenta Hale, 1924;

P. bakeri Hale, 1924; P. torus sp. nov.; P. tingara

sp. nov.

Remarks. The flagellum of the second antenna

consists of two articles, not one as described by

Hale (1924). Hale believed that all four anterior

pereopods consisted of six articles, ending in a

rounded setose propodus. Examination has

shown that in all species the second pereopod has

a small dactylus and unguis. Some specimens

have a secondary unguis on the dactylus of

pereopod 2.

Key to species of Parastacilla

1. Pereonites without large dorsal elevations. P. truculenta

Pereonites with dorsal elevations.2

2. Head without dorsal elevation; antenna 2 articles 4 and 5 with tuberculate

elevations. P. bakeri

Head with dorsal elevation; antenna 2 articles mostly smooth, may have

some tuberculation...3

3. Female pereonite 4 expanded laterally; large anteriorly directed dorsal ele¬

vations on pereonites 3 and 4; male without elevation on pereonite 3, eleva¬

tion on pereonite 4 with 2 apices... P. torus

Female pereonite 4 not expanded laterally; elevations on pereonites 3 and 4

of equal size to elevation on head. P. tingara

Parastacilla truculenta Hale

Parastacilla truculenta Hale, 1924: 210-211, fig.

1.—Hale, 1946: 187-188, fig. 15 A.

Material examined. I lolotype. South Australia. Beach-

port, 5-7 m, dredge, H.M. Hale, SAM C237 (1 female,

18.5 mm).

Other material. South Australia. Flinders Island,

“The Hotspot” reef, 5 n. miles W of Flinders Island

(33°40.80'S, 134°22.50’E), 21 m, large red algae,

SCUBA, G.C.B. Poore on FV Limnos , 20 Apr 1985 (stn

SA-69), NMV J16696 (manca 2, 6.0 mm).

Diagnosis. Female: Head with dorsal elevation.

Fusion of head and pereonite 1 indicated by a dor¬

solateral groove with the lateral margin extended

anteriorly and incised laterally. Pereonites 2 to 7

smooth. Pereonite 4 more than 9 times as long

as pereonite 3. Pereonites 5 to 7 progressively

shorter. Pleon length greater than the

combined lengths of pereonites 5 to 7; with small

dorsolateral wings.

Eyes lateral. Antenna I extending to end of

peduncle article 2 of antenna 2; uniarticulate flag¬

ellum with lateral and distal aesthetascs. Antenna

2 stout, more than half length of body; flagellum

of 2 articles and claw, with a medial row of comb¬

like spines full length.

Pereopod I dactylus barely longer than

wide, without unguis, bearing 2 medial and 3

oblique setae. Pereopod 2 with dactylus and

unguis. Pereopods 3 and 4 without dactylus.

Pereopods 5 to 7 dactlyus with unguis and

secondary unguis.

Uropodal exopod oblique, not reaching mid¬

point of endopod, with 2 distal setae of equal

length.

Distribution. South Australia; subtidal.

THE ARCTURID GENUS PARASTAC1LLA

127

Remarks. This redcscription was based on the

holotype (SAM C237). This specimen is an

immature female in which the oostegites are not

fully formed. Male specimens have not been

found. A specimen from Museum Victoria (NMV

J16696) is a manca 2 stage: a juvenile with a

developed pereopod 7 and undeveloped sexual

appendages.

Parastacilla bakeri Hale

Figure 1

Parastacilla bakeri Hale, 1924: 211-212, fig. 2.—

Hale, 1946: 187-188, fig. 15 B.

Material examined Holotype, South Australia. Marino

Reef, W.H. Baker, SAM C238 (1 male, 9.5 mm).

Diagnosis. Male: Head with small dorsal eleva¬

tion. Fusion of head and pereonite 1 indicated by

dorsolateral groove with lateral margin slightly

extended anteriorly and incised laterally. Pere¬

onite 2 smooth. Pereonite 3 with dorsal elevation.

Pereonite 4 about 9 times as long as pereonite 3

with midlength dorsal elevation of similar height

to elevation on pereonite 3. Pereonites 5 to 7 pro¬

gressively shorter, smooth. Pleon longer than

combined lengths of pereonites 5 to 7, with small

lateral expansions, small dorsal elevation at two-

thirds length and 2 dorsolateral wings.

Eyes lateral. Antenna 1 extending to end of the

peduncle article 2 of antenna 2; uniarticulate flag¬

ellum with aesthetascs attached laterally and dis-

tally. Antenna 2 stout, more than half as long as

body, tuberculate elevations with setae on dorsal

surface of peduncle articles 3-5; flagellum of 2

articles and claw, with a medial row of spines

along full length.

Pereopod 1 dactylus barely longer than wide,

without unguis, bearing 2 medial setae and 3

oblique setae on the distal margin. Pereopods 2 to

4 tuberculate. Pereopod 2 with dactylus on medial

surface, with unguis. Pereopods 3 and 4 without

dactylus. Pereopods 5 to 7 with an unguis and a

secondary unguis. Pereopods 5 and 6 dactylus

denticulate; pereopod 7 dactylus smooth.

Uropodal exopod not reaching midpoint of

endopod, with 2 distal setae of equal length.

Male pleopod 2 appendix masculina with ridge

on posterior face, apex simple, curved and short

(see Remarks).

Distribution. South Australia; subtidal.

Remarks. Hale’s original description stated that

the type of Parastacilla bakeri (SAM C238) was

‘mounted in Balsam’ and so it was only possible

to figure the whole animal and antenna 1. In 1946

Hale re-illustrated the flagellum of antenna 2 and

pereopod 1 of what he recorded was the same

specimen, apparently not mounted. The specimen

(SAM C238) redescribed here was unmounted

and undissected before being partially dissected

for this redescription. It is possible that Hale’s

(1946) illustrations were done without any dis¬

secting and mounting. 1 assume that this is the

holotype as a second specimen was never men¬

tioned. The holotype designated by Hale was a

male and is the only specimen available.

The appendix masculina was examined in situ

from the holotype but not illustrated.

Parastacilla tingara sp. nov.

Figures 2^1

Material examined. Holotype. Tasmania, Waterhouse

Point (40°49.25'S, 47°40.04'E), 5 m, seagrass: Amphi-

bolus antarctica, G. Edgar, 24 Apr 1992, NMV J39333

(1 female, 13.5 mm).

Paratype. South Australia. Edithburgh (35°05'S,

137°45'E), 3 m. red algae: Caulocystis cephalomithos,

hand collection. R.A. King, 14 Mar 1999, NMV J39334

(I juvenile, 8.0 mm).

Diagnosis. Head with dorsal elevation. Pereonite

3 with dorsal elevation, smaller than elevation on

head. Pereonite 4 with elevation, smaller than

elevation on pereonite 3. Antenna 2 with some

granulation on peduncular articles 3 and 4.

Description. Female: Anterolateral margins of

head with small tubercles, rostral point very

small. Head with large dorsal blunt elevation,

with setae extending out of elevation. Fusion of

head and pereonite 1 indicated by dorsolateral

groove with lateral margin extended slightly ante¬

riorly and incised laterally. Pereonite 2 smooth

with small tuberculate anterolateral extensions,

with some dorsal setae. Pereonite 3 with large

dorsal elevation of a similar size to elevation on

head, with dorsal setae; anterolateral extensions

small and tuberculate. Pereonite 4 about 7 times

as long as pereonite 3; with row of dorsolateral

tubercles along anterior margin and small ante¬

rolateral expansions; not markedly wider than

previous pereonites; with smaller dorsal elevation

at midlength, with dorsal setae. Pereonites 5 to 7

lateral margins not expanded but slightly tuber¬

culate, progressively shorter posteriorly, dorsal

surfaces with numerous setae. Pleon longer than

combined lengths of pereonites 5 to 7, with small

lateral expansions, a small dorsal elevation at

quarter length and 2 dorsolateral wings.

Eyes lateral. Antenna 1 extending almost to end

of peduncle article 2 of antenna 2; uniarticulate

128

RACHAEL A. KING

Figure 1. Parastacilla bakeri male holotype (SAM C238): a, lateral view; distal end of antenna 2; pereopods 1 to

7. Scales: a = 1.0 mm; b (P1-P7) = 0.5 mm.

THE ARCTURID GENUS PARASTACILLA

129

Figure 2. Parastacilla tingara female holotype (NMV J39333): a, dorsal view; b, lateral view; c, ventral view with

oostegites detailed; antennae 1 and 2. Scales: a, b = 1.0 mm; c = 1.0 mm; A1 =0.5 mm; A2 = 1.0 mm.

130

RACHAEL A. KING

Figure 3. Parastacilla tingara female (NMV J39333): mouthparts; distal end of uropod. Scales a (MP, MX1, MX2,

1MD, rMD) = 0.5 mm; b (U) = 0.5 mm.

flagellum with distal aesthetascs. Antenna 2 stout,

more than half length of body, margins with some

tuberculation; flagellum of 2 articles and claw,

with medial row of comb-like spines along full

length.

Maxilla 1 inner lobe with 3 terminal setae;

outer lobe with 10 distal robust setae. Maxilla 2

inner lobe with 16 plumose setae; middle lobe

with 4 setae; outer lobe with 3 setae. Maxillipedal

endite with 19 mesial setae; 1 coupling hook pre¬

sent; palp article 2 with mesial seta) rows; article

3 with mesial and lateral setal rows; article 4 nar¬

rower than article 3 and with mesial and lateral

setal rows; article 5 with distal setae.

Pereopod 1 included within margin of head;

propodus as long as carpus; dactylus barely

longer than wide, without unguis, with 3 medial

setae and 3 setae on the distal oblique margin.

Pereopod 2 with dactylus, unguis and secondary

unguis. Pereopods 3 and 4 without dactylus. Pcre-

opods 5 to 7 with unguis and secondary unguis

two-thirds length of primary unguis; dactylus

barely denticulate with raised setose area close to

dactylus/propodus suture; pereopod 5 basis length

twice pereopod 6 basis.

Uropodal exopod oblique, not reaching mid¬

point of endopod, with 2 setae of subequal length.

Oostegites present on pereopods 1 to 4;

oostegite 4 without suture.

Distribution. South Australia to Tasmania;

subtidal.

Etymology. "Tingara " is an Australian aboriginal

word meaning the sea.

Remarks. This species most closely resembles

Parastacilla bakeri. Males are generally only

slightly smaller than females in this genus but the

male specimen of P. bakeri is much smaller than

the female of/’, tingara (9.5 mm vs 13.5 mm) and

yet both are fully mature. Ornamentation of the

head, usually highly species specific, differs

between P. bakeri and P. tingara. Antenna 2 of

P. bakeri is more slender than antenna 2 of

P. tingara The morphology of antenna 2 is usu¬

ally extremely similar between sexes of the same

species. The scales on the antenna 2 flagellum in

P. bakeri and P. tingara are different. P. bakeri

has simple scales and P. tingara has comb-like

scales as does P. torus. Denticulation of the

THE ARCTURID GENUS PARASTACILLA

131

Figure 4. Parastacilla tingara female (NMV J39333): pereopods 1 to 7. Scale = 0.5 mm.

132

RACHAEL A. KING

dactylus of pereopods 5 to 7 also differs between

P. bakeri and P. tingara. Examination of other

species of Parastacilla leads me to believe that

there may be differences in the dcnticulation of

the dactylus of pereopods 5 to 7 between the

sexes but this variation is not as pronounced as

the differences between P. bakeri and P. tingara.

Although the male of P. tingara and female of

Parastacilla bakeri are unknown, I believe the

differences between specimens is more than can

be expected from sexual dimorphism.

Parastacilla torus sp. nov.

Figures 5-8

Material examined Holotype. Victoria. Venus Bay

(38°39.57'S, 145“42.00'E), 9 m, SCUBA, 6 Mar 1982,

NMV J16691 (1 female, 9.5 mm ).

Paratypes. Victoria. Twin Reefs, 11 m. 4 Mar 1982,

NMV J16693 (1 male. 6.0 tnm). Venus Bay, 8 m, 5 Mar

1982, NMV J16695 (2 juvenile males). Cape Paterson

(38°40.22'S, 145-36.53'E), 6 m, 5 Mar 1982 NMV

J16692 (1 juvenile). Manners Haven, 6 m. 6 Mar 1982,

NMV J16690 (2 juveniles). Nepean Bay (38"18.26'S,

144“39.57’E), 8 Apr 1998, NMV J39297 (I female, 8.5

mm; 1 immature female, 7,0 mm).

South Australia. Flinders Island (39° 52.17'S 148°

01.02'E), SCUBA, 18 m, 19 Apr 1985, NMV J16688 (1

male, 7.0nun). "Hotspot Reef’, 5 n. miles W of Flinders

Island (33°40.80'S, 134°22.50'E), 21 m, 20 Apr 1985,

NMV J16689 (1 female. 8.0 mm).

Tasmania. Pegleg Cove, Deal Island (43°56.3rs,

147°18.59'E), 8 m, 13 Apr 1983, NMV J16687 (1 male,

6.5 mm).

Western Australia. North Lumps, 2 km Off Mullaloo

(31°47.12'S, 115“43.54’E), 8 m, 2 May 1986, NMV

J39296 (1 female, 8.0 mm).

Diagnosis. Head with dorsal elevation. Pereonite

3 in female with an anteriorly directed dorsal ele¬

vation. Pereonite 4 in female with large anteriorly

directed elevation covering the entire dorsal

surface, with anterolateral expansions. Males

without the elevation on pereonite 3 or the ante¬

rolateral expansions; with a dorsal elevation on

pereonite 4 with 2 apices.

Description. Female. Anterolateral margins of

head rounded, rostral point very small. Head with

large blunt elevation. Fusion of head and pere¬

onite 1 indicated by dorsolateral groove with lat¬

eral margin extended anteriorly and incised later¬

ally. Pereonite 2 smooth, lateral margins visible in

dorsal view with small tubercles. Pereonite 3 with

large anteriorly directed dorsal elevation, half

height of elevation on head, lateral margins

visible in dorsal view with small mbercles. Pere¬

onite 4 about 8 times as long as pereonite 3;

anterolateral expansions rounded and tuberculate,

projecting around pereonite 3; markedly wider

than previous pereonites; with large dorsal ele¬

vation; small posterior lateral projections also

present. Pereonites 5 to 7 relatively smooth with

row of tubercles along each posterior dorsal

margin; progressively shorter posteriorly. Pleon

longer than combined lengths of pereonites 5 to 7;

with small anterior lateral expansions, a small

dorsal elevation and with dorsolateral wings.

Eyes lateral. Antenna 1 extending midway

along peduncle article 2 of antenna 2; small tuber¬

cles on dorsal surface of peduncle article 1; flag¬

ellum uniarticulate with aesthetascs along distal

and lateral edge. Antenna 2 stout, more than half

as long as body; flagellum of 2 articles and claw,

with medial row of comb-like spines full length.

Maxilla 1 inner lobe with 3 terminal setae;

outer lobe with 11 distal robust setae. Maxilla 2

inner lobe with 18 plumose setae; middle lobe

with 4 setae; outer lobe with 3 setae. Maxillipeda!

endite with 19 mesial setae; 2 coupling hooks

present; palp article 2 with mesial setal rows;

article 3 with mesial setal rows; article 4 narrower

than article 3, with mesial and lateral setal rows;

article 5 with distal setae.

Pereopod 1 included within margin of head;

propodus as long as carpus; dactylus barely

longer than wide, without unguis, with 3 medial

setae and 4 setae on distal oblique margin. Pereo¬

pod 2 with small dactylus with unguis and some¬

times secondary unguis (see Remarks). Pereopods

3 and 4 dactylus absent. Pereopod 5 to 7 with

unguis and secondary unguis, dactylus denticu¬

late; pereopod 5 basis length twice pereopod 6

basis.

Uropod exopod oblique, not reaching midpoint

of endopod. with 2 setae of subequal length.

Oostegites present on pereopods 1 to 4;

oostegite 4 with suture.

Male: Anterolateral lobes of head rounded, rostral

point undetected. Fusion of head and pereonite 1

indicated by dorsolateral groove with lateral

margin extended anteriorly and incised laterally.

Head with large blunt elevation. Pereonites 2 and

3 smooth, lateral margins not expanded. Pereonite

4 around 8 times length of pereonite 3, with large

forward facing dorsal elevation with 2 apices; as

wide as previous pereonites; lateral margins not

expanded. Pereonites 5 to 7 relatively smooth,

lateral margins not expanded. Pleon length

greater than combined lengths of pereonites 5 to

7, with small anterior lateral expansions, large

dorsolateral wings present.

Eyes small, subtriangular and positioned later¬

ally. Antenna 1 and antenna 2 as for female.

THE ARCTURID GENUS PARASTACILLA

133

Figure 5. Parastacilla torus, female holotype (NMV J16691): a, lateral view; b, dorsal view; e, ventral view with

oostegites detailed. Male: c, dorsal view; d, lateral view. Scale = 1.0 mm

134

RACHAEL A. KING

Figure 6. Parastacilla torus, female holotype (NMV J16691): left maxilliped, left maxillae 1 and 2, left and right

mandibles, antennae 1 and 2, distal end of uropod. Scales: a (MP, MX1, MX2,1MD, rMD, U) = 0.5 mm; b (Al) =

0.5 mm; c (A2a) = 1.0 mm; d (A2b) = 0.5 mm.

THE ARCTURID GENUS PARASTAC1LLA

135

Figure 7. Parastacilla torus, female holotype (NMV J16691): pereopods 1 to 7. Scale = 0.5 mm

136

RACHAEL A. KING

Figure 8. Parastacilla torus, male (NMV J16688): pleopods 1 and 2, penes. Scales: a (PL1, PL2) = 0.5 mm; b (Pe)

= 0.5 mm.

Mouthparts as for female. Pereopods as for

female.

Pleopod I exopod with lateral notch and 2

plumose setae of unequal lengths on posterior

face. Pleopod 2 appendix masculina with ridge on

posterior face, apex simple, curved and short.

Penes simple, straight.

Distribution. Tasmania, South Australia and

Western Australia; subtidal.

Etymology. “Torus” is Latin for round elevation

or protuberance, referring to the ornamentation of

the head.

Remarks. Two specimens were found with one

unguis on the dactylus of the second pcreopod.

Acknowledgments

1 gratefully acknowledge Dr Wolfgang Zeidler

(SAM) for access to material. Thanks to Gary

Poore, Museum Victoria, for reading the

manuscript and offering advice.

References

Bate, C.S. and Westwood, J.O., 1868. A history of the

British sessile-eved Crustacea. Vol. 2. John Van

Voorst: London. 536 pp.

Hale, I I.M.. 1924. Notes on Australian Crustacea. No.

3. Transactions of the Roval Society of South Aus¬

tralia 48: 209-225.

Hale, H.M., 1946. Isopoda — Valvifera. B.A.N.Z.

Antarctic Research Expedition. 1929-1931.

Reports-Series B (Zoology and Botany) 5:

161-212.

Memoirs of Museum Victoria 58(1): 137-148 (2000)

LEV1NEBALIA MARIA , A NEW GENUS AND NEW SPECIES OF LEPTOSTRACA

(CRUSTACEA) FROM AUSTRALIA

Genefor K. Walker-Smith

Crustacea Laboratory, Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia

and

Zoology Department, The University of Melbourne, Victoria 3010, Australia

(gwsmith@museum.vic.gov.au)

Abstract

Walker-Smith, G.K., 2000. Levinebalia maria , a new genus and new species of Leptostraca

(Crustacea) from Australia. Memoirs of Museum Victoria 58 (1): 137-148.

The genus Paranebalia historically contained three species but with the discovery of a new

species from southeastern Australia a new genus, Levinebalia , has been erected for

Paranebalia fortunata Wakabara, 1976 and L. maria sp. nov. (type species). Species of

Levinebalia differ from Paranebalia in: having the surface oftheir eyes smooth; lacking a setal

brush on the mandible incisor; having smooth pleopod peduncle margins and having crenella-

tions on the margin of pleonites 5,6, and 7. Tire margin of pleonite 5 is smooth in Paranebalia.

Of significance is antenna 2 of Levinebalia. It has several rows of small and medium-sized

spines and patches of tiny spines on the third peduncular article and flagellum. In addition,

there are conical sensory structures proximally on the third peduncle article. The conical

sensory structures and spines on antenna 2 have not been observed in any other genus of

Leptostraca.

Introduction

Historically there are six genera in the lept-

ostracan family Nebaliidae Samouelle, 1819:

Nebalia Leach, 1814; Paranebalia Claus, 1880;

Nebaliella Thiele, 1904; Dahlella Hessler, 1984;

Sarsinebalia Dahl, 1985; and Speonebalia

Bowman, Yager and lliffe, 1985. Paranebalia

was recognised by Claus (1880) as being distinct

from Nebalia, having thoracopods that extend

beyond the ventral margin of the carapace, a

greatly reduced epipod on the thoracopods and a

rostrum without a keel but with a subterminal

spine. Paranebalia contains three species,

P. longipes (Willemoes-Suhm, 1875), P. fortu¬

nata Wakabara, 1976, and P. belizensis Modlin,

1991.

Paranebalia longipes and P. belizensis are

most similar to one another having denticulate

eyes, denticles or crenellations on the pleopod

peduncles, and crenellations only on the dorsal

margin of pleonites 6 and 7. They also share the

presence of a setal brush on the mandible incisor.

The setal brush is unique to P. longipes and

P. belizensis.

Paranebalia fortunata and a new similar

Australian species do not have denticulate eyes or

crenellations on the pleopod peduncles and the

dorsal margin of pleonite 5 and pleonites 6 and 7

have very small crenellations. The new species,

here described as Levinebalia maria gen. et sp.

nov., and Paranebalia fortunata have the unique

structures on antenna 2 (Fig. 4a). Both species

have several rows of small and medium spines,

and patches of tiny spines, on the third peduncle

article and the flagellum (Figs 5c, d, e) as well

as conical structures proximally on the third

peduncular article (Figs 5a, b). These may be

chemosensory organs as vents can be seen around

the base (Fig. 5b). Wakabara (1976: 300)

observed spines on the third article of antenna 2

and the flagellum of P. fortunata and described

them as “short, strong spines”. She did not how¬

ever, mention any conical structures on antenna

2, but my examination of a paratype revealed

their existence. The conical sensory structures

and spines on antenna 2 have not been observed

in any other genus of Leptostraca. For these rea¬

sons the two species are placed in a new genus,

Levinebalia.

137

138

GENEFOR K. WALKER-SMITH

Methods

All specimens examined came from the collec¬

tions of Museum Victoria (NMV). Specimens

were dissected and mounted in glycerol and slides

were viewed under an Olympus BH-2 or BX-50

compound microscope. Whole specimens and

body parts were drawn with the aid of a camera

lucida. Plumose setae are numerous on many

body parts, but in most cases they have been fig¬

ured without their setules so as not to obscure

other details. Abbreviations used in figures are:

RO, rostrum; Al, antenna 1; A2, antenna 2; MD,

mandible; MX1, maxilla 1; MX2, maxilla 2; Tl,

T3, T8, thoracopods 1, 3, and 8; P1-P6, pleopods

1 -6; and CR caudal rami or furca. Scale bars are

1 mm.

Terminology’. 1 follow the usage of Watling

(1989) for setal classification. Spine has been

used for a non-articulating cuticular process, seta

for an articulating cuticular process and setule for

a flexible extension of the shaft of a seta. A

plumose seta is a seta with a regular row of long

setules on each side (Watling, 1989: Fig. 4g). The

comh-row consists of a row of bi-pectinate setae

(Fig. 6b) and is equivalent to the spine-row of

other authors (e.g., Dahl. 1985: Fig. 155). The

term bi-pectinate setae refers to setae that have a

comb-like row of projections along each side of

the central shaft of the seta.

Scanning electron microscopy (SEM). Leptostra-

can specimens examined under the scanning elec¬

tron microscope (SEM) were dehydrated in an

ethanol series (70%-absolute EtOH, with 10%

increments). Specimens were left in each alcohol

concentration for 30 minutes. Specimens were

then placed in a 50:50 mixture of absolute ethanol

and hexamethyldisilazane (HMDS), followed by

a 25:75 solution of absolute ethanol and HMDS,

then placed in 100% HMDS before being air-

dried. Specimens were left in each HMDS solu¬

tion for 10 minutes. Specimens were mounted on

SEM stubs with double-sided Scotch™ tape. Stubs

were sputter-coated with gold and examined

under a Phillips 505 (tungsten filament) scanning

electron microscope at 20 and 20.1 kv. The SEM

was linked to a computer and the program

Spectrum was used to capture the images.

Levinebalia gen. nov.

Type species. Levinebalia maria sp. nov.

Diagnosis. Carapace emarginate, not sculptured.

Rostrum with subterminal spine, keel absent.

Surface of eyes smooth, without denticles or

cuticular outgrowths. Eyes elongate and with

ommatidia. Antenna 1 4-articulatc, article 4 with¬

out robust setae but with a variable number of

teeth along anterior margin. Male flagellum mod¬

ified, either swollen (in juveniles) or transformed

into a callynophore. Antenna 2 with 2 rounded

cuticular outgrowths on peduncle article 3 and

conical sensory organs (Fig. 4a), surface of

peduncle article 3 and flagellum with patches of

spines (Fig. 4a), dorsal spine absent. Mandible

incisor without setal brush. Maxilla 1 palp long,

well developed. Maxilla 2 endopod uni-articulate.

Thoracopods closely spaced, long and tapering,

extending beyond ventral margin of carapace;

cpipod small, shorter than exopod; endopods

showing some articulation. Pleopods 1 -4 pedun¬

cle margins smooth. Pleopod 1 exopod with or

without comb-row of bi-pectinate setae (Fig. 6a).

Pleonites 5, 6, and 7 margins with tiny crenel la-

tions. Pleonites 5, 6, and 7 all approximately

equal in size. Pleopod 5 longer than pleopod 6.

Caudal furca short and stout.

Composition. Levinebalia maria sp. nov.;

Levinebalia fortunata (Wakabara, 1976) comb,

nov.

Etymology. Levi (Latin) means smooth, referring

to the absence of denticles on the surface of the

eyes.

Remarks. Levinebalia differs from Paranebalia in

a number of ways. Species of Paranebalia have

denticulate eyes, distinctive crenellate margins

only on pleonites 6 and 7 and crenellate pleopod

peduncles. They also have a setal brush on the

mandibular incisor. Levinebalia species have

none of these characters. The unique characters of

Levinebalia are the rows of small and medium

spines and patches of tiny spines on the third

peduncle article and flagellum of antenna 2 (Figs

5c, d, e) and the conical structures proxinially on

the third peduncle article (Figs 5a, b). These may

be chemosensory organs as vents can be seen

around the base (Fig. 5b).

Claus (1880) used the absence of a rostral keel

and the presence of a rostral spine as characters

defining Paranebalia. Both species of Levine¬

balia also have these characters. However.

Dahlella and Speonebalia also lack a rostral keel

and Sarsinebalia has a rostral spine. Their pres¬

ence in species of Paranebalia and the two

species assigned to Levinebalia suggests all are

probably related but it is likely that these charac¬

ters have arisen more than once. Indeed.

Paranebalia and Levinebalia do share three

unique synapomorphies: the elongate thoracopod;

LEVINEBALIA MARIA, NEW GENUS AND SPECIES OF LEPTOSTRACAN

139

reduced epipod; and rounded cuticular out¬

growths on article 3 of antenna 2 (Figs 4a, b).

However, the differences between Paranebalia

and Levinebalia described above, especially the

unique armature of antenna 2 which is not present

in any other Leptostraca, suggest more strongly

that the new genus Levinebalia should be erected

for Paranebalia fortunata and the new Australian

species.

Unfortunately, Wakabara (1976) did not men¬

tion the comb-row on the exopod of pleopod 1.

Her illustration (Fig. 2f) indicates only one type

of seta on the lateral margin of the pleopod 1 exo¬

pod. However my examination of a paratype of

Paranebalia fortunata revealed the presence of a

comb-row, half as long as the exopod. The finely

bi-pinnate setae are approximately equal in length

to the smooth setae found distally on the lateral

margin of the exopod. I have also examined sev¬

eral other undescribed species of Levinebalia

from the collection of Museum Victoria and all

have comb-rows on the exopod of pleopod 1.

The most recent key to extant families and

genera of Leptostraca (Martin et al., 1996) needs

to be emended and a new key to the genera of the

Leptostraca is in preparation (Walker-Smith and

Poore).

Levinebalia is represented by L, maria from

Tasmania, Australia and L. fortunata from New

Zealand. Examination of material from Museum

Victoria collections reveals Levinebalia is found

throughout Australia; in New South Wales,

Queensland, north Western Australia, South Aus¬

tralia, and Victoria but these specimens may not

belong to the new species described here.

Paranebalia has two described species; P.

longipes originally identified from the Atlantic

but also recorded from Bermuda (Willemoes-

Suhm, 1975; Sars, 1887; Verrill, 1923; Clark,

1932), Virgin Islands (Thiele, 1904), southern

Florida (Brattegard, 1970), West Indies (Thiele,

1905), Japan (Thiele, 1905), Gulf of Siam

(Thiele, 1905) and Torres Strait (Thiele, 1905);

and P. belizensis , from Belize, Central America.

The first record of Paranebalia in Australia was

by Hale (1929; 367) as P. longipes “or a closely

allied species” from the coast of South Australia.

Dahl (1985: 135) suggested Nebalia and

Paranebalia which were originally “assumed to

be composed of a small number of highly variable

species, with more or less cosmopolitan distribu¬

tion” actually contained a considerable number of

species. The existence of many undescribed

species of lcptostracans in Australian museum

collections (Dahl’s and own unpublished obser¬

vations in litt.; Walker-Smith, 1998) suggest that

records of Paranebalia longipes remote from the

type locality are dubious.

Levinebalia maria sp. nov.

Figures 1-6

Material examined. Holotype. Tasman Sea, 15 km E of

Maria 1., Tasmania (42°37'S, 148°2(rE), 102 m, WHOI

epibenthic sled, R.S. Wilson, on RV Soeia, 9 Oct 1984

(sin S05/84/1), NMV J34661 (1 female).

Paratypes. Collected with holotype. NMV J34663 (1

immature male, allotype). Australia. Tasmania, Maria

I., 5 km NE of Mistaken Cape (42°37'S, 148°12.5'E),

100 m, fine muddy bryozoan sand, WHOI epibenthic

sled, R.S. Wilson on RV Challenger, 23 Apr 1985 (stn

TAS 31), NMV J34256 (10 females and 1 male). 15 km

E of Maria I., (42°37'S. 148°20’E), 102 m, WHOI

epibenthic sled R.S. Wilson, on RV Soeia, 9 Oct 1984,

(sin SOS/84/1), NMV J34574 (19 immature males);

NMV J13282 (40 specimens; mean carapace length,

3.1940.43 mm); NMV J34573 (1 specimen; carapace

length, 3.7 mm). 20 km E of Falmouth (41°32.9 S,

148°35.0'E), 122 m, WHOI epibenthic sled, R.S. Wil¬

son, on RV Soeia, 10 Oct 1984 (stn S05/84/05). NMV

J 13283 (4 specimens; mean carapace length. 3.1540.03

mm). Off Freycinct Peninsula (42°2.20 S,

148°38.70'E), 800 m, course shelly sand, WHOI

epibenthic sled, M.F. Gomon et al. on RV Franklin, 27

Jul 1986 (stn SLOPE 45), NMV J34576 (3 specimens;

mean carapace length, 3.3740.18 mm). Off Freycinet

Peninsula (41°56.50 S, I48°37.90'E), 200 m. coarse

bryozoan sand, WHOI epibenthic sled, M.F.Gomon

M.F. et al. on RV Franklin, 27 Jul 1986 (stn SLOPE

49), NMV J34580 (4 specimens).

Description of holotype. Female (Fig. 1) with 3

embryos, entire length 4.8 mm. Carapace

length 3.0 mm, depth 2.0 mm, emarginate,

dorsally convex, anterior and posterior margin

rounded, 3.6 times length of rostrum, posterior

margin reaching pleonite 4, surface not

sculptured.

Rostrum (Fig. 1) length 2.79 times width,

greatest depth 0.23 times length, subterminal

spine present, keel absent.

Eyestalks (Fig. 1) pigmented, ommatidial

region 0.51 times length of eye, 0.69 times length

of rostrum, width 0.41 times length, dorsal

margin slightly convex, without denticles, dorsal

papilla absent, tapering, rounded distally,

supraocular scale absent.

Antenna 1 (Fig. 1) article 2 length 2.8 times

width, 9 mesiodistal plumose setae; article 3 0.64

times length of article 2, with 10 distomedial

plumose setae; article 4 medial flange with 19

teeth, each tooth with denticulate surface, numer¬

ous plumose setae over lateral and medial surface;

140

GENEFOR K. WALKER-SMITH

Figure 1. Levinebalia maria. Ferrule holotype J34661: dorsal view of rostrum; medial view of antenna 1; lateral

view of antenna 2; lateral view of mandible palp.

LEVINEBALIA MARIA , NEW GENUS AND SPECIES OF LEPTOSTRACAN

141

Figure 2. Levinebalia maria. Female holotype J34661: thoracopod 1 anterior view; thoracopod 3 posterior view;

thoracopod 8 anterior view; maxillae 1 and 2.

142

GENEFOR K. WALKER-SMITH

Figure 3. Levinebalia maria. Anterior section of male allotype J34663. Female holotype J34661: pleopods 1-4

anterior view; dorsal view of caudal furca.

LEVINEBALIA MARIA , NEW GENUS AND SPECIES OF LEPTOSTRACAN

143

swollen scale length 2.7 times width, heavily

setose, with plumose setae; flagellum with 4

articles, article 1 0.8 times length of flagellum, 4

aesthetascs anteriorly.

Antenna 2 (Figs 1, 4a, 5) article 2 length 1.52

times width, without dorsal spine; articles 3 and 4

fused, combined length 2.19 times length of

article 2, heavily setose on anterior surface, with

2 rounded cuticular outgrowths and a row of con¬

ical sensory organs, 1 row of spines, several rows

of smaller spines and patches of tiny spines on the

outer (lateral) surface of articles 3 and 4 and flag¬

ellum; flagellum with 6 articles, approximately 3

plumose setae per article.

Mandibular palp (Fig. 1) of 3 articles; article 2

with long medial seta; article 3 approximately

equal in length to article 2, margins tapering, 3

rows of plumose setae along posterior and distal

margin, increasing in length distally, terminal row

of short setae; well developed molar process,

without setal brush; mandible incisor with I

tooth.

Maxilla 1 (Fig. 2) endite 1 rounded with 1 row

of plumose marginal setae; endite 2 rectangular

with 2 rows of simple robust setae and 3 plumose

setae; palp long, well developed, with 12 lateral

setae and 2 terminal setae.

Maxilla 2 (Fig. 2) with 5 endites; endite 1

expanded distally, margin with 3 rows of plumose

setae, fine setae along proximal margin; endite 2

rectangular with 3 rows of plumose supracuticu-

lar setae; endite 3 approximately equal to endite 2,

with 2 rows of plumose setae; endite 4 0.4 times

length of endite 3, with 2 plumose setae; endite 5

same size as endite 4 with I long smooth seta;

endopod 1.16 times the length of exopod, tapering

distally, of 1 article, 3 plumose setae terminally;

exopod with plumose setae along lateral and

terminal margins.

Thoracopod endopods and exopods tapering

distally (Fig. 2); epipods reduced. Thoracopod 1,

exopod lateral and distomedial margins setose,

0.52 times length of endopod; endopod anterior

margin with 2 rows of plumose setae; epipod 0.31

times length of exopod. Thoracopod 3, exopod

lateral margin setose, 0.65 times length of endo¬

pod; endopod anterior margin with 2 rows of

plumose setae, posterior margin with several

plumose setae distally; epipod 0.3 times length of

exopod, dorsal lobe longer than ventral lobe, dor¬

sal lobe tapering distally, ventral lobe rounded,

margin with fine hair-like setae. Thoracopod 8

exopod lateral margin setose, 0.41 times length of

endopod; endopod anterior margin with 2 rows of

plumose setae, posterior margin with single row

of setae distally; epipod 0.73 times length of

exopod, dorsal lobe narrow and elongate, longer

than ventral lobe, ventral lobe distally rounded,

margin of epipod with thin hair-like setae.

Pleonites with tiny pointed denticles along

dorsal margin of pleonites 5, 6, and 7; pleonite 7

0.67 times length of telson, 1.20 times length of

pleonite 6, equal to pleonite 5, 1.20 times length

of pleonite 4.

Pleopod 1 (Figs 3, 6) exopod 0.71 times length

of peduncle, 0.67 times length of endopod, lateral

margin with comb-row of bi-pectinate setae,

comb-row 0.38 times length of exopod, 6 robust

simple setae distally along lateral margin, 2

smooth robust setae terminally, medial margin

with numerous long, fine plumose setae; endopod

with long fine plumose setae on lateral and medial

margins, 1 smooth seta and one short, stout spine

terminally.

Pleopod 2 (Fig. 3) peduncle with 5 distal

plumose setae, posterior margin not denticulate;

exopod 0.82 times length of peduncle, 0.82 times

length of endopod, lateral margin with 5 pairs of

smooth setae, medial margin with long, fine

plumose setae; endopod lateral and medial mar¬

gins with long, fine plumose setae, robust smooth

seta and stout spine terminally; reticulum present.

Pleopod 3 (Fig. 3) posterior margin of peduncle

not serrate; exopod 0.73 times length of peduncle,

0.79 times length of endopod, 6 pairs of smooth

setae, longer plumose setae between each pair, 3

terminal smooth setae, medial margin with

long, fine plumose setae; endopod lateral and

medial margins with long, fine, plumose setae,

robust smooth setae and stout spine terminally;

reticulum present above endopod.

Pleopod 4 (Fig. 3) peduncle without serrate

posterior margin; exopod approximately equal to

length of peduncle, 0.86 times length of endopod,

lateral margin with 5 pairs of simple setae, longer

plumose setae between smooth setae in each pair,

3 smooth setae terminally, medial margin with

long, fine plumose setae; endopod lateral and

medial margins with long, fine plumose setae,

smooth robust seta and stout lateral spine

terminally; reticulum present.

Pleopod 5 (Fig. 3) length 4.5 times width, with

8 simple setae terminally, fine plumose setae on

lateral margin.

Pleopod 6 (Fig. 3) length 2.5 times width, with

9 simple setae on terminal margin, fine plumose

setae on lateral margin.

Caudal furca (Fig. 3) 2.36 times long as wide,

1.44 times as long as telson, 0.17 times as long as

carapace; with 9 short, subcuticular smooth setae

144

GENEFOR K. WALKER-SMITH

Figure 4. Antenna 2 in situ (left), peduncle article 3 and first article of flagellum, a, Levinebalia maria paratype

J13282, note rows of small, spines. Arrows point to conical sensor organs and large cuticular projections, b,

Paranebalia sp. J13278, note absence of small, spines. Arrows indicate large cuticular projections.

LEVINEBALIA MARIA , NEW GENUS AND SPECIES OF LEPTOSTRACAN

145

Figure 5. a-c, Levinebalia maria paratype J13282. a, antenna 2 (left) in situ. Arrows indicate conical sensory organs

and large cuticular outgrowths, b, conical sensory organ, c, spines on article 3 of antenna 2. d-e, Levinebalia maria

paratype, J34256. d, antenna 2 (right) in situ (proximal end of antenna at bottom of photo), e, area of spines,

including patch of tiny spines.

146

GENEFOR K. WALKER-SMITH

Figure 6. a b, Levinebalia maria paratype J13282. a, pleopod I, exopod and endopod. Exopod with two types of

marginal setae, b, proximal marginal setae, forming the comb-row (arrow in a).

LEVINEBALIA MARIA, NEW GENUS AND SPECIES OF LEPTOSTRACAN

147

on lateral margin; numerous plumose setae on

medial margin and terminally, 4 denticulate setae

terminally.

Description of allotype. Juvenile male. Entire

length 4.41 mm. Carapace length 2.84 mm, depth

1.85 mm. Antenna 1 (Fig. 3) flagellum swollen,

articles fused, numerous aesthctascs, article 4 as

for female. Antenna 2 with small cuticular

outgrowth on anteroproximal margin. All other

characters same as for female.

Etymology. For Maria Island, Tasmania, type

locality (noun in apposition).

Remarks. The posterior margin of the carapace of

Levinebalia maria is rounded. Wakabara (1976)

illustrated the carapace o f Paranebalia fortunata

with a straight posterior margin but my examina¬

tion of a paratype of P. fortunata reveals the cara¬

pace is a similar shape to that of Levinebalia

maria. The lateral margin of the exopod of pleo-

pod 2 of L. maria possesses pairs of smooth setae

and this is the setal arrangement found in P. for¬

tunata (paratype examined). Wakabara's (1976)

illustration ofpleopod 2 suggested the setae occur

singly along the lateral margin. No fully mature

male of L. maria has been found.

Acknowledgments

l thank Dr Gary Poore, Museum Victoria, and an

anonymous reviewer for their helpful comments

and criticisms of the draft manuscript. 1 also thank

Keith Probert for the loan of the paratype from

Portobello Marine Laboratory, Dunedin, New

Zealand.

References

Bowman, T.E., Yager, J., and Iliffe, T.M., 1985. Spe-

onebaiia cannoni, n. gen., n. sp., from the Caicos

Islands, the first hypogean leptostracan (Nebali-

acea: Nebaliidae). Proceedings of the Biological

Society of Washington 98(2): 439-446.

Brattegard, T., 1970. Marine biological investigations

in the Bahamas 13. Leptostraca from shallow water

in the Bahamas and southern Florida. Sarsia

44:1-7.

Clark, A.E., 1932. Nebaliella caboti n. sp„ with obser¬

vation on other Nebaliacea. Transac tions of the

Royal Society of Canada 26(5): 217-235.

Claus, C., 1880. Grundzuge der Zoologie, Vol 2. 4th

edn. [not seen]

Dahl, E., 1985. Crustacea Leptostraca, principles of

taxonomy and a revision of European shelf species.

Sarsia 70: 135-165.

Hale, H.M., 1929. The crustaceans of South Australia.

fart 2. South Australian Government Printer

Adelaide. Pp. 202-380.

Hessler, R.R., 1984. Dahlella caldariensis , new genus

new species: a leptostracan (Crustacea, Malacos-

traca) from deep-sea hydrothermal vents. Journal

of Crustacean Biology 4: 655-664.

Leach, W.L., 1814. Nebalia. The Zoological

Miscellany, I: 99 100.

Martin. J.W.. Vetter, E.W. and Cash-Clark, C.E., 1996

Description, external morphology, and natural his¬

tory observ ations of Nebalia hessleri, new species

(Phyllocarida: Leptostraca), from southern Califor¬

nia, with a key to the extant families and genera of

the Leptostraca. Journal of Crustacean Biology

16(2): 347-372. **

Modlin, R.. 1991. Paranebalia belizensis , a new

species from shallow waters off Belize, Central

America (Crustacea: Malacostraca: Lepto¬

straca). Proceedings of the Biological Society of

Washington. 104(3): 603-612.

Sars, G.O., 1887. Report on the Phyllocarida collected

by H.M.S. Challenger during the years 1873-76.

Report on the Scientific Results of the Voyage of

H.M.S. Challenger during the years 1873-76.

Zoology 19: 1—38.

Thiele, J, 1904. Die Leptostraken. Wissenchaftliche

Ergebnisse der Deutschen Tiefsee-Expedition auf

dem Dampfer "Valdiva", 1898 1899 8: 1-26.

Thiele. J, 1905. Ober die Leptostraken der Deutschen

Siidpolar-Expedition 1901-1903. Deutsche

Siidpolar-Expedilion 1901-1903. 9 (Zoology) L

59-68.

Verrill, A.E., 1923. Crustacea of Bermuda: Schizopoda,

Cumacea, Stomatopoda and Phyllocarida. Trans¬

actions of the Connecticut Academy of Arts and

Sciences 6: 204—211.

Walker-Smith. G.K., 1998. A review of Nebaliella

(Crustacea: Leptostraca) with description of a new

species from the continental slope of southeastern

Australia. Memoirs of the Museum of Victoria 57:

39-56.

Wakabara, Y., 1976. Paranebalia fortunata n. sp. from

New Zealand (Crustacea, Leptostraca, Nebali¬

acea ). Journal of the Roval Society of New Zealand

6(3): 297-300.

Watling, L., 1989. A classification system for crus¬

tacean setae based on the homology concept. In:

B.E. Felgenhauer, A.B. Thistle and L. Watling

(eds), Functional morphology of feeding and

grooming in Crustacea. Crustacean Issues 6:

15-26.

Willemoes-Suhm. R.. 1875. On some Atlantic Crus¬

tacea from the 'Challenger' Expedition. Transac¬

tions of the Linnean Society of London, Series 2,

Zoology 1: 23-59.

Memoirs of Museum Victoria 58(1): 149-178 (2000)

REVISION OF THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION

(CRUSTACEA: AMPHIPODA) WITH DESCRIPTION OF FOUR NEW SPECIES

J0RGEN BERGE 1 AND WlM VADER 2

Department of Zoology, Tromso Museum, University of Tromso, 9037 Tromso, Norway

('joergenb@imv.uit.no; 2 wi m@imv.uit.no)

Abstract

Berge, J. and Vader, W., 2000. Revision of the stegocephalid genera Phippsia and Tetradeion

(Crustacea: Amphipoda) with description of four new species. Memoirs of Museum Victoria

58(1): 149-178.

The stegocephalid (Crustacea: Amphipoda) genus Phippsia Stebbing, 1906 is revised to

include six species, of which three are new to science. One new species of the genus

Tetradeion Stebbing, 1899 is also described. A key to all species of the two genera is

presented.

Contents

Introduction...149

Key to species of Phippsia and Tetradeion .150

Phippsia Stebbing, 1906 . 150

P. gihbosa (Sars, 1883) .151

P. angustipalpa sp. nov.151

P. dampieri sp. nov.155

P. roemeri Schellenberg, 1925 . 158

P. unihamata sp. nov.158

P. vanhoeffeni (Schellenberg, 1926) . 166

Tetradeion Stebbing, 1899 . 170

T. crassum (Chilton, 1883) .171

T. quatro sp. nov.172

Acknowledgements .177

References.177

Introduction

In 1883, Sars described Aspidopleums gibbosus

(the name was later changed to Phippsia

gibbosa by Stebbing in 1906) from the west coast

of Norway, and in 1925, Schellenberg described

P. roemeri. Both species have a strictly northern

distribution, found only in the North Atlantic and

the Arctic (Berge and Vader, 1997). From the lit¬

toral zone in New Zealand, Cyproidea crassa

Chilton, 1883 was placed in Tetradeion by

Chilton (1924), and redescribed by Hurley (1955)

and Barnard (1972). Although the latter species is

clearly distinct from both P. gibbosa and P. roe¬

meri, all three species do share some striking sim¬

ilarities that appear to have been overlooked in

the literature. Pending a total revision and a phy¬

logenetic analysis of the family (Berge and

Vader, in prep.), the two genera are kept separate

to minimise changes in classification. However, a

preliminary cladistic analysis indicates that the

eight species treated herein do form a mono-

phyletic group but that their relationships are still

unclear.

In the present paper, one new species of

Tetradeion is described. The number of species in

Phippsia is increased from two to six. The present

study is primarily based on material from

Museum Victoria, Melbourne, Australia (NMV)

but additional material also comes from the Nat¬

ural History Museum, London, UK (NHM).

Museum fur Naturkunde, Berlin, Germany

(ZMB), and Darling Marine Center, University of

Maine, Walpole, USA (DMC).

All dissected appendages were mounted in

polyvinyl-lactophenol, and stained with rose-

bengal. Figures of these appendages were made

using a Leica compound microscope. Mature and

149

150

J0RGEN BERGE AND W1M VADER

immature females were distinguished from males

by the presence of oostegites. The classification

of setae follows that of Berge (in press a, b).

Scales attached are all 0.1 mm. Symbols used in

the figures are as follows: Al. A2: antennae 1,2;

EP3: epirneral plate 3; L: labium: LBR: labrum;

LMND: left mandible; M: male; MX1, MX 2:

maxillae 1, 2; MXP: maxilliped; P1-P7: pere-

opods 1-7; PLP: palp; RMND: right mandible;

ST: setal-teeth; T: telson; U1-U3: uropods 1-3.

Key to species of Phippsia and Tetradeion

1. Pereopod 7 with number of articles reduced. Tetradeion .. 2

Pereopod 7 with all articles present. Phippsia .. 3

2. Pereopod 7 with 2 articles (plus coxa). Tetradeion crassum

Pereopod 7 with 4 articles (plus coxa). Tetradeion quatro

3. Metasome segment 3 dorsaily smooth.4

Mctasome segment 3 dorsaily produced. Phippsia gibbosa

4. Telson entire.5

Telson cleft.6

5. Maxilliped palp article 2 rectangular (not produced distally), labrum

elongate and triangular (clearly longer than broad). Phippsia angnstipalpa

Maxilliped palp article 2 distally produced, labrum not longer than broad,

rounded. Phippsia dampieri

6. Epirneral plate 3 distally without serrations. Phippsia roemeri

Epirneral plate 3 distally serrate.7

7. Antenna 2 peduncle article 3 shorter than broad, epistomal plate absent.

. Phippsia vunhoeffeni

Antenna 2 peduncle article 3 elongate, epistomal plate large.

. Phippsia unihamata

Phippsia Stebbing

Aspidopleurus Sars, 1895: 203 (homonym, Pisces)

Phippsia Stebbing, 1906: 89 (replacement name)

Type species. Stegocephalus gibbosus Sars, 1883

(original designation).

Species. Phippsia angnstipalpa sp. nov., P.

dampieri sp. nov., P. gibbosa (Sars, 1883), P.

roemeri Schellenberg, 1925, P. unihamata sp.

nov. and P. vanhoeffeni (Schellenberg, 1926).

Distribution. North Atlantic and Arctic Oceans

(P. gibbosa and P. roemeri), Antarctic (P. uni¬

hamata and P. vanhoeffeni), and Australia (P.

angustipalpa, P. dampieri and P. vanhoeffeni).

Remarks. Sars (1895) erected Aspidopleurus as a

monotypic genus for Stegocephalus gibbosus

mainly based on the morphology of the mouth-

parts. Phippsia gibbosa has the second article of

the maxilliped palp distally produced, and its

inner plate is long and rectangular. Furthermore,

the setae on the outer plate of maxilla 2 have dou¬

ble hooks distally (referred to in the descriptions

as a distal cleft, in addition to the hooks, see MX2

in Fig. 6), and the palp of maxilla 1 is reduced. All

these characters, together with the conspicuous

epistomal plate are characters that have, until

present, separated the two northern species of the

genus from all other known stegocephalid

species.

However, with the inclusion of Stegocephalop-

sis vanhoeffeni (Schellenberg, 1926) and the three

new species, the genus no longer appears so well

separated from other genera in the family. None

of the last mentioned four species possesses the

“doubled hooked setae” on maxilla 2, and all

other character states described above vary

between the different species. Furthermore, the

accessory flagellum of antenna 1 varies between

well developed (e.g., type species) and rudimen¬

tary (e.g., P. dampieri), and the telson varies

between 80% cleft (e.g., P. vanhoeffeni) and

entire (e.g., P. angustipalpa). On the other hand,

the labrum is partly fused with the epistome in all

six species, and the arrangements of different

maxillipedal setal-groups (Berge, in press a) indi¬

cate a strong relationship between the taxa, and

between Phippsia and Tetradeion. Most species

of the two genera possess an elongate and genic¬

ulate peduncle of antenna 2. The only character

that separates Phippsia (as treated herein) from

Tetradeion is the lower number of articles on

pereopod 7. Otherwise, all characters discussed

above are also (variably) present in Tetradeion.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

151

Thus, the genus appears a weakly defined

group kept together mainly by a large coxa 4,

short and stout antennae and narrow basis on

pereopod 6. As a revision of the family and

a cladistic analysis is in preparation by us,

phylogenetic relationships will not be discussed

further.

Stegoceplialopsis pacifica (Bulycheva, 1952)

probably also belongs in Phippsia, but due to

the lack of material and adequate descriptions,

S. pacifica is not transferred. Stegoceplialopsis

pacifica was originally described in Phippsia but

later transferred to Stegoceplialopsis by Gur-

janova (1962). The maxillipcd and maxilla 1

resemble those of both P. angustipalpa and P.

vanhoeffeni. Furthermore, the mouthparts seem to

be elongate and the species has strong affinities to

P. angustipalpa.

Phippsia gihhosa (Sars)

Stegocephalus gibbosus Sars, 1883.

Aspidopleurus gibbosus .— Sars, 1895: 203.

Phippsia gibbosa. —Stebbing, 1906: 89.—Berge et

al., in press.

Remarks. The species is figured and described by

Berge et al. (in press).

Phippsia angustipalpa sp. nov.

Figures 1,2

MateriaI examined. Holotype. Australia, Tasmania,

eastern Bass Strait, 100 km NE of North Point, Flinders

Island (38°52.6'S. 148 C, 25.2 E). 130-52 m, fine sand,

R. Wilson on RV Tangaroa , 15 Nov 1981 (stn BSS

170), NMV J45336 (immature female, 3 mm).

Paratypes. Collected with the holotype. NMV J47018

(4 immature specimens).

Description. Rostrum very small.

Antennae short. Antenna I shorter than antenna

2; flagellum 4-articulate; accessory flagellum

rudimentary. Antenna 2 peduncle (articles 3-5 )

longer than flagellum; article 3 elongate, articles

3 and 4 geniculate; article 4 longer than article 5.

Epistome curved (convex) and smooth.

Epistomal plate (medial keel) produced, but

small.

Mouthparts elongate; pointed and narrow.

Mandible incisor lateral; toothed; left lacinia

mobilis powerful, laterally expanded. Maxilla 1

palp 1-articulate; rectangular; apex not reaching

above the apex of outer plate; outer plate distally

rectangular; ST in a pseudocrown; ST first row

with 6 setae (ST 1-5, ST 7); ST 1 ordinary (simi¬

lar to ST 2-4); gap between ST 5 and ST 7 pre¬

sent; ST A present; located distally, part of first

row; ST B and C present; part of second row;

inner plate with a well developed shoulder; setae

pappocuspidate. Maxilla 2 gaping and geniculate;

outer plate setae with distal hooks present; distal

cleft absent; inner plate setal row A covering the

entire margin; appressed to row B; row A setae

pappopectinate; row B setae proximally pappose;

distally with cusps present; row C present; row D

absent.

Maxilliped palp 4-articulate; article 2 distally

unproduced; dactylus distally simple (pointed);

inner plate not exceeding base of palp article 4; 2

nodular setae; medial setal-row present; not

reduced; vertical; setae pectinate; distal setal-row

present; inner setal-row present; row reduced to 1

or 2 setae; outer plate outer setal-row present;

submarginal; setae attached normally; setae long;

strongly curved upwards (hooks); inner setal-row

absent; distal setal-group present; setae attached

in a deep hollow; setae short simple. Labrum

elongate; lobes symmetrical; right lobe reduced;

left lobe reduced. Labium distally narrowing.

Coxal plates and basis on pereopods smooth.

Coxae 1 -3 contiguous. Pereopod 1 coxal plate not

as deep as basis; propodus subovate. Pereopod 2

general appearance like pereopod I; ischium not

elongate, ratio length:breadth not exceeding 1.5;

distal posterior margin plumose setae present;

propodus subovate; palm absent. Pereopod 4 coxa

posteroventral lobe very large, reaching beyond

the base of the perconite 7; basis anterior margin

with long setae absent; posterior margin with long

setae present; plumose setae on distal anterior and

posterior margins present; ischium plumose setae

on posterior distal margin present. Pereopod 6

basis posteriorly unexpanded; with a row of long

plumose setae present. Pereopod 7 basis anterior

margin straight; distally rounded; medial row of

setae present; setae short and robust.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod I peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus as long as

inner. Uropod 3 peduncle longer than half length

of rami; outer ramus 2-articulate, longer than

inner.

Telson longer than broad; longer than peduncle

uropod 3; submarginal setae on apex absent;

entire; apically rounded.

Males: Unknown.

Etymology. Angustipalpa describes the un¬

produced article 2 on the palp of the maxilliped,

152

J0RGEN BERGE AND WIM VADER

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

153

Figure 2. Phippsia angustipalpa sp. nov. Holotype.

154

J0RGEN BERGE AND WIM VADER

a character state not found in any other species of

the genus.

Distribution. Australia, Bass Strait (known only

from the type locality).

Remarks. This is a not a typical member of the

genus in that the palp of the maxilliped is slender

(hence the name) and the mouthparts are elongate

and thus are more similar to those of both

Tetradeion crassum (Chilton, 1883) and a new

species of Stegocephalina (Berge, in press b).

Based on arrangement of the different setal-

groups on the maxilliped, rudimentary accessory

flagellum, rounded and entire telson, and genicu-

Iated peduncle (between articles 3 and 4) on

antenna 2, the species is assigned for the moment

to Pltippsia.

Phippsia dampieri sp. nov.

Figures 3-5

Material examined. Holotype. Australia, Western Aus¬

tralia. Northwest Shelf, between Port Hedland and

Dampier (18°41 S, I18°39'E), 134 m, muddy sand,

WIIOI epibenthic sled, G.C.B. Poore and H.M. Lew

Ton on RV Soela. 4 Jun 1983 (stn NWA 21), NMV

J470I9 (female, 8 mm).

Paratypes. Collected with holotype, NMV J47019 (7

specimens, male and females).

Description. Rostrum very small.

Antennae short. Antenna 1 shorter than antenna

2; flagellum 6-articulate: accessory flagellum

rudimentary. Antenna 2 peduncle (articles 3-5 )

longer than flagellum; article 3 elongate, articles

3 and 4 geniculate; article 4 longer than article 5.

Epistome curved (convex) and smooth; epis-

tomal plate produced into a large conspicuous

medial keel.

Mouthparts not elongate or pointed. Mandible

incisor lateral; toothed; left lacinia mobilis

powerful, laterally expanded. Maxilla 1 palp 2-

articuiate; rectangular; apex not reaching above

the apex of outer plate; outer plate distally rectan¬

gular; ST in a pseudocrown; ST first row with 6

setae (ST 1-5, ST 7); ST 6 absent; gap between

ST 5 and ST 7 present; ST A present; located dis¬

tally, part of first row; ST B present; part of

second row; ST C present; inner plate with a well

developed shoulder; setae pappocuspidate. Max¬

illa 2 gaping and geniculate; outer plate setae with

distal hooks present; distal cleft absent; inner

plate setal row A covering the entire margin;

appressed to row B; row A setae pappopectinate;

row B setae proximally pappose; distally with

cusps present; row C present; row D absent. Max¬

illiped palp 4-articulate; article 2 distally conspic¬

uously produced; dactylus distally simple

(pointed); inner plate not exceeding base of p a |p

article 3; 3 nodular setae; medial setal-row p re _

sent; not reduced; vertical; setae pectinate; distal

setal-row present; inner setal-row present; row

reduced to 1 or 2 setae; outer plate outer setal-row

present; submarginal; setae attached normally-

setae long robust; straight; inner setal-row pre¬

sent; well developed; setae long robust; pappo Se -

proximally parallel to outer, distally transverse;

distal setal-group present: setae attached in a deep

hollow; setae long robust. Labrum about as long

as broad; lobes asymmetrical; right lobe not

reduced; left lobe reduced. Labium distally

narrowing.

Coxal plates and basis on pereopods covered

with setae; setae very short. Coxae 1-3 contigu¬

ous. Pereopod 1 coxal plate not as deep as basis;

propodus subovate. Pereopod 2 general appear¬

ance like pereopod 1; ischium not elongate, ratio

length:breadth not exceeding 1.5; ischium distal

posterior margin plumose setae present; propodus

subovate; palm absent. Pereopod 4 coxa p os _

teroventral lobe very large, reaching beyond the

base of the pereonite 7; basis anterior margin with

long setae absent; posterior margin with long

setae present; plumose setae on distal anterior and

posterior margins present; ischium plumose setae

on posterior distal margin present. Pereopod 6

basis posteriorly unexpanded; with a row of long

plumose setae present. Pereopod 7 basis anterior

margin straight; distally rounded; medial row of

setae present; setae short and robust.

Oostegites on pereopods 2-5, gills 0 n

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod 1 peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus longer

than inner. Uropod 3 peduncle longer than half

length of rami; outer ramus 2-articulate, longer

than inner.

Telson longer than broad; longer than peduncle

uropod 3; submarginal setae on apex present;

entire; apically rounded.

Males: Pereopod 2 propodus larger in males

than in females; Urosome ordinary (similar to

females).

Etymology’. Named after the British pirate and

explorer William Dampier who visited this shore.

Distribution. Northwestern Australia. Known

only from type locality.

Remarks. Phippsia dampieri is, to the authors'

knowledge, the first stegocephalid reported from

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

155

Figure 3. Phippsia dampieri sp. nov. Paratype male, 5 mm, except habitus: holotype female, 8 mm.

156

J0RGEN BERGE AND WIM VADER

Figure 4. Phippsia dampieri sp. nov. Paratype male.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

157

Figure 5. Phippsia dampieri sp. nov. Paratype male.

158

J0RGEN BERGE AND WIM VADER

the west coast of Australia or the eastern Indian

Ocean. There are records of the family from the

western and northern Indian Ocean and from

Papua New Guinea. The species is separated

from its congeners by the combination of a entire

telson and a distally produced maxilliped palp

article 2.

Phippsia roemeri Schellenberg

Figures 6-8

Phippsia roemeri Schellenberg, 1925: 197.—

Stephensen, 1925: 133.

Material examined. About 200 specimens from BIO-

FAR and BIOICE programmes (Berge and Vader,

1997).

Description. Rostrum very small.

Antennae short. Antenna 1 as long as antenna

2; flagellum 6-articulate; accessory flagellum

well developed, article 2 absent. Antenna 2

peduncle ( articles 3-5 ) as long as flagellum;

article 3 short, about as long as broad; article 4

shorter than article 5.

Epistome curved (convex) and smooth; epis-

tomal plate produced into a large conspicuous

medial keel.

Mouthparts not elongate or pointed. Mandible

incisor lateral; toothed; left lacinia mobilis

powerful, laterally expanded. Maxilla 1 palp 2-

articulate; rectangular; apex not reaching above

the apex of outer plate; outer plate distally rectan¬

gular; ST in a pseudocrown; ST first row with 6

setae (ST 1-5, ST7); ST 6 absent; gap between ST

5 and ST 7 absent; ST A present; located distally,

part of first row; ST B present; part of second

row; ST C present; inner plate with a well devel¬

oped shoulder; setae pappocuspidate. Maxilla 2

gaping and geniculate; outer plate setae with dis¬

tal hooks present; distal cleft present; inner plate

setal row A covering about two-thirds of the mar¬

gin; appressed to row B; row A setae pappopecti-

nate; row B setae proximally pappose; distally

with cusps present; row C present; row D present;

expanded, row elongated towards and beyond

row A; with many small cusps distally. Maxil¬

liped palp 4-articulate; article 2 distally conspicu¬

ously produced; dactylus distally simple

(pointed); inner plate not exceeding base of palp

article 4; 4 nodular setae; medial setal-row pre¬

sent; not reduced; vertical; setae pectinate; distal

setal-row present; inner setal-row present; row

reduced to 1 or 2 setae; outer plate outer setal-row

present; submarginal; setae attached normally;

setae long; straight; inner setal-row present; well

developed; setae long robust; pappose; proxi¬

mally parallel to outer, distally transverse; distal

setal-group present; setae attached in a deep

hollow; setae short simple. Labrum about as long

as broad; lobes asymmetrical; right lobe not

reduced; left lobe reduced. Labium distally

narrowing.

Coxal plates and basis on pereopods smooth

Coxae 1-3 contiguous. Pereopod 1 coxal plate not

as deep as basis; propodus subovate. Pereopod 2

longer and thinner than pereopod 1; ischium not

elongate, ratio length:breadth not exceeding 1 .S-

ischium distal posterior margin plumose selu<5

present; propodus subrectangular; palm absent

Pereopod 4 coxa posteroventral lobe very large

reaching beyond the base of the perconite 7; basis

anterior margin with long setae absent; posterio r

margin with long setae present; plumose setae on

distal anterior and posterior margins absent-

ischium plumose setae on posterior distal margin

present. Pereopod 6 basis posteriorly unex-

panded; with a row of long plumose setae present

Pereopod 7 basis anterior margin straight; distally

rounded; medial row of setae absent.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod 1 peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus longer

than inner. Uropod 3 peduncle longer than half

length of rami; outer ramus 2-articulate, longer

than inner.

Telson longer than broad; as long as peduncle

uropod 3; submarginal setae on apex of each lobe

absent; cleft; apically rounded.

Males: Pereopod 2 propodus equally sized in

males and females. Urosome ordinary (similar to

females).

Distribution. An abundant species in North

Atlantic and Arctic Oceans that appears to be

limited to cold waters (Berge and Vader, 1997),

Remarks. Phippsia roemeri appears to be close to

the type species both taxonomically and zoogeo-

graphically. The two species possess a similar

maxilliped and maxilla 2, in addition to the non-

geniculate peduncle of antenna 2 and a cleft

telson. Furthermore, they are both endemic to the

North Atlantic (and subarctic), and are the only

species from the Northern Hemisphere.

Phippsia unihamata sp. nov.

Figures 9-12

Material examined. Holotype. Weddell Sea, Antarctica

(72°25.40'S, 16°26.63'W), 198-260 m, ZMB A 25

(female, 7 mm).

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

159

Figure 6. Phippsia roemeri Schellenberg, 1925. BIOFAR stn 172, female.

160

J0RGEN BERGE AND WIM VADER

Figure 7. Phippsia roemeri Schellenberg, 1925. BIOFAR stn 172, female.

THE STEGOCEPHALID GENERA PH1PPSIA AND TETRADEION (AMPHIPODA)

161

Figure 8. Phippsia roemeri Schellenberg, 1925. BIOFAR stn 172, female.

162

J0RGEN BERGE AND WIM VADER

Figure 9. Phippsia unihamata sp. nov. Holotype.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

163

Figure 10. Phippsia unihamata sp. nov. Holotype.

164

J0RGEN BERGE AND WIM VADER

Figure 11. Phippsia unihamata sp. nov. Holotype.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

165

Figure 12. Phippsia unihamata sp. nov. Holotype.

166

J0RGEN BERGE AND WIM VADER

Additional material. Weddell Sea, Antarctica

(72°27.28'S, 17°32.94'W), 455-473 m, ZMB A 111/41

(female, 6 mm); Weddell Sea, Antarctica (72°53.10'S,

19°29.33 W), 421-429 m, ZMB A 111/44 (immature, 3

mm); Antarctica (66°S, 49°E), NHM unregistered

(female, 6 mm).

Description. Rostrum very small.

Antennae short. Antenna 1 shorter than antenna

2; flagellum 6-articulate; accessory flagellum

rudimentary. Antenna 2 peduncle ( articles 3-5 )

shorter than flagellum; article 3 elongate, articles

3 and 4 geniculate; article 4 longer than article 5.

Epistome curved (convex) and smooth; epis-

tomal plate produced into a large conspicuous

medial keel.

Mouthparts not elongate or pointed. Mandible

incisor lateral; toothed; left lacinia mobilis

powerful, laterally expanded. Maxilla 1 palp 2-

articulate; rectangular; apex not reaching above

the apex of outer plate; outer plate distally rectan¬

gular; ST in a pseudocrovvn; ST first row with 6

setae (ST 1-5, ST 7); ST 6 absent; gap between

ST 5 and ST 7 present; ST A present; located

distally, part of first row; ST B present: part of

second row; ST C present; inner plate with a well

developed shoulder; setae pappocuspidate.

Maxilla 2 gaping and geniculate; outer plate

setae with distal hooks present; distal cleft absent;

inner plate setal row A covering about two-thirds

of the margin; appressed to row B; row A setae

pappopectinate; row B setae proximally pappose;

distally with cusps present; row C present; row D

present; expanded, row elongated towards and

beyond row A; with many small cusps distally.

Maxilliped palp 4-articulate; article 2 distally pro¬

duced; distal inner margin greatly produced;

dactylus distally simple (pointed); inner plate not

exceeding base of palp article 3; nodular setae 4;

medial setal-row present; not reduced; vertical;

setae pectinate; distal setal-row present; inner

setal-row absent; outer plate outer setal-row pre¬

sent; submarginal; setae attached normally; setae

long; straight; inner setal-row present; well

developed; setae long robust; pappose; proxi¬

mally parallel to outer, distally transverse;

distal setal-group present; setae attached in

a deep hollow; setae short simple. Labrum

about as long as broad; lobes symmetrical; right

lobe reduced; left lobe reduced. Labium distally

narrowing.

Coxal plates and basis on pereopods covered

with setae; setae very short. Coxae 1-3 contigu¬

ous. Pereopod 1 coxal plate not as deep as basis;

propodus subovate. Pereopod 2 longer and thin¬

ner than pereopod 1; ischium not elongate, ratio

length:breadth not exceeding 1.5; ischium distal

posterior margin plumose setae present; propodus

subovate; palm absent.

Pereopod 4 coxa posteroventral lobe very large,

reaching beyond the base of pereonite 7; basis

anterior margin with long setae absent; posterior

margin with long setae present; plumose setae on

distal anterior and posterior margins present;

ischium plumose setae on posterior distal margin

present. Pereopod 6 basis posteriorly expanded;

rudimentary; with a row of long plumose setae

present. Pereopod 7 basis anterior margin straight;

distally rounded; medial row of setae absent.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth;

Urosome: articulation between urosomites 2

and 3 absent. Uropod 1 peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle shorter than rami; outer ramus as long as

inner. Uropod 3 peduncle longer than half length

of rami; outer ramus 2-articulate, longer than

inner.

Tclson longer than broad; longer than peduncle

uropod 3; submarginal setae on apex of each lobe

absent; cleft; apically rounded.

Males: Unknown.

Etymology. The name refers to the fact that this

was the first species we discovered without a

double-hooked setae on the outer plate on maxilla

2 (MX2, Fig. 6).

Distribution. Antarctica, 198—473 m (possibly

circumpolar).

Remarks. This and P. vanhoeffeni are the only

species in the genus occurring in Antarctica. Both

are separated from their congeners by the strongly

produced and serrate hind margin of the third

epimeral plate. Phippsia unihamata differs from

P. vanhoeffeni in the long and geniculate

peduncle of antenna 2.

Phippsia vanhoeffeni (Schellenberg, 1926)

comb. nov.

Figures 13-15

Stegocephaloides vanhoeffeni Schellenberg, 1926:

299.—K.H. Barnard, 1930: 328.

Stegocephalopsis vanhoeffeni .—Barnard and

Karaman, 1991: 681.

Material examined. Syntype. Eastern Antarctica

(Wilhelm II land), 385 m, ZMB 20388 (female, 6 mm).

Additional material. Australia, Victoria, S of Point

Hicks (38°21.90'S, 149°20.00'E). 1000 m, WHOI

epibenthic sled, G.C.B. Poore et al. on RV Franklin 23

Jul 1986 (stn SLOPE 32), NMV J24057 (4 males and

females, 4-5mm).

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

Figure 13. Phippsia vanhoeffeni (Schellenberg, 1926). NMV J24057, female A.

168

J0RGEN BERGE AND WIM VADER

Figure 14. Phippsia vanhoeffeni (Schellenberg, 1926). NMV J24057, female A.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

169

Figure 15. Phippsia vanhoeffeni (Schellenberg, 1926). NMV J24057, female A.

170

J0RGEN BERGE AND WIM VADER

Description. Rostrum very small.

Antennae short. Antenna 1 longer than antenna

2; flagellum 5-articulate; accessory flagellum

well developed, article 2 absent. Antenna 2

peduncle ( articles 3-5 ) longer than flagellum;

article 3 short, about as long as broad; article 4

longer than article 5.

Epistome curved (convex) and smooth;

epistomal plate (medial keel) not produced.

Mouthparts not elongate or pointed. Mandible

incisor lateral; toothed; left lacinia mobilis

powerful, laterally expanded. Maxilla 1 palp 1-

articulate; rectangular; apex not reaching above

the apex of outer plate; outer plate distally rectan¬

gular; ST in a pseudocrown; ST first row with 6

setae (ST 1-5, ST 7); ST 6 absent; gap between

ST 5 and ST 7 absent; ST A present; located

distally. part of first row; ST B present; part of

second row; ST C present; inner plate with a well

developed shoulder; setae pappocuspidate. Max¬

illa 2 gaping and geniculate; outer plate setae with

distal hooks present; distal cleft absent; inner

plate setal row A covering about two-thirds of the

margin; clearly separated from row B; row A

setae pappopectinate; row B setae proximally

pappose; distally with cusps present; row C pre¬

sent; row D present; expanded, row elongated

towards and beyond row A; with many small

cusps distally. Maxillipcd palp 4-articulate;

article 2 distally produced; distal inner margin

weaklyproduced; dactylus distally simple

(pointed); inner plate not exceeding base of palp

article 3; nodular setae 2; medial setal-row pre¬

sent; not reduced; vertical; setae pectinate; distal

setal-row present; inner setal-row absent; outer

plate outer setal-row present; marginal; setae

attached in a deep hollow; setae short; strongly

curved upwards (hooks); inner setal-row present;

well developed; setae long robust; pappose; prox¬

imally parallel to outer, distally transverse; distal

setal-group present; setae attached in a deep hol¬

low; setae short simple. Labrum about as long as

broad; lobes symmetrical; right lobe not reduced;

left lobe not reduced. Labium distally narrowing.

Coxal plates and basis on pereopods covered

with setae; setae very short. Coxae 1-3 contigu¬

ous. Pereopod 1 coxal plate deeper than basis;

propodus subovate. Pereopod 2 longer and thin¬

ner than pereopod 1; ischium not elongate, ratio

length:breadth not exceeding 1.5; ischium distal

posterior margin plumose setae present; propodus

subrectangular; palm absent. Pereopod 4 coxa

posteroventral lobe large, reaching about the base

of pereonite 7; basis anterior margin with long

setae absent; posterior margin with long setae

present; plumose setae on distal anterior and

posterior margins present; ischium plumose setae

on posterior distal margin present. Pereopod 6

basis posteriorly expanded; rudimentary; with a

row of long plumose setae present. Pereopod 7

basis anterior margin straight; distally rounded;

medial row of setae present; setae short and

robust.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod I peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus as long as

inner. Uropod 3 peduncle longer than half length

of rami; outer ramus 1-articulate, as long as inner.

Telson longer than broad; longer than peduncle

uropod 3; submarginal setae on apex of each lobe

present; cleft; apical ly pointed.

Males: Pereopod 2 propodus equally sized in

males and females; Urosome ordinary (similar to

females).

Distribution. Eastern Antarctica (type locality)

and south of Point Hicks, Victoria, Australia.

Remarks. This species is distinct from others of

Phippsia in several respects, by its uniarticulate

outer ramus of uropod 3, the weakly produced

article 2 of the maxillipcd palp, and the arrange¬

ment of setal-rows A, B and D on maxilla 2

(Bergc, in press a). However, the species is trans¬

ferred from Stegocephalopsis Sehellenberg, 1925

on the following characters: 1, articulation

between labrum and epistome partly absent; 2,

inner plate of maxillipcd long and rectangular; 3,

articulation absent between urosomites 2 and 3;

and 4, epimeral plate 3 produced and strongly ser¬

rate.

Tetradeion Stebbing

Tetradeion Stebbing, 1899: 207

Type species. Cyproidea crassa Chilton, 1883.

Species: Tetradeion crassum (Chilton, 1883); T.

quatro sp. nov.

Distribution. Australia, New Zealand and

subantarctic region.

Remarks. Stegocephalus latus Haswell, 1879 was

described from shallow water Tasmania, Aus¬

tralia. Although the species was inadequately

described and the two syntypes no longer exist

(Springthorpe and Lowry, 1994), the descriptions

by Haswell (1879, 1885) strongly resemble those

of T. crassum. The conspicuous eyes, large coxae

1-4, reduced pereopod 7 (although all articles

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

171

appears to be present in S. lalus), and absence of

an articulation between urosomites 2 and 3 are

similar. Stegocephalus lalus was transferred to

Stegocephalopsis by Barnard and Karaman

(1991), but there are however, good reasons to

suspect that it belongs in the same genus as T.

crassum. Pending a revision of the family (Berge

and Vader, in prep.), and due to inadequate infor¬

mation of this species’ morphology, it is not

herein transferred to Tetradeian.

Tetradeion crassum (Chilton)

Figure 16

Cyproidea crassa Chilton, 1883: 80.

Tetradeion crassum .—Stebbing, 1899: 207.—Steb-

bing, 1906: 157— Chilton, 1924: 631—K.H. Barnard,

1930: 329.—Hurley, 1955: 197.—J.L. Barnard, 1972:

155.

Material examined. New Zealand. North Cape (precise

location unknown), 3 m, 1 Sep 1911, Terra Nova stn

135 (see K.H. Barnard, 1930: 329), NHM

1930.8.1.95-104 (15 specimens, 2.5-3.5 mm). Auck¬

land Islands (50°52'S, 166°42'E), 135-139 m. 9 Feb

1965, Elantin Cruise 16 (stn 1425), DMC (female, 2

mm).

South Atlantic, S of Falkland Islands (54°43'S,

56°37'W), 339-357 m, 14 Mar 1966, Eltanin Cruise 22

(stn 1593), DMC (immature. 2 mm).

Description. Rostrum very small.

Antennae short. Antenna 1 shorter than antenna

2; flagellum 6-articulate; accessory flagellum

rudimentary; peduncle elongate. Antenna 2

peduncle (articles 3-5 ) longer than flagellum;

article 3 elongate, articles 3 and 4 geniculate;

article 4 shorter than article 5.

Epistome curved (convex) and smooth;

epistomal plate (medial keel) produced, but small.

Figure 16. Tetradeion crassum (Chilton, 1883). NHM 1930.8.1.95-104 female.

172

J0RGEN BERGE AND WIM VADER

Mouthparts elongate; pointed and narrow.

Mandible incisor lateral; toothed; left lacinia

mobilis powerful, laterally expanded. Maxilla 1

palp 2-articulate; rectangular; apex not reaching

above the apex of outer plate; outer plate distally

rectangular; ST in a pseudocrown; ST first row

with 6 setae (ST 1-5, ST 7); ST 6 absent; gap

between ST 5 and ST 7 present; ST A present;

located distally, part of first row; ST B present;

part of second row; ST C present; inner plate with

a well developed shoulder; setae pappocuspidate.

Maxilla 2 gaping and geniculate; outer plate setae

with distal hooks present; distal cleft present;

inner plate setal row A covering about two-thirds

of margin; appressed to row B; row A setae pap-

popectinate; row B setae proximally simple; dis¬

tally with cusps present; row C present; row D

present; expanded, row elongated towards and

beyond row A; with many small cusps distally.

Maxilliped palp 4-articulate; article 2 distally pro¬

duced; distal inner margin greatly produced;

dactylus distally simple (pointed); inner plate not

exceeding base of palp article 3; 3 nodular setae;

medial setal-row present; not reduced; vertical;

setae pectinate; distal setal-row present; inner

setal-row present; row reduced to 1 or 2 setae;

outer plate outer setal-row present; submarginal;

setae attached normally; setae long; straight; inner

setal-row present; well developed; setae long

robust; pappose; proximally parallel to outer, dis¬

tally transverse; distal setal-group present; setae

attached in a deep hollow; setae long robust.

Labrum elongate; lobes asymmetrical; right lobe

not reduced; left lobe reduced. Labium distally

narrowing.

Coxal plates and basis on pereopods covered

with setae; setae very short. Coxae 1-3 contigu¬

ous. Pereopod 1 coxal plate not as deep as basis;

propodus subovate. Pereopod 2 general appear¬

ance like pereopod 1; ischium not elongate, ratio

length.breadth not exceeding 1.5; ischium distal

posterior margin plumose setae present; propodus

subovate; palm absent.

Pereopod 4 coxa posteroventral lobe very large,

reaching beyond the base of the pereonite 7; basis

anterior margin with long setae absent; posterior

margin with long setae present; plumose setae on

distal anterior and posterior margins present;

ischium plumose setae on posterior distal margin

present. Pereopod 6 basis posteriorly unex¬

panded; with a row of long plumose setae present.

Pereopod 7 strongly reduced, only 2 articles pre¬

sent; basis anterior margin straight; distally

rounded; medial row of setae present; setae short

and robust.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod 1 peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus longer

than inner. Uropod 3 peduncle at least as long as

rami; outer ramus 1 -articulate, longer than inner.

Telson longer than broad; shorter than peduncle

uropod 3; submarginal setae on apex absent;

entire; apically rounded.

Males; Unknown.

Distribution. Circumpolar in the subantarctic

region including New Zealand, 0-357 m.

Remarks. Tetradeion crassum was thoroughly

described and figured by Chilton (1924), Hurley

(1955) and Barnard (1972).

The strongly reduced pereopod 7 and large red

eyes arc two distinctive features distinguishing

this species from all other known stegocephalid

species. The morphology of maxillae I and 2 and

the maxilliped indicate phylogenetic relationships

with Phippsia (see also above).

Tetradeion quatro sp. nov.

Figures 17-20

Material examined. Holotype. Australia, Victoria, east¬

ern Bass Strait, 8 km S of Southeast Point, Wilsons Pro-

motory, (39°12.9’S, 146°27.3'E), 65 in, medium sand,

R. Wilson on RV Tangaroa , 18 Nov 1981 (stn BSS

180), NMV J47020 (female, 7 mm).

Paratypes. Collected with holotype, NMV J45337

(30 specimens, males and females, 3-8 mm).

Additional material. Australia, Victoria, eastern Bass

Strait, 8 km S of Southeast Point, Wilsons Promotory,

(39°12.9'S, 146°27.3’E), 65 m, medium sand, R. Wil¬

son on RV Tangaroa , 18 Nov 1981 (stn BSS 180),

NMV J45342 (female, 6 mm).

Description. Rostrum very small.

Antennae short. Antenna 1 shorter than antenna

2; flagellum 7-articulate; accessory flagellum

rudimentary. Antenna 2 peduncle (articles 3-5 )

longer than flagellum; article 3 elongate, articles

3 and 4 geniculate; article 4 as long as article 5.

Epistome curved (convex) and smooth; epis-

tomal plate produced into a large conspicuous

medial keel.

Mouthparts not elongate or pointed. Mandible

incisor lateral; toothed; left lacinia mobilis

powerful, laterally expanded.Maxilla 1 palp

2-articulate; rectangular; apex not reaching

above the apex ofouter plate; outer plate distally

rectangular; ST in a pseudocrown; ST first row

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

Figure 17. Tetradeion quatro sp. nov. Holotype except habitus: paratype female, 4 mm.

174

J0RGEN BERGE AND WIM VADER

Figure 18. Tetradeion quatro sp. nov. Holotype.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

Figure 19. Tetradeion quatro sp. nov. Holotype.

176

J0RGEN BERGE AND WIM VADER

Figure 20. Tetradeion quatro sp. nov. Holotype.

THE STEGOCEPHALID GENERA PHIPPSIA AND TETRADEION (AMPHIPODA)

177

with 6 setae (ST 1-5, ST 7); ST 6 absent; gap

between ST 5 and ST 7 present; ST A present;

located distally, part of first row; ST B present;

part of second row; ST C present; inner plate with

a well developed shoulder; setae pappocuspidate.

Maxilla 2 gaping and geniculate; outer plate setae

with distal hooks present; distal cleft absent; inner

plate setal row A covering the entire margin;

appressed to row B; row A setae pappopectinate;

row B setae proximally pappose; distally with

cusps present; row C present; row D present;

reduced, 1-3 long setae distally; with many small

cusps distally. Maxilliped palp 4-articulatc; arti¬

cle 2 distally produced; distal inner margin

greatly produced; dactylus distally simple

(pointed); inner plate not exceeding base of palp

article 4; nodular setae 4; medial setal-row pre¬

sent; not reduced; vertical; setae pectinate; distal

setal-row present; inner setal-row present; row

reduced to 1 or 2 setae; outer plate outer setal-row

present; submarginal; setae attached normally;

setae long robust; straight; inner setal-row pre¬

sent; well developed; setae long robust; slender;

proximally parallel to outer, distally transverse;

distal setal-group present; setae attached in a deep

hollow; setae long robust. Labrum about as long

as broad; lobes symmetrical; right lobe reduced;

left lobe reduced. Labium distally narrowing.

Coxal plates and basis on pereopods covered

with setae; setae very short. Coxae 1 3 contigu¬

ous. Pereopod 1 coxal plate not as deep as basis;

propodus subovate. Pereopod 2 general appear¬

ance like pereopod 1; ischium not elongate, ratio

length:breadth not exceeding 1.5; ischium distal

posterior margin plumose setae present; propodus

subovate; palm absent.

Pereopod 4 coxa posteroventral lobe very large,

reaching beyond the base of pereonite 7; basis

anterior margin with long setae absent; posterior

margin with long setae present; plumose setae on

distal anterior margin absent; plumose setae on

distal posterior margin present; ischium plumose

setae on posterior distal margin present. Pereopod

6 basis posteriorly unexpanded; with a row of

long plumose setae present. Pereopod 7 strongly

reduced, only 4 articles present; basis anterior

margin straight; distally rounded; medial row of

setae present; setae short and robust.

Oostegites on pereopods 2-5, gills on

pereopods 2-7.

Pleonites 1-3 dorsally smooth.

Urosome: articulation between urosomites 2

and 3 absent. Uropod 1 peduncle longer than

rami; outer ramus longer than inner. Uropod 2

peduncle longer than rami; outer ramus longer

than inner. Uropod 3 peduncle longer than half

length of rami; outer ramus 2-articulate, longer

than inner.

Telson longer than broad; longer than peduncle

uropod 3; submarginal setae on apex present;

entire; apically rounded.

Males: Pereopod 2 propodus larger in males

than in females, urosome ordinary (similar to

females).

Etymology. The species name reflects the

presence of just four articles on pereopod 7.

Distribution. Eastern Bass Strait, Victoria (known

only from type locality).

Remarks. The presence of just four articles (in

addition to coxa) is a unique character state within

the family; T. crassum has just two articles, all

other stegocephalid species have six articles on

pereopod 7 (in addition to coxa).

Acknowledgements

We thank Drs Jim Lowry and Penny Berents

(Australian Museum, Sydney), Dr Gary Poore

(Museum Victoria, Melbourne), Dr Oliver Cole¬

man (Zological Museum, Berlin), Dr Les Watling

(Darling Marine Center, Maine), Dr Geoff

Boxshall and Ms Miranda Lowe (The Natural

History Museum, London) for their kind help in

providing the material for this study.

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