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CONTENTS

New genus and species of southern Australian and Pacific Asterinidae (Echinodermata,

Asteroidea)

P. M O'Loughlin .277

Report on selected species of BANZARE and ANARE Holothuroidea, with reviews of

Meseres Ludwig and Heterocuciimis Panning (Echinodermata)

P.M. O'Loughlin . 297

A new species oiMeanthes (Polychaeta: Nereididae) from southern Australia

T. Bakken .327

Con’slalona, a new hydroid genus (Leptolida; Leptothecatae) from southern Australia

J.E. Watson . 333

Hydroids (Cnidaria: Hydrozoa) from southern Queensland

J.E. Watson . 337

A revision of the Australian genus Umbilia (Gastropoda: Cypraeidae)

T.A.Dairagh . 355

The Australian species of Chimarra Stephens (Trichoptera; Philopotamidae)

D.I.Cartwright . 393

Five artotrogids (Crustacea: Copepoda: Siphonostomatoida) from Eastern Antarctica

R. Johnsson anti C.E.F. Rocha . 439

New Phreatoicidea (Cnistacea: Isopoda) from Grampians National Park, with revisions of

Synamphisopus and Phreatoicopsis

G.D.F. Wilson and S.J. Keable . 457

Redescriptions of A.sphalidesmus leae Silvestri, 1910 and A. parx’us (Chamberlin, 1920)

comb. nov. from Tasmania, Australia (Diplopoda: Polydesmida: Haplodesmidae)

R. Mesibov . 53 ]

Pycnogonum (Pycnogonida: Pycnogonidae) from Australia with descriptions of two new species

D. A. Staples . 541

Memoirs of Museum Victoria 59(2): 277—296 (2002)

NEW GENUS AND SPECIES OF SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE

(ECHINODERMATA, ASTEROIDEA)

P. Mark O’Loughlin

Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E, Melbourne,

Vic. 3001, Australia (e-mail: pmo@bigpond.net.au)

Abstract

O’Loughlin, P.M., 2002. New genus and species of southern Australian and Pacific

Astcrinidae'(Echinodermata, Asteroidea). Memoirs of Museum Victoria 59(2): 277-296.

The diagnostic characters of the Astcrinidae species Aslerimi atyphoiiia H.L. Clark from

southern Australia and Aslerina gihhosa (Pennant) from the Atlantic, type species of Asterina

Nardo, are reviewed. Asterina atyphoida is referred to Meridiastra gen. nov., with four other

species: Meridiastra fissiira sp. nov. and Meridiastra nigranota sp. nov. from southern

Australia, Meridiastra rapa sp. nov. from the central south Pacific, and Meridiastra modesta

(Verrill) from the Pacific coast of Panama and Mexico. Asterina agustincasoi Caso is

synonymised with Meridiastra modesta (Verrill). Asterinides Verrill is recognised as a valid

genus. Meridiastra fissura is flssiparous. Meridiastra is distributed from southern Australia to

the Pacific coast of Mexico and Panama. A key is provided for the species of Meridiastra.

Introduction

Rowe (in Rowe and Gates, 1995 and in a note in

Campbell and Rowe, 1997) considered that

species of the Asterinidae from Australian waters

which were assigned to the genus Asterina

Nardo, 1834 were not congeneric with the type

species Asterina gihbosa (Pennant, Mil), and

required reassignment to an existing or new

genus. Rowe and Berents (in an unpublished draft

manuscript, cited in this work as pers. com.)

anticipated assigning one of these species, Aster¬

ina atyphoida H.L. Clark, 1916, to a new genus

together with a new flssiparous southern Aus¬

tralian species. This anticipated genus is estab¬

lished here, and a second new species from

southern Australia is included. Tlie remain¬

ing Australian species assigned to Asterina are

not reallocated to this new genus, and their

reassignment is not treated in this work.

Marsh (1974) reported an Aslerina sp. from

Rapa I. in the South Pacific, and this species is

described below and assigned to the new genus.

The eastern Pacifle species Asterina agustincasoi

Caso, 1977 and Asterina modesta Verrill, 1870

are synonymised, and A. modesta is reassigned to

the new genus.

Rowe and Berents (pers. com.), in their antici¬

pated review of the genera of the Asterinidae,

considered that the presence or absence of super-

ambulacral internal plates, the internal alignment

of abactinal and actinal plates towards the body

margin, and the arrangement of cleared actinal

plates were helpful in distinguishing the genera.

These diagnostic features are included in the

descriptions below, without judgement as to their

significance. Clark and Downey (1992) used the

flrst two of these characters in their diagnosis of

Aslerina, referring to an absence of super-

ambulacral plates and to the reinforcement of the

ventrolateral angle by an internal overlapping of

the adjacent abactinal and actinal plates often

with additional plates, but did not describe the

arrangement of actinal plates. In this study the

arrangement of cleared actinal interradial plates

was found to be inconsistent for a specimen and a

species, and the apparent arrangement on

uncleared specimens was sometimes different to

that revealed on cleared specimens of the same

species. Vor Asterina gihhosa the actinal series on

one uncleared specimen appeared to curve at an

acute angle distally from furrow to margin (Fig.

le), while on a cleared specimen the series curved

obliquely (obtusely) from furrow to margin

(Fig. Hj.

Verrill (1913) reassigned Aslerina modesta to

his new genus Asterinides Verrill, 1913. Fisher

(1919) synonymised Asterinides with Asterina.

A.M. Clark (1983) and Clark and Downey (1992)

277

278

P, M. O’LOUGHLIN

considered that Asterinides (type species Aster-

iscus folium Lutken, 1860, from the West Indies)

could be revived from its synonymy with Asterina

and recognised at least as a valid subgenus. Rowe

(in Rowe and Gates. 1995 and in a note in Camp¬

bell and Rowe, 1997) supported the recognition

of Asterinides as a valid genus. Rowe and Bcrents

(pers. com.) considered that their anticipated new

genus, described below, was close to Asterinides,

but distinguished the two genera. Asterinides is

recognised below as a valid genus, but not the

appropriate genus for Asterina modesta.

Molecular phylogeny studies of Asterinidae

species, such as by Byrne et al. (1999), are pro¬

viding further evidence on which to review the

assignment of species to the many existing Aster¬

inidae genera. In the current absence of adequate

molecular evidence a review of the assignment of

species to Asterina and Asterinides is not under¬

taken here.

Terminology here follows that defined in the

glossary and illustrated in Figs 2 and 3 of Clark

and Downey (1992), except that “papular space"

is used for "papular area" (“restricted area with

papular pores”) and "papulate areas” is used to

refer to the parts of the abactinal surface where

papulae occur. "Ad-disc" refers to a location adja¬

cent and distal to the plates bordering the disc.

Abbreviations of institutions are as follows:

AM, Australian Museum, Sydney, Australia;

BPBM, Bishop Museum, Honolulu, USA; ICML-

UNAM. Instituto de Ciencias del Mar y Limnolo-

gia, Universidad Nacional Autonoma de Mexico,

Mexico; MCZ, Museum of Comparative Zool¬

ogy, Harvard University, Massachusetts, USA;

NMV, Museum Victoria, Melbourne, Au.stralia;

NTM, Northern Territory- Museum, Darwin, Aus¬

tralia; SAM, South Australian Museum, Ade¬

laide, Australia; TM, Tasmanian Museum,

Hobart. Australia; USNM, Museum of Natural

History. Smithsonian Institution, Washington.

USA; WAM, Western Australian Museum, Perth,

Australia; YPM, Peabody Museum of Natural

History, Yale University, Connecticut, USA;

ZMUC. Zoological Museum, University of

Copenhagen. Denmark.

Asterinidae Gray. 1840

/l.vreri//tf Nardo, 1834

Synonymy. See Clark and Downey (1992: 177).

Type species. A<iterina gibhosa (Pennant, 1777)

Diagnosis. See Clark and Downey (1992: 177).

Remarks. Rowe (in Rowe and Gates, 1995) listed

nine Asterinidae species from Australian waters

which were assigned to Asterina: A. alba H.L.

Clark, 1938 from the Tasman Sea; A. anomala

H.L. Clark, 1921 from Torres Strait; A. atypboida

H.L. Clark, 1916 from southern Australia; A.

cephetis (Muller and Tro.schel, 1842) from north-

eni Australia; A. coronata von Martens, 1866

from northern Australia; A. heteractis H.L. Clark,

1938 from the Tasman Sea; A. inopinata Living¬

stone, 1933 from eastern Australia; A. sarasini

(de Loriol, 1897) from northern Australia; and A.

scobinata Livingstone, 1933 from south-eastern

Australia. Rowe (in Rowe and Gates, 1995) con¬

sidered that these species assigned to Asterina

were not congeneric with the type species Aster¬

ina gibbosa (Pennant, 1777) and required reas¬

signment to an exi.sting or new genus. Of the

above nine species only Asterina atypboida H.L.

Clark is assigned here to a new genus. None of the

remaining eight species is congeneric with A. aty-

phoida (pers. obs. of AM specimens ol' all

species). Possible reassignment of these eight

species is not undertaken in this work. Rowe (in

Rowe and Gates, 1995) considered that the genus

Asterina might be restricted to Atlantic waters.

With the description of a new genus below and

inclusion of two species previously assigned to

Asterina. and the raising below of Asterinides out

of synonymy with .Asterina, the diagnosis of Aste¬

rina requires emendment. But a re-examination of

the many species currently assigned to Asterina

should be appropriately undertaken when the

emerging molecular evidence is adequate, and an

emended diagnosis is not given here.

Asterina gibhosa (Pennant, 1777)

Figures la-f, 7f

Synonymy. See Clark and Downey (1992: 184).

Material examined. Atlantic Ocean, European seas (no

other data), AM J1250(1); Ireland, Co. Down, Granagh

Bay, 5 Aug 1959, NMV F8724I (3); Wales, Gwynedd,

Anglesey. Rhosneigr, under rocks, 28 Sep 1971. NMV

F87242 (5). Mediterranean Sea. France. Marseilles, Jun

1978, AM J 12408 (3); Italy, Bay of Naples. AM

G11524 (2); donation from Naples Aquarium, 1890.

NMV F45I08 (3); Mediterranean (no other data), AM

G7575 (2).

Diagnosis. See Clark and Downey (1992:

184-185).

Remarks. Asterina gihho.sa material examined in

this study differed in minor ways from the diag¬

nosis by Clark and Downey (1992), and these dif¬

ferences and additional observations arc noted.

The first five radials involved in forming a pen¬

tagon bordering the disc were separated by single

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA)

279

Figure I. Aslorimi gihhoxa (Pennant) (Mediterranean specimens. lot NMV F45I()8). a. cleared abactinal ray

(R 19 mm); b. cleared disc and madreporite (lower right), with one non-papular proximal inlerradius (arrow);

c. abactinal spinelets (0..^ mm long); d. abactinal pedicellariae (one at arrow. 0.25 mm long); e. actinal interradius

with gonopores (arrows); F. eleared actinal interradius, with plates in series curving obliquely from Furrow towards

margin, and with gonopores (arrows).

280

P. M. O’LOUGHLIN

interradial plates. Papulate areas were more

extensive than non-papulate areas. The

madreporite was oval to lobed, not triangular. A

carinal series of plates in the mid-third of each ray

was distinct, the carinals typically separated by

pairs of secondary plates. Secondary' plates num¬

bered up to six in some pore areas. The supero-

marginals were the smallest of the abactinal inter¬

radial plates, were generally aligned with the

proximal edge of the projecting inferomarginals,

and were interspersed with varying numbers of

secondary plates. Associated with the distinctive

disc there were four ad-disc triangular interradial

papular spaces each with more than two sec¬

ondary plates and more than two papulae (fifih

space occupied predominantly by the madre¬

porite), and distal to these were five interradial

areas which had secondary plates and pedicellar-

iae but which lacked papulae and spines. There

were some conspicuous glassy convexities

around the margin but not on the crown of abacti¬

nal plates. Abactinal spinelets were typically

0.3 mm long. There were clusters of up to 6-8

papular pores. Suboral spines were noticeably

tall, and some plates had up to three. The first row

ol actinal adradial plates bore a complete series of

typical actinal spines. There was an irregular

series of secondary plates at the actinal edge of

the inferomarginal series. There were no internal

superambulacral plates, but there were internal

tapered contiguous projections on the distal

actinal and abactinal interradial plates giving

structural support to the ventrolateral angle.

Cleared actinal interradial plates curved in

oblique (obtuse) series from the furrow to the

margin.

Meridiastra gen. nov.

Type species. Asterinci atvphoida H.L. Clark,

1916

Diagnosis. Small, up to R = 13 mm; R/r =

1.2-1.3; rays 5 (6-8 in fissiparous species), ends

rounded, interradial margins straight or slightly

incurved: pentagonal or near-pentagonal if

5-rayed: body fiat orally, thin, acute angle at

margin, body slightly domed or low pyramidal

aborally; pedicellariae absent; triangular

madreporite.

Abactinal plates closely imbricating, in regular

scries; disc poorly defined or not distinct; papular

spaces small; projecting plates fan-shaped, except

radially, never predominantly crescentic or

deeply notched; papulate areas smaller than non-

papulate areas; papulae few, proximal, pre¬

dominantly radial, mostly single; abactinal plates

granular, covered with glassy convexities; abacti¬

nal plates with readily-detached glassy spinelets,

smaller than 0.3 mm long, across free margin on

proximal plates, not in tufts on proximal plates;

superomarginal plates not extending to margin,

generally aligned w'ith proximal edges of project¬

ing inferomarginals; lacking superambulacral

internal plates; distal actinal and abactinal inter¬

radial plates with internal tapered vertical

contiguous projections.

Projecting inferomarginal plates form margin,

with distal fine spinelets forming marginal fringe;

actinal plates in regular series; actinal interradial

plates with 1-3 spines; adambulacral plates

with 1-2 subambulacral spines, 2-3 webbed

adambulacral (furrow) spines.

Species. Meridiastra atyphoida (H.L. Clark,

1916); M Jissiira sp. nov.; M modesta (Verrill,

1870); M. nigranota sp. nov.; M rapa sp. nov.

Etymology. From the Latin meridies (south) with

astnirn (star), referring to the southern Australian

and Pacific occurrence of this seastar genus

(feminine).

Distribution. From southern Western Australia

to the Pacific coasts of Mexico and Panama;

0-59 m.

Remarks. Rowe (in Rowe and Gates, 1995) con¬

sidered that A. atyphoida w'as not congeneric

with A. gibhosa, the type species of Asterina, and

this judgement is supported here. The combina¬

tion of characters distinguishing the Asterina type

species, A. gibbosa. and Meridiastra arc listed in

Table 1. The diagnostic distinctions between

Meridiastra and the Asterinides type species, A.

folium, are listed in Table 3. Distinguishing char¬

acters are evident in small to large specimens.

Key to species of Meridiastra

Arms 6-8, form irregular (post fissipary); more than 1 madreporite; fis¬

siparous; only distal abactinal plates in regular series.

. Meridiastra fissura sp. nov.

Pentagonal fomi; single madreporite; not fissiparous; abactinal plates in

regular series.2

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA)

281

2. Secondary plates numerous proximally; abaclinal gonopores each with

I -3 proximal spinous secondary plates; actinal interradial plates with

predominantly 2 spines. Meridiasira modesta (Vcrrill, 1870)

— Secondary plates rare or absent proximally; abactinal or aetinal gonopores

lacking secondary plates; actinal interradial plates with predominantly 1

spine.3

3. Dise not distinct; proximal carinal series very irregular, extending less than

half ray length; actinal adradial row of interradial plates with complete series

of spines. Meridiaslru rapa sp. nov.

— Disc distinct; regular proximal carinal series extending three quarters ray

length; aetinal adradial row of interradial plates with spines irregularly

present or lacking.4

4. Gonopores abactinal; abactinal spinclets predominantly in single series on

proximal plate edge; subambulacral spines predominantly single; single sub¬

oral spine frequently present; live colour variable cream, yellow, orange,

mauve, red. brown. Meridiasira atyphoida (11.L. Clark. 1916)

— Gonopores actinal; abactinal spinelets frequently in irregular double series

on proximal plate edge; subambulacral spines predominantly paired; suboral

spines rarely present; live colour white with small black spots abactinally.

. Meridiasira nigranota sp. nov.

Meridiasira atyphoida (ILL. Clark) comb. nov.

Figures 2a-f

Aslerina ulyphnkki ILL. Clark. 1916: 57. pi. 17.—

ILL. Clark, 1928: 389.—Cotton and Godfrey, 1942:

201.—ILL. Clark. 1946: 130.—A.M. Clark. 1966:

324-325.—Shepherd, 1968: 745.—Dartnall. 1970a:

73.—Dartnall, 1980: 34. map 6( I).—Zeidler and Shep¬

herd. 1982: 41HI2, lig. 10.6(d).—Rowe and Vail,

1982: 222.—O'Loughlin. 1984: 134.—Clark. A.M..

1993: 207.—Marsh .and Paw.son, 1993: 281 (part.

Esperance specimens).—Rowe (in Rowe and Gates),

1995: 34.—Edgar. 1997: 344-345.

Malcrial examined. Ilolotype. South Australia, 15

miles NW of Cape Jerv'is, 31 m, AM E6303.

Paratypes. Kangaroo L.otTCape Marsden. 30 m. AM

E859 (2).

Other material (selection for distribution and depth

range). Victoria. East Gippsland Scallop Survey St. 41,

38"50 S, I47’4rE. 22 m. 28 Jan 1971. NMV l•73005

(2); Port Albert, donated 23 May 1906 (no other data).

NMV 1-71937 (1); San Remo, I Apr 1990. NMV

F58684 (2); Crib Point. 15 m. 12 Aug 1970, NMV

F71939 (3); Shoreham, 20 Aug 1972. TM II84I (I);

Flinders ocean platforms. 12 Dec 1993, NMV F87229

(I): Port Phillip Bay, Portsea Jetty, 5 ni. May 1975, AM

J9246 (I); Portland, Sa.xon Reef. 11 m, 5 Mar 1992,

NMV F87235 (I). Tasmania. Tamar River, Greens

Beach, 13 Jul 1969, TM II1059 (I); Deal L, East Cove,

5 m. 4 May 1974, AM J16573 (I): Frith L. 21 m, 1974.

NMV F87I60 (1); Bass Strait Survey St. 138, olT

Flinders I., 52 m, 6 Feb 19X1. NMV F87228 (9); St.

160,otTDeal I.. 59 m, l3Nov 1981. NMV F87236 (I).

South Australia [SA material referred to by ILL. Clark

(1916. 1928): SAM K103 (I), K104 (6), Kl()5 (I),

KI878 (6), KI879 (14)]; Victor Harbour, 5 m, 2 Apr

1980, NMV F87I68 (I); Kangaroo L, The Pages, NE of

Cape Willoughby, 27 m, 12 Apr 1941, SAM K1890(1),

Cape Jervis Jetty, 2 m. 9 Mar 1984, SAM K1887 (I);

Investigator Strait, 15 Jan 1971, NMV F87163 (2);

Yorke Peninsula, Edithburgh Jetty. 3-4 m. 10 Sep 1995.

SAM KI872 (I); Sir Jo.scph Banks Group, Lusby 1., 6

m. 11 Jan 1984, SAM K1885 (2); Port Lincoln, 19 Mar

1968. NMV F87233 (I); Elliston. 18 m, 12 May 1971.

NMV F87I5S (I); Nuyts Archipelago, Franklin L, 6-8

m. 15 Apr 1983, SAM K1S7I (I):'Point Sinclair, 10

May 1973. NMV F8716I (3). We.stcrn Australia,

Esperance, between Sandy Hook 1. and Cape Le Grand.

31-35 m. 23-25 Jun 1986. WAM Z6X55 (5).

Description of material. Small, up to R = 13 mm;

form pentitgonal, interradial edges straight, some¬

times near-pentagonal with interradial margins

incurved, typically R/r - 1.3; rays 5, rarely 6, ends

rounded; body Hat orally, thin, slightly domed or

low pyramidal aborally, acute angle at margin;

single triangular madreporite; not llssiparous;

pedicellariae absent; paired interradial gonopores

abactinal.

Abactinal plates closely imbricate, in regular

series; papular spaces small; papulale areas less

extensive than non-papulate areas; secondary

plates rare, disc and margin only, never separating

carinal plates; disc variably distinct, bordered by

irregular pentagon comprising 5 proximal carinal

and 5 interradial plates; each radius with 5 longi¬

tudinal series of plates in mid-ray; distinct proxi¬

mal carinal series comprises up to 10 (rarely 11)

plates extending three quarters of ray length, dis-

tally typically 6 (up to 14) zig-zag radial plates;

carinals with proximal edge slightly convex or

straight or concave, some with slight median

lobe; some radials indented proximally for I or

rarely 2 papulae; projecting interradial plates

f(D

282

P. M. O’LOUGHLIN

Clark) comb. nov. a, abaclinal ray and interradii (R 10 mm) (NMV

), b, cleared abaclinal ray and interradii (R = 11 nun) (NMV F45092); c, cleared proximal abaclinal plates,

madreporite (lower right of disc), and indistinct disc (arrows at two proximal carinal plates) (NMV F45092);

d, abactmal plates with spinelcts (0.1 mm long), glassy convexities, and madreporite (upper right) (NMV F73465);

e cleared abactmal mterradial plates with two gonoporcs (arrows) (NMV F45092); F. proximal actinal view, with

absence of spines on adradial actinal plates (arrow) (NMV F87229)

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA) 283

fan-shaped; abactinal plates granular, covered

with conspicuous glassy convexities: plates with

commonly up to about 10 (rarely up to 20) fine

bluntly-pointed readily-detached glassy spinelets,

up to 0.15 mm long, mostly across proximal edge

in irregular single transverse series, rarely over

plate, decreasing in number distally; papulae

small, mostly single, on proximal two-thirds ot

radial areas, in proximal third only ol carinal

series, up to 6 longitudinal series in mid-ray, few

on proximal interradial areas; superomarginals

not extending to margin, generally aligned with

proximal edge of projecting inferomarginals. each

with up to 6 vertical spinelets on projecting crown

of plate; some secondary plates proximal to

superomarginal series; lacking internal superam-

bulacral plates between ambulacrals and actinals;

distal abactinal and actinal interradial plates with

internal tapered vertical contiguous projections.

Projecting inferomarginal plates form margin,

each with marginal fringe of 6-7 distal line

spinelets, typically 0.25 mm long, proximal trans¬

verse row of shorter spinelets (typically 5); some

secondary plates at actinal edge of inferomarginal

series; cleared actinal plates in regular series,

curv ing acutely from furrow to margin or perpen¬

dicular to furrow; actinal interradial plates with I

spine, rarely 2 in mid-interradius, I .3 very' small

spines distally; adradial row of actinal interradial

plates lacking spines; adambulacral plates with 1

thick to tall subambulacral spine (sometimes 2 in

mid-ray and distally), .3 (rarely 4) webbed furrow

spines (2-1 distally); oral spines 4-6 (commonly

5); I suboral spine frequently present on every

oral plate, rarely none.

Live colour. Aboral colour varies, unifomi or

spotted or mottled, with white, pale brown, pink,

red, red-brown, red-orange, mauve, yellow; disc

and ends of rays dark grey-green to black; orally

white with some irregular orange to reddish-

brown llccking (Victorian specimens). Shepherd

(1968) reported pale cream, yellow or light brown

for South Australian specimens.

Distrihution. Southern Australia, East Gippsland.

Victoria, to Espcrance, Western Australia; Bass

Strait coast of Tasmania; 0-59 m.

Remarks. Material from Rottnesi I., Western Aus¬

tralia, reported by Marsh and Pawson (1993) as 4.

alyphoicki, and a single specimen from Flinders,

Victoria, reported by Dartnall (1970b) as A. aly¬

phoicki, are the closely related M. nigranola

described below. No evidence was found to con¬

firm a distribution of M alyphoiJa as lar we.st as

Rottnest I., as reported by Rowe (in Rowe and

Gates, 1995). Nor was evidence found to confirm

a distribution as far south as SE Tasmania, as

reported by Dartnall (1980). Within Meridiastra

the combination of diagnostic characters which

distinguish M. alyphoida are the pentagonal lorm;

long regular proximal carinal series of plates

extending more than hall" the ray length; absence

of proximal secondary plates except rarely within

the disc; absence of spines on the adradial actinal

plates; predominantly single subambulacral

spines; frequent presence of suboral spines; and

abaetinal gonopores.

Meridiastra Jissiira sp. nov.

Figures 3a-f

MeUeriat examined, llolotype. Victoria, Flinders, ocean

platfonns, sublittoral rocky shallows, M. O’Loughlin

and J. Ortenburg. 9 May 1993, NMV F87157.

Paralypes. Type locality, 16 Nov 1980. NMV

F71878 (1): 13 Apr 1985, NMV F86021 (7); 5 Oct

1991, NMV F87227 (I); 22 Feb 1992, NMV F7I880

(1); 12 Apr 1993. NMV F71879 (3); 12 Dec 1993,

NMV F86019(l).

Other material (selection for distribution and depth

range). New South Wales, Little Bay, Sydney, 17 Jiin

2001, NMV F8916l(l); Eden, Twofold Bay, Yallumgo

Cove, 24 Nov 1984 (pers. obs.). Victoria. Cape Conran,

5-6 ni. 15 Apr 1998. NMV F87155 (1); Waterloo Bay.

10 m. 27 Feb 1996. NMV F87066 (1); Venus Bay,

Twin Reefs, 4 Mar 1982. NMV F71755 (1); Cape

Paterson. 14 Feb 1981. NMV F87237 (1); llarmers

Haven. 24 Feb 1983, NMV F71736 (2+2 cleared);

Phillip L. Kitty Miller Bay. 8 Jan 1986, NMV F87411

(1); Cheviot Beach, 31 Mar 1998, NMV F87065 (14);

Port Phillip Heads, 15 m. I Jtil 1982, NMV F86022 (2):

Portsea Jetty, 5 m. May 1975. AM J9243 (I); Torquay,

Point Danger. 28 May 1982, NMV F87239 (1); Mullet

Holes, 10 km NF. Apollo Bay, 2 Jan 1988 (pers. obs.);

Portland, 28 Feb 1992. NMV F7I883 (4). Tasmania,

Alonnah. 12 Jan 1989 (pers. obs.); Maria I., 30 m, 23

Apr 1985, NMV F86020(T); Deal L. East Cove. 15 m.

26 Mar 1981. NMV F7I877 (5+3 cleared). South Aus¬

tralia. Robe. 9 Jan 1990 (pers. obs.); Victor Harbour. 9

Nov 1988 (pers. obs.); Cape Jervis. 10 Nov 1988, NMV

F87412 (1): Normanville, 11 Nov 1988 (pers. obs.);

Point Labatt. under rocks, intertidal. 15 Jan 1976, AM

J10867 (7). Western Australia. Cheyne Beach. Lookout

Point, 10 Nov 1969. NMV F87238 (2); Perth, Garden 1..

1-2 m, 10 Dec 1983. WAM Z9695 (3).

Description. .Small, up to R = 9 mm; rays 6-8, fre¬

quently 7, ends rounded, interradial edges slightly

incuiA'ed, typically R/r = 1.3; form frequently

asymmetrical (post fissipary), rays may be 3-4;

body fiat orally, thin, slightly domed aborally,

acute angle at margin; madreporites up to 4,

inconspicuous; anal openings up to 5; fissiparous;

pedicellariae absent; 1-2 gonopores in each

abactinal interradius.

284

P. M. O’LOUGHLIN

Figure 3. MerUtuistraftssura sp. nov. a. abactinal view of holotype (R = 5 mm) (NMV F87I57); b, actinal view of

holotype; c, cleared proximal abactinal plates, and two madreporites (arrows) (NMV F71736); d. abactinal spinelets

( . mm long) (holotype); e, cleared abactinal rays and interradius (NMV F71736); f, actinal view of two specimens

immediately after fissipary (NMV F87240).

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA) 285

Abactinal plates closely imbricate except prox-

imally, in regular series distally only, in irregular

mosaic proximally; papular spaces small; papu-

late areas similar in extent to non-papulate areas;

secondary plates apically and distally, never sep¬

arating carinals; lacking distinct disc; 5 longi¬

tudinal series of radial plates mid-ray; carinal

series in mid-ray only; distal projecting abactinal

plates fan-shaped, some radials with slightly con¬

cave proximal edge; size and fonn of proximal

abactinal plates irregular; single (rarely paired)

large papulae irregularly over proximal abactinal

surface, absent from disRil radial and interradial

areas, proximal plates frequently slightly notched

for single papula; abactinal plates granular, cov¬

ered with fine glassy convexities; plates with up

to about 30 small rugose frequently-webbed

readily-detached glassy spinelets, typically 0.1

mm long, upright, scattered over plate, concen¬

trated marginally, some plates with series on

proximal edge pointing apically; superomarginals

not extending to margin, generally aligned with

proximal edges of projecting inferomarginals,

some secondary plates along proximal edge of

superomarginal series; lacking internal super-

ambulacral plates between ambulacrals and

actinals; distal abactinal and actinal interradial

plates with internal tapered vertical contiguous

projections.

Projecting inferomarginal plates form margin,

each with marginal fringe of up to 7 distal fine

spinelets, typically 0.2 mm long; non-calcareous

actinal interradial areas proximally; cleared acti¬

nal interradial plates in regular series distally,

curving acutely from furrow to margin; some sec¬

ondary plates at actinal edge of inferomarginal

series; actinal interradial plates frequently with

2-3 spines proximally; adradial row of actinal

interradial plates with incomplete series of single

often-reduced spines; adambulacral plates with 2

webbed subambulacral spines, 3 (rarely 4)

webbed furrow spines; 6-4 oral spines; lacking

.suboral spines.

Live colour. Aborally faun to cream to white

proximally, white distally, with a few irregular

orange to red to dark reddish-brown spots, some

spots brownish distally; madreporites yellow; rare

dark greenish-black markings around ends of

rays; some anal openings surrounded by dark

greenish-black or red; white orally.

Distribution. Southern Australia, from Sydney

(central New South Wales) to Garden I. (SW

Western Australia); D’Entrecasteaux Channel,

Alonnah (southern Tasmania); rock substrate,

0-30 m.

Etymology. From Latin fissura (split), referring to

the fissiparous reproductive habit.

Remarks. This small species is fissiparous, and

post-fissipary forms are frequently found. The

number of arms (6-8), fissiparous habit, irregular

form, multiple inconspicuous madreporites, very

irregular proximal abactinal plates, and proximal

abactinal spinelets not predominantly across the

free margin of the plates distinguish this species

and make it exceptional within the genus. But the

small size, slightly incurved margins, distal

abactinal plate arrangement, papular arrange¬

ment, small glassy readily-detached abactinal

spinelets, actinal spination and absence of pedi-

cellariae justify placement in Meridiasira rather

than the establishment of a new monotypic Aster-

inidae genus.

Meridiasira modesta (Vcrrill) comb. nov.

Figures 4a-f

Aslerina (Asteriscus) modesta Verrill, 1870: 277.

Asterinides modesta. —Verrill, 1913: 482.—Verrill,

1915: 61.

Asterimi modesta. —Fisher, 1919: 410.—ILL. Clark.

1916: 57.—ILL. Clark. 1946: 130.—A.M. Clark, 1993:

211 - 212 .

Asterina agiistincasoi Caso, 1977: 209-231, figs

8-16, this 2-3 (syn. nov.)

Material examined. Asterina modesta. Leclotype (YPM

24175, alcohol). Panama. F.ll. Bradley, 1866.

Paraleclotypes. Type scries, YPM 605A (3, dry,

mounted); YPM 605B (3. alcohol); MCZ 776 (1, dry);

Panama, Pearl I.. F.ll. Bradley. 1866. YPM 1536 (I,

alcohol).

Other material. Panama, Panama Bay, Taboga L, Jun

1914, USNM 39110(1).

Asterina agnstincasoi. Paraleclotypes. Type series.

Mexico, Guerrero. 12 km W of Zihuatanejo, Ixtapa I.,

Jan 1963. ICML-UNAM 2.42.0 (9).

Description of material. Small, up to R = 10 mm;

form pentagonal, interradial edges straight, some¬

times near-pentagonal with interradial margins

incurved, typically R/r = 1.2-1.3; rays 5, rarely 4,

ends rounded; body fiat orally, thin, slightly

domed aborally, acute angle at margin; single tri¬

angular madreporite; not fissiparous; pedicellar-

iae absent; paired interradial gonopores abactinal,

under proximal edge of same or separate plates,

with 1-3 proximal spinous secondary plates.

Abactinal plates closely imbricate, in regular

series; papular spaces small; papillate areas less

extensive than non-papulate areas; numerous sec¬

ondary plates proximally, 1-5 in papular spaces

radially and interradially, frequently between

carinal plates, proximal to abactinal gonopores in

286

P. M. O’LOUGHLIN

Figure 4. MciUliastni modeskt (Verrill) comb. nov. a, abaclinal view of leclolype (R = 7 mm) (YPM 24175);

b. abactinal primary and seeondary plates, and spinclcts (0.15 mm long) (U.SNM 39110); c. abaclinal rays and

interradius (R = 10 mm) (USNM 39110); d. cleared abactinal ray and interradii (R ^ 7 mm) (paralectotype of

cigusiincasoi); e. margin (lower) and distal inlerradial primary and secondary plates and spines, with spinous

secondary plates proximal to two gonopores (arrows) (USNM 39110); f, actinaf surface (USNM 39110).

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA) 287

mid-inlcrradius, adjacent proximally to supero-

marginal plates; disc variably distinct, bordered

by irregular pentagon comprising 5 proximal cari-

nal and 5 smaller interradial plates; each radius

with 4 longitudinal scries of plates in mid-ray;

radial plates with proximal edge slightly convex

or straight or slightly concave, sometimes with

slight median lobe; some radials indented prox¬

imally for 1 or commonly 2 or rarely 3 papulae;

short proximal carinal series comprises typically

5 plates extending third to half ray length, dislally

12-14 zig-zag radial plates; projecting interradial

plates fan-shaped; abactinal plates granular, cov¬

ered with conspicuous glassy convexities; plates

with up to 12 short bluntly-pointed readily-

detached glassy spinelets, 0.10-0.15 mm long, on

proximal edge of plates in 1 -2 irregular trans¬

verse rows, pointing apically; 4-6 vertical

spinelets on crown of distal abactinal interradial

plates; papulae commonly paired, small, longi¬

tudinal scries along three quarters of ray laterally,

few in proximal interradial areas; supcromarginal

plates not extending to margin, aligned with prox¬

imal edge of projecting infcromarginals except in

mid-ray, each with up to 5 vertical spinelets

concentrated distal ly on plate; secondarj' plates

along proximal edge of supcromarginals; lacking

internal superambulaeral plates between ambu-

lacrals and actinals; distal abactinal and actinal

interradial plates with internal tapered vertical

contiguous projections.

Projecting inferomarginal plates form margin,

each with marginal fringe of double oblique

combs each up to 7 spinelets, up to 0.3 mm long;

secondary plates adjacent to infcromarginals acti-

nally; cleared actinal interradial plates in scries

curv ing obliquely or acutely from furrow to mar¬

gin (from mid-ray); actinal interradial plates with

predominantly 2 spines, variably 1-3; actinal

adradial row of interradial plates lacking spines

proximally, then irregular series of spines, pre¬

dominantly I, sometimes 2 mid-ray; adambu-

lacral plates w ith predominantly 2 webbed sub-

ambulacral spines proximally, I distally; 4 furrow

spines proximally, 3 mid-ray, 2 dislally, webbed

to tips; oral spines typically 5, subspalulate, not

pointed; suboral spines I, tall, tapered.

Live colour. Ivory, almost white (Caso, 1977).

Disirihiilion. Pacific coasts of Panama and Mex¬

ico; rocky shallows.

Remarks. Caso (1977) distinguished Asterina

agii.stlnca.soi from Asterina miniala Brandt, but

did not distinguish the new species trom Asterina

moclesta. Type material of A. agustincasoi and A.

mociesla w'as examined, and the species arc syn-

onymised. A lectotype and paralectotype series

are e.stablished for A. moclesta, and this species is

reassigned to Mericliasira. ILL. Clark (1916,

1946) noted the similarity of A. moclesta to A. aty-

phoicla. Within the genus M moclesta is distin¬

guished by: the short proximal carinal series ol

plates, some of which are doubly papillate;

numerous proximal secondary plates; spinous

secondary plates proximal to the abactinal gono-

pores; and predominantly paired actinal inteira-

dial spines. Having numerous proximal secondary

plates makes the species unique within the new

genus. The form of the abactinal spinelets, and

disposition across the proximal edge of the plates,

are characters quite similar to the type species M.

atyphoicla.

.Mericliasira nigranota sp. nov.

Figures 5a- f

Asterina alyplioiclci. —Darlnall, 1970b: 19, fig. 1 [non

Asterina atyphoicla H.L. Clark, 19161.—Marsh and

Pawson, 1993: 281 (part, Rolincsl 1. specimens) [non

Asterina atyphokia ILL. Clark. 19161.

Material examined. Ilolotype. Australia. Victoria,

Flinders, ocean platforms, shallow rocky sub-littoral,

M. O'Loughlin, 13 Apr 1985. NMV F874I4.

Paratypes. Type locality, 6 Jun 1969, TM HI058 (1);

17 Dec 1969. NMV F87232 (2-H cleared): TM H493

(3); 6 Feb 1972. NMV F71938 (1): 10 Apr 1970, NMV

F874I3 (1): 7 Apr 1980. NMV F73fl02 (1): 13 Apr

1985. NMV F72030 (Cl cleared); 12 Dec 1985. NMV

F71930 (I); 17 Feb 1990, NMV F7193I (2); 11 Aug

1990. NMV F71941 (1); 5 Oct 1991. NMV F87234 (1);

22 Feb 1992. NMV F659I4 (1); 12 Apr 1993. NMV

F87174 (I I); 9 May 1993. NMV F87226 (6); 12 Dec

1993, NMV F8723I (I).

Other material (selection for distribution and depth

range). Victoria. Wilsons Promontory. 10 Mar 1984.

NMV F71936 (I): Cape Paterson, 6 Mar 1982, NMV

F71932 (2); Western Port. Crawfish Rock. 15 Feb 1969,

NMV F7314() (I); Port Phillip Bay, Portarlington. 13

Mar l977.NMVF73003(l);CapeNelson,3Mar 1984.

NMV F7I934 (1). Tasmania. NE between Ringarooma

Bay and Waterhouse Point, Mar 1970. TM 111330 (1);

King I.. Currie. 10 Mar 1980, NMV F87162 (2); Tas¬

man Peninsula, N of Thumbs Point, 6-10 m. 31 Jul

1991, SAM K1894 (5). South Australia. Port MacDon-

nell, 18 Mar 1976. .AM J9909 (5): olTCape Northum¬

berland, 30 m, 14 Jul 1974, SAM KI892 (I). Western

Australia, Rottnest L, Little Armstrong Bay, 0 m, 9 Jan

1991. WAM Z68.56 (I); Ricey Beach, 0-1 m, 14 Jan

1991. WAM Z6857 (l).

Description. Small, up to R = 13 mm; form pen¬

tagonal, interradial edges straight, sometimes

near-pentagonal with interradial margins

incurved, typically R/r = 1.3; rays 5, ends

288

P. M. O’LOUGHLIN

Figure 5 Meridiaslni nigraiwla sp. nov. a, abactinal view of holotype (R 7 mm) (NMV F87414); b, cleared

abactmal ray and mlerradii (paratype NMV F87232; R = 8 mm); c, abactinal view of dissected and cleared inter-

radius. showing ambulacral plates and absence of superambulacral plates (paratype NMV F87232); d, abactinal

^ (holotype); e. proximal actinal view, with single spine (arrow) on adradial plates

(NMV F87162); f, cleared actinal view, with gonopores (arrows) (paratype NMV F87232).

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA)

289

rounded; body flat orally, thin, slightly domed or

low pyramidal aborally, acute angle at margin;

single triangular madreporite; not fissiparous;

pedicellariae absent; paired interradial gonopores

actinal.

Abactinal plates closely imbricate, in regular

series; papular spaces small; papulate areas less

extensive than non-papulatc areas; ,secondary

plates rare, apical and marginal only, never

separating carinal plates; disc variably distinct,

bordered by iiTCgular pentagon comprising 5

proximal carinal and 5 interradial plates; each

radius with 5 longitudinal series of plates in mid¬

ray; distinct proximal carinal series comprises up

to 12 (rarely 13) plates extending three quarters of

ray length, distal ly 2-12 zig-zag radial plates;

carinals with proximal edge slightly convex or

straight or concave, some with slight median

lobe; some radials indented proximally for 1 or

rarely 2 papulae; projecting interradial plates fan¬

shaped; abactinal plates granular, covered with

conspicuous glassy convexities; plates with fre¬

quently more than 20 fine pointed rugose ter-

minally-spinous frcquently-w'ebbed readily-

detached glassy spinelets, up to 0.2 mm long,

clustered across proximal edge in irregular single

and double transverse series, frequently also scat¬

tered over plate, on projecting crown of distal

interradial plates; papulae small, mo.stly single, on

disc and proximal two-thirds of radial areas, in

proximal third only of carinal series, 4, longi¬

tudinal series in mid-ray, few on proximal inter¬

radial areas; supcromarginals not extending to

margin, aligned with proximal edge of projecting

inferomarginals, each with up to 6 vertical

spinelets; some secondary plates along proximal

edge of superomarginal series; lacking internal

superambulacral plates between ambulacrals and

actinals; distal abactinal and actinal interradial

plates with internal tapered vertical contiguous

projections.

Projecting infcromarginal plates form margin,

each with marginal fringe of 5 6 distal fine

webbed spinelets, typically 0.25 mm long, prox¬

imal transverse row of shorter spinelets (typically

4); some secondary plates at actinal edge of infcr¬

omarginal series; cleared actinal plates in regular

series, curving obliquely or acutely from I'urrow

to margin or perpendicular to furrow; actinal

interradial plates with 1 spine, frequently 2 in

mid-interradius, 1-3 very small spines distally;

actinal adradial row of interradial plates with

none to very few to ineomplete series of fre¬

quently reduced spines; adambulacral plates with

1-2 tall subambulacral spines proximally, fre¬

quently 2 from mid-ray distally; 3 webbed furrow

spines, 2-1 distally; oral spines 4-6 (commonly

5); suboral spines rarely present, frequently

reduced if present.

Live colour. Aboral colour white to faun to pale

grey, w ith irregular black to dark reddish-black to

dark greenish-black spotting, some brownish-red

to red spots distally; disc dark greenish-black;

madreporite white; faint red flecking around ends

of rays; white orally.

Disirihulion. Southern Australia, Wilsons

Promontory, Victoria, to Rottnest I., Western

Australia; Tasmania south to Tasman Peninsula;

0-30 m.

Etyinolog}’. From Latin niger (black) and nola

(spot), referring to the fine black aboral spotting

on live animals.

Remarks. Within Meridiastra the combination of

diagnostic characters which distinguish M. iiigra-

nota arc the pentagonal form; long regular prox¬

imal carinal series of plates extending more than

half the ray length; absence of proximal sec¬

ondary plates except rarely within the disc;

absence or iiTegular presence of spines on the

adradial actinal plates; predominantly paired sub¬

ambulacral spines; frequent absence of suboral

spines; and actinal gonopores. Within the new

genus M nigranoia is most closely related to

M atypbokku from which it is distinguished by

live colour and actinal gonopores.

Meridiastra rapa sp. nov.

Figures 6a-f

Asteritm —Marsh, 1974: 92-93.

Material e.xaniined (all dry). Holotypc. South Pacific,

Austral Ridge, Rapa I., Ilaurei Bay, off Point Tura-

gainuu, under basal! boulders, 1-2 rn. D.M. Devaney

Feb 1971, BPBM Wd.W.

Paratypes. Type series, BPBM W3033 (3); WAM

Z6875 (2).

Other material. Type locality, 27 Nov 1963, USNM

E53()99 (I).

Description of material. Small, up to R = 5.5 mm

(type series; R = 7 mm for flattened USNM

E53099); fomi pentagonal, interradial edges

straight, or near-pentagonal with interradial mar¬

gins incurv'cd, typically R/r = 1.3; rays 5, ends

rounded; body flat orally, thin, slightly domed

aborally, acute angle at margin; single triangular

madreporite; not fissiparous; pedicellariae absent;

paired interradial gonopores abactinal.

Abactinal plates closely imbricate, in regular

series except proximally; papular spaces small;

papulate areas slightly less extensive than

290

P. M. O’LOUGHLIN

Figure 6. Meridiaslra rapa sp. nov. a, abactinal view of liolotype (R = 5.5 mm) (BPBM W3507); b, abactinal view

of cleared paratype (R = 5 mm) (BPBM W3033); c, abactinal interradius of liolotype showing spinelets (0.1 mm

long); d, distal abactinal interradial spinelets (0.1 mm long) (paratype WAM Z6875); e, abactinal rays and intcrra-

dius (paratype WAM Z6875); f. actinal view (paratype WAM Z6875).

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA)

291

non-papulate areas; secondary plates absent

abactinally: lacking distinct disc; apical plates

large, irregular; lacking regular carinal scries of

plates, sometimes about 4 irregular proximal cari¬

nal plates; ray defined by 2 lateral longitudinal

series of plates singly notched for proximal two-

thirds of ray; mid-ray longitudinally comprises 2

series of alternating plates for length of ray, least

regular proximally, sparsely papillate with prox¬

imal plate edge slightly notched or concave for

proximal half of ray; each ray with 4 longitudinal

series of plates in mid-ray; projecting interradial

plates fan-shaped; abaclinal plates granular, cov¬

ered with fine glassy convexities; plates with up

to 14 fine pointed readily-detached glassy

spinelels, typically 0.15 mm long, vertical across

free edge of proximal plates in irregular double

scries, sometimes webbed, on crown of distal

plates; papulae single, papular spaces small,

papulae principally radial, distinct single series

along proximal two-thirds of ray laterally, 3 irre¬

gular longitudinal series of papulae in mid-ray;

some on proximal interradial areas; supero-

marginals not extending to margin, aligned with

proximal edge of projecting inferomarginals, each

with typically 2-3 vertical spinelels; lacking

internal superambulacral plates between ambu-

lacrals and actinals; distal abactinal and actinal

interradial plates with some internal vertical con¬

tiguous projections.

Projecting inferomarginal plates fomi margin;

marginal fringe of oblique combs of spinelels,

sometimes webbed, sometimes double series;, up

to 8, typically 0.2 mm long; few secondary plates

adjacent to inferomarginals; actinal plates

(cleared) in series curving obliquely (obtusely)

from furrow to margin; actinal interradial plates

with 1 2 spines, predominantly 1; actinal adradial

row of interradial plates all with 1 spine; adambu-

laeral plates with I 2 tall subambulacral spines,

predominantly 2; predominantly 3 webbed furrow

spines, 2 distally; oral spines 5; suboral spines

1 - 2 .

Elymolog}’. Named after the type locality, Rapa I.,

noun in apposition.

Distribution. Rapa I.; 0-2 m.

Remarks. Within Meridiastra the combination of

diagnostic characters which distinguish M. rapa

are the close to pentagonal form; very small size

(up to R = 5.5 mm); very irregular short proximal

carinal series of plates; irregular proximal plates

and absence of a distinct disc. M. rapa and M. Jh-

siira both have irregular proximal abactinal

plates, and lack a distinct disc.

Astcrinides Verrill

Figures 7a-e

Asterinides Verrill, 1913: 479.—Verrill. 1915: 58.—

Fisher. 1919: 410.—A.M. Clark, 1983: 364.—Clark

and Downey. 1992: 178.—A.M. Clark, 1993: 214.—

Rowe (in Rowe and Gates). 1995: 33.—Rowe (in

Campbell and Rowe), 1997: 131.

Type species. Asteriscus folium Liitken, 1860.

Diagnosis. Small, up to R = 15 mm; form near¬

pentagonal, interradial margins slightly incurved,

R/r = 1.3; rays 5, ends rounded; body flat orally,

low dome aborally, acute angle at margin;

madreporile plate with pores centrally only;

pcdicellariae absent; abaclinal gonopores.

Abactinal plates imbricate, in regular series;

papular spaces prominent; papillate areas more

extensive than non-papulate areas; disc clearly

delineated, 4 (one occupied by madreporile) ad-

disc interradial papular spaces with 2 papulae

separated by I -2 secondary plates; proximal sec¬

ondary plates rare; proximal radial and interradial

plates with distinct papular notch creating near-

cre.scentic appearance; some proximal and distal

interradial plates with small dome on crest;

abactinal plates granular, covered with conspicu¬

ous glassy convexities; abaclinal proximal plates

with central lufi of up to 20 slender readily-

detached glassy spinelels, up to 0.25 mm long,

distal and midinlcrradial superomarginal plates

with vertical clusters of near-paxilliform radiating

spinelels; superomarginal plates not extending to

margin, not always aligned with inferomarginals;

distal interradial and superomarginal plates small,

irregular in size and form; papulae numerous,

large, single, in distinct papular notch; lacking

superambulacral internal plates; distal actinal and

abactinal interradial plates with internal tapered

vertical contiguous projections.

Projecting inferomarginal plates fomi margin,

each with clusters of crowded radiating spinelels,

up to 0.25 mm long; cleared actinal interradial

plates in regular series, curving acutely from fur¬

row to margin; actinal interradial plates with 3^

spines midradially; adambulacral plates with 3-4

webbed subambulacral spines; 3-5 webbed fur¬

row spines; 1-6 suboral spines.

Remarks. Verrill (1913) distinguished Asterinides

Irom Asterina by the former’s lack of pcdicellar-

iac. Verrill (1913) assigned A. cepbeus (Miillcr

and Troschel) from northern Australia, and A.

modesta (Verrill) from the eastern Pacific, to

Asterinides. Fisher (1919) did not regard the pres¬

ence or absence of pcdicellariae as a reliable diag¬

nostic character, and based on the similarity of the

292

P. M. O’LOUGHLIN

Figure 7. a-e. AslermUles folium (Lutken). a, uncleared abaclinal view (R - 7 mm) (USNM 38811); b, cleared

abaciinal proximal plates, showing disc (USNM 38811); c, uncleared disc with madreporite (lop, arrow), ad-disc

papular spaces with longitudinal sccondarj' plates (one at white arrow) and subtending pairs of concave plates (one

at black arrow) (USNM 38811); d (upper), abactinal proximal intcrradial plates with tufts of spinelets (up to 0.25

Ii'rcvtxr®* (USNM 38811); d (lower), abactinal distal intcrradial plates w ith near-paxilliform clusters of spinelets

(USNM 38236); e, actinal view (U.SNM 38811); f, Asterina gihhosa (Pennant), dissected and cleared distal

mtcrradius with contiguous internal projections of abactinal and actinal plates (NMV F8724I).

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA) 293

abactinal skeleton and spines synonymised Aster-

inides with Aslerina. A.M. Clark (1983) and

Clark and Downey (1992) considered that Aster-

inides might be revived, based on the delicate

abactinal spinelets and relatively low and near¬

pentagonal body Ibim. Rowe and Bercnts (pers.

corn.) and Rowe (in Rowe and Gates, 1995 and in

a note in Campbell and Rowe. 1997) supported

the recognition of Asierinides as a valid genus. A.

modest a is reassigned above to Meridiastni gen.

nov. Specimens of A. cepheus from Queensland

(AM J2333I) and New Guinea (AM .122934)

were examined. Although the amis were distinct

and secondary plates absent, having a combina¬

tion of slender glassy abactinal spinelets, numer¬

ous actinal spines on each plate, abactinal plates

notched for single papulae, small papular spaees,

papulate more extensive than non-papulate areas,

irregular carinal series, abactinal gonopores, and

no pedicellariae supports a retention of this

species in Asierinides.

Based on the two type species Asterina gihhosa

(Pennant) and Asierinides folium (Liltken), the

combination of characters distinguishing Asterina

Nardo and Asierinides Verrill are listed in Table

2. Based on the type species Asierinides folium

(Liltken), the combination of characters distin¬

guishing Asierinides Verrill and Meridiastra gen.

nov. are listed in Table 3.

Asierinides folium (Liitkeii, 1860)

Figures 7a-e

Synonymy. See Clark and Downey (1992: 182-183).

Material examined. Holotype. Virgin Is, St Thomas.

Consul Krebs (ZMUC).

Other material. Atlantic Ocean, Gulf of Mexico,

Florida, Florida Key.s. off Key West, Sand Key Reef,

J.B. Henderson, no date, USNM 38811 (3); Bemiuda.

Hungry Bay, Sep 1901. USNM 38236 (4).

Description of material. Small, up to R = 15 mm;

fomi near-pentagonal to arms slightly pctaloid,

typically R/r = 1.4; rays 5, ends rounded; body

llat orally, low dome ahorally, acute angle at

margin; single madreporite, not triangular, pores

centrally only on plate; not fissiparous; pedi¬

cellariae absent; paired interradial gonopores

abactinal.

Abactinal plates imbricate, in regular series;

papular spaces conspicuous; papulate areas more

extensive than non-papulate areas; secondary

plates sometimes in papular spaces, some along

supcromarginal series, some next to infero-

marginals actinally; disc inconspicuous but dis¬

tinctive, bordered by pentagon comprising 5

proximal carinal and 5 interradial plates, 5 ad-disc

interradial papular spaces each with 2 lateral

papulae separated by 1-2 secondary plates or

madreporite plate; carinal plates irregularly pre¬

sent proximally, series up to 7 extending one third

of ray distally. midradials zig-zag longitudinally

for distal two-thirds of ray; proximal radial and

interradial plates with papular notch creating

near-crescentic appearance, up to 2 carinals dou¬

bly notched; some proximal and distal interradial

plates with small dome on crown; distal interradi-

als and superomarginals small, irregular in size

and arrangement, superomarginals not always

aligned with inferomarginals; abactinal plates

granular, covered with conspicuous glassy con¬

vexities; proximal plates with central tuft of up to

20 very slender, pointed, very readily-displaced,

glassy spinelets, up to 0.25 mm long, orientated

apically over papular spaee, some spinelet clus¬

ters near-paxilliform; distal interradial plates and

midinterradial supcromarginal plates with vertical

clusters of near-paxillifonti radiating spinelets;

papulae large, single except for 1-2 carinals, dis¬

tinct papular notches, some papular spaces with

1-3 secondary plates, papulae extending four

fifths of ray distally, up to three quarters of inter¬

radius distally; longitudinal papular series up to 8-

wide in mid-ray; lacking internal superambulacral

plates between ambulacrals and actinals; distal

abactinal and actinal interradial plates with inter¬

nal tapered vertical contiguous projections.

Projecting inferomarginal plates forni margin,

each with clusters of crowded laterally-radiating

spinelets up to 0.25 mm long; cleared actinal

interradial plates in regular series, curving acutely

from furrow to margin from mid-ray distally; acti¬

nal interradial plates with 2-5 slender webbed

spines, predominantly 3-4 in midinterradius;

adradial row of actinal interradial plates with 3^

webbed spines; adambulacral plates with 3-4

webbed subambulacral spines, 3-5 webbed fur¬

row spines; oral plates with 5-6 spines; 1-6

suboral spines, some webbed.

Distribution. See Clark and Downey (1992; 183).

Remarks. Verrill (1915) noted the absence of

“gyri" (wavy perforations) around the base of the

madreporite plate. The abactinal plates of the

Bermuda specimens (USNM 38236) frequently

had domes on the crests, which were not evident

on the holotype or in the Florida material (USNM

38811). Subsequent to the synonymy of Clark and

Downey (1992), Rowe (in Rowe and Gates, 1995

and in a note in Campbell and Rowe, 1997) dis¬

cussed the referral of A. folium to Asierinides, as

noted above.

294

P. M. O’LOUGHLIN

Table I. Characters distinguishing/titerwa (Pennant) and Meridiastra gen. nov.

Asterina gibbosa (Pennant)

Meridiastra gen. nov.

larger size; R up to 35 mm

short-rayed stellate form; arms distinct

crescentic, openly imbricating, abactinal plates

oval or lobed madreporite

clearly delineated disc

ad-disc triangular interradial papular spaces with

more than two secondary plates

papulate areas greater than non-papulate

large papular spaces with numbers of papulae

pedicellariae present

blunt opaque lirmly-attached abactinal spinelets

abactinal spinelets on crown of plates

small size; R up to 13 mm

pentagonal or near-pentagonal form

fan-shaped, closely imbricating, abactinal plates

triangular madreporite

disc poorly delineated or not distinct

lacking ad-disc triangular interradial papular

spaces

non-papulate areas greater than papulate

small papular spaces with single papulae

pedicellariae absent

fine glassy readily-detached spinelets

abactinal spinelets on free margin of plates

Table 2. Characters distinguishingNardo and Asterinides Verrill,

Asterina gibbosa (Pennant)

Asterinides folium (Liitken)

larger size; R up to 35 mm

short-rayed stellate fonn; arms distinct

numerous secondary plates abactinally

whole madreporite plate with pores

ad-disc triangular interradial papular spaces

with more than 2 secondary plates

papular spaces large, with numbers of papulae

pedicellariae present

blunt opaque firmly-attached abactinal spinelets

abactinal spinelets over crown of plates

gonoporcs actinal

small size; R up to 15 mm

pentagonal or near-pentagonal fonn

rare secondary plates abactinally

madreporite plate with pores centrally only

ad-disc triangular interradial papular spaces

divided by 1 -2 secondary plates

papular spaces smaller, with single papulae

pedicellariae absent

fine glassy readily-detached spinelets

abactinal spinelets in tufts on plates

gonopores abactinal

Table 3. Characters distinguishing Asterinides Verrill and Meridiastra gen. nov.

Asterinides folium (Lutken)

Meridiastra gen. nov.

abactinal plates prominently notched, near-,

crescentic

madreporite plate with pores centrally only

ad-disc triangular interradial papular

spaces divided by 1-2 secondary plates

papular spaces conspicuous

papulate areas greater than non-papulate

abactinal spinelets in tufts on plates

actinal interradial plates with 3-4 spinelets in

mid-ray

abactinal plates fan-shaped, not near-

crescentic

whole madreporite plate with pores

lacking ad-disc triangular interradial papular

spaces

papular spaces inconspicuous

non-papulate areas greater than papulate

abactinal spinelets across free margin

actinal interradial plates with 1-2 spinelets in

mid-ray

NEW SOUTHERN AUSTRALIAN AND PACIFIC ASTERINIDAE (ECHINODERMATA) 295

Acknowledgements

I am most grateful to: Dr Frank Rowe (fomierly

of the AM) and Dr Penny Berents (AM) who gen¬

erously allowed me to use an unpublished

manuscript which they were developing on the

Asterinidac; Loisette Marsh (WAM) and Dr

David Pawson (USNM) for providing infonna-

tion about the location of material; Eric Lazo-

Wasem (YPM) for providing photographs of type

material; Cynthia Aheam (USNM), Dr Penny

Berents (AM), Fred Collier (MCZ), Gavin Dally

(NTM), Lucius Eldridge (BPBM), Dr Alfredo

Laguarda Figueras and Israel Fabricio Barbosa

Ledesma (ICML-UNAM), Erie Lazo-Wasem

(YPM). Loisette Marsh (WAM), Dr Claus Nielsen

(ZMUC), Elizabeth Turner (TM), Wolfgang Zei-

dler and Thierry Laperousaz (SAM) for assistance

by making loan material available; Georgina Cun¬

ningham, Clarrie Handreck, Melanie Mackenzie,

Rebecca McIntosh, Tim O’Hara, Janinc Orten-

burg and the Marine Research Group of Victoria

for the contribution of fieldwork to this study; Sue

Boyd, Melanie Mackenzie and Chris Rowley for

assistance and use of the resources of Museum

Victoria; Peter Tutera for assistance with photog¬

raphy; Ben Boonen for assistance with the pre¬

sentation of the figures; Dr Frank Rowe and Tim

O’Hara for critical comments on the manuscript.

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Memoirs of Museum Victoria 59(2): 297-325 (2002)

REPORT ON SELECTED SPECIES OF BANZARE AND ANARE HOLOTHUROIDEA, WITH

REVIEWS OF MESERES LUDWIG AND HETEROCUCUMIS PANNING (ECHINODERMATA)

P. Mark O’Loughlin

Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E,

Melbourne, Vic, 3001, Australia

(e-mail: pmo@bigpond.net.au)

Abstract

O’Loughlin, P.M.. 2002. Report on selected species of BANZARE and ANARE

Holothuroidca, with reviews of Meseres Ludwig and Heterocucumis Panning (Echinoder-

mata). Memoirs of Museum Victoria 59(2): 297-325.

Selected Holothuroidca species collected by The British, Australian and New Zealand

Antarctic Research Expedition (BANZARE) 1929-1931 from Kerguelen I. and eastern

Antarctica, are listed. Relevant species of Holothuroidca also collected from the Kerguelen

region (Heard I.) and eastern Antarctica by the Australian National Antarctic Research Expe¬

ditions (ANARE) 1985-1997 are reported and discussed. The genera Meseres Ludwig and

Heterocucumis Panning arc reviewed. The genus Ekmocucumis Hcding and subgenus

Ekmocucumis Panning are junior synonyms of Staurocucumis Ekman. A new' Antarctic genus

of the Cucumariidac, Psolicrtix, and a new species of Synallactidac, Meseres spiculifenis, are

described. Systematic and distribution notes are provided for Chiridota pisanii Ludwig, Tae-

niog)'nis contortus (Ludwig), Bathvpiotes hongraini Vancy, B. gourdoni (Vancy), Meseres

globigeriiiae (Herouard), M. hyalegerus Sluiter, M. iuvolutus Sluiter. M. macdonaldi Ludw ig,

M. occultatus (Marenzeller), M. peripatus Sluiter, M. propinquus (Fisher), M. toreus (Theel),

M. traehus (Sluiter), M. villosus (Theel), Mesothuria bifurcata Herouard, Pseudostichopiis

atlanticus Perrier, Heterocucumis godejfroyi (Semper), //. steineni (Ludwig). PsoUcrux coatsi

(Vancy), Staurocucumis liouvillei (Vaney), Traebythyone lechleri (Lampert), Paracucumis

turrieata (Vaney), Amperima robusta (Theel), Rhipidothuria racovilzai Herouard and Laet-

mogone wyyillethomsoiii Thcc\. Pseudostichopus giobigerinae Herouard is removed Irom the

synonymy of Pseudostichopus villosus Theel. and assigned to Meseres. Pseudostichopus

propinquus Fisher is removed from the synonymy of P.seudostichopus pustulostis Sluiter, and

assigned to Meseres. A list of species with original and current name combinations, and a sum¬

mary of new distributions, are tabulated.

Introduction

The British, Australian and New Zealand Antarc¬

tic Research Expedition (BANZARE), under the

command of Sir Douglas Mawson, comprised

two voyages by the Discovery. The first voyage

left Cape Town in October 1929 and worked off

soulhert) Africa, Crozet. Kerguelen, Heard, east¬

ern Antarctica (40° to 80°E), and south-western

Australia. The second voyage left Hobart in

November 1930 and worked off southern Tas¬

mania, Macquarie, Scott, eastern Antarctica (60°

to 180°E), and western Tasmania. Station data

were detailed by Johnston (1937). In the Reports

following the Expedition, John (1939) reported

on the Crinoidca, Mortensen (1950) on the

Echinoidea, A.M. Clark (1962) on the Aster-

oidea, and Madsen (1967) on the Ophiuroidea.

The remaining Echinodermata from BANZARE,

the Holothuroidca, are reported in part here.

Material examined comprised 20 lots of 12

species, including all of the Aspidochirotida and

Elasipodida with some Apodida and Dendro-

chirotida. BANZARE Reports are no longer

published by the University of Adelaide. This

collection is lodged in the South Australian

Museum.

Recent holothurian collections, taken by the

Australian National Antarctic Research Expedi¬

tions (ANARE) 1985-1997 from Heard I. and

eastern Antarctica, are held in part by Museum

Victoria. Some ANARE species arc referred to in

this report to augment the data provided by the

BANZARE material, for the purposes of refining

the on-going systematic account of the Antarctic

and Kerguelen region holothurians and clarifying

which species have a distribution including the

southern coasts of South America, Antarctic

297

298

P. M. O'LOUGHLIN

coast, and Kerguelen region. Detennination of

some BANZARE lots necessitated a review of the

genera Meseres Ludwig, 1894 and Heiero-

cuctimis Panning, 1949. A new Antarctic genus

of the Cucumariidae, Psolkrux, and new species

of Synallactidae, Meseres spiculiferus, arc

described. Pseiuhstichopus globigerinae

Herouard, 1923 is removed from the synonymy of

Pseudostichopiis villosiis Theel. 1886. and

referred to Meseres. P.seuihslichopus propinquus

Fisher, 1907 is removed from the synonymy of

Pseiuhstichopus pustulo.sus Sluitcr, 1901. and

referred to Meseres. The genus Ekmocucumis

fleding. 1942 and subgenus Ekmocucumis Pan¬

ning. 1949 are synonymised with Staurocucumis

Ekman. 1927, and Heterocucumis Panning, 1949

is raised to generic status. All species considered

in this report, with new synonymies and new

combinations, are listed in Table I; BANZARE

species are listed in Table 2; and new extensions

of distribution are listed in Table 4.

Some of the material e.xamined provided evi¬

dence of possible regional variations for some

species, but in the absence of growth series and

adequate numbers of specimens the evidence is

considered to be insufficient for the establishment

of additional species. Variations, principally in

size, are noted.

Abbreviations for institutions are: AM, Aus¬

tralian Museum, Sydney; BMNH, The Natural

History Museum. London; IRSNB, Institut Royal

des Sciences naturelles de Belgique, Brussels;

MNHN. Museum National d'Histoire Naturelle,

Paris; NMV. Museum Victoria, Melbourne,

Australia; SAM, South Australian Museum. Ade¬

laide; USNM, US Museum of Natural History,

Smithsonian Institution. Washington; ZIM,

Zoological Institute and Museum, Hamburg;

ZMA. Zodlogisch Museum, Amsterdam; ZMUC,

Zoological Museum, University of Copenhagen.

The Magellanic region refers to southern Chile,

southern Argentina, and the Falkland Is. The Ker¬

guelen region refers to depths of less than 600 m

around Kerguelen, Heard, and the McDonald Is.

Bathyal refers to depths of 200-2600 m; abyssal

to 2600-6000 m (following Hansen, 1975).

Order Apudida Brandt

Chiridotidae Ostergren

Chiridota pisanii Ludwig

Chirodotapkanii Ludwig. 1886; 29-30, pi. 2 fig. 14.

Chirodotapurpurea.^Msic:^ 1886a: 15, 35. pi. 2 fig.

1 [non Trochodotapurpurea (Lesson. 1830)].

Chiridota pisanii. — Ludwig. 1892: 359.

Complete synonymy. .See Pawson (1964: 464).

.Material examined. Heard I.. 52"41'-53’13'S

72”56’-73‘4rE, 120-228 ni [ANARL, NMV

F84977-84979 (3 specimens)]. Argentina. Tierra del

Fuego, 54’00'S, 67’24'W. 0 ni, 1999 [NMV F860I6

(2)].

Type locality. Calbuco, Chile.

Distribution. Magellanic region, from Seno

Reloncavi in Chile to 43°S on the coast of

Argentina, Falkland Is. 0-102 m (summary by

Pawson. 1969). Kerguelen region. Heard L,

120-228 m (this paper).

Remarks. The specimens of C. pisanii from Fleard

l. and Tierra del Fuego were identical in tentacle

number) 12), tentacle digits (5-7 pairs, distalmost

pair largest), and ossicle complement [wheels, up

to 0.10 mm diameter, in papillae clusters; short

unbranched rods (miliary granules), 0.04-0.0S

mm long, in longitudinal muscles; curved tenta-

cles rods, typically 0.08-0.10 mm long; lacking

sigmoid hooks]. Theel (1886a) gave a similar

description ol specimens of C. pisanii from the

Falkland Is (as Chirodotapurpurea), but reported

wheel diameters up to 0.16 mm. Pawson (1964)

reported bracket-shaped tentacle rods with an

average length of 0.05 mm in material from

southern Chile. ANARE specimens of C. pisanii

taken off Heard I. are the first record for the Ker¬

guelen region, where the species is reported at a

greater depth. C. pisanii has not been reported for

Antarctica.

Taeniogyrus contortus (Ludwig)

Chirodota conlorla Ludwig. 1875: 80-81, pi. 6 figs

6a-c.

7'ac'n(ogvn/i cw;to/7M.v.—ILL. Clark, 1907: 121-123,

pi. 7 figs 8-13.

Taeniogyrus cf. contortus. —O'Loughlin et al.. 1994;

553, 554.

Complete synonymy. See Pawson (1964; 466-467).

.Material examined. Svntypcs ('!). Argentina. Santa

Cruz, east of Grande Bay, 51''34 S, 68'00'W, 91 m,

1888 [Albatross Stn 2771. USNM 19826 (3)].

Other material. Chile, Inutil Bay, 53*35'S. 69*45 W,

37-46 m, 1969 [USNM E33679 (13)]; 53*34'S,

69*59'W. 82-91 m. 1970 [USNM E33715 (9)|. Eastern

Antarctica. olT Wilkes Land. 66'18'.S, 110*32 E. 101

m. 1961 [USNM E33725 (25)]; MacRobertson Shelf,

66*55'-67-|6'S. 62*32'-68*59'E. 109-216 m

[ANARE, NMV F68691 (20), F69099 (2). F69100 (2)1.

Kerguelen L. Royal Sound, 49'28'S, 70*04'E. 4-5 m.

1929 [BANZARE Stn 12, SAM KI839 (l)J; 49*30 S,

69’48'E, 2-20 m, 1930 [BANZARE Stn 49, SAM

KI840(I)].

BANZARE AND ANARE HOLOTHUROIDEA

299

Table 1. Index ol'speeies in paper, ineluding new synonymies and new eombinations.

Original combination

Current combination or senior synonym Family

ahyssorum, Ciicumaria, Theel, 1886 Slaurocucumi.i ahyssomm (Theel, 1886) Cucumariidae

acaudum. Molpadiodemas, Heding. 1935 Pseudostichopus allcinlicus Perrier, 1898 Synallactidae

alalus, Pseudostichopus (Pseudostichopus), Pseudostichopus akitiis Imaoka, 1990 Synallactidae

Imaoka. 1990

cileutianus, Pseudostichopus, Ohshima. \9\5Meseresperipalus Sluitcr, 1901 Synallactidae

antarctica, Cucumaria, Vaney, 1906 Ileterocucumis steineni {Ludwig, 1898) Cucumariidae

autarctica, Paracucumis, MoTicuscn, 1925 Parcicucumis luiricata {Vancy, 1906) Paracucumidae

arenosus, Pseudostichopus, Ohshima, 1915 Pseudostichopus arenosus Ohshima, 1915 Synallactidae

citlanticus, Pseudostichopus, PcTricT, 1898 Pseudostichopus otlanticus PuTviev, 1898 Synallactidae

belyaevi. Amperima, Gebruk, 1988 Amperima helyaevi Gehruk, 1988 Elpidiidae

hifurcata, Mesothuria, Wtxouatd, 1901 Mesothuria bifurcata Wcrouard, 1901 Synallactidae

histriatum. Psolidium. Ludwig and Heding, Psolicnvc coatsi (Vaney, 1908) Cucumariidae

1935

hongraini. BcUhyplotes, Vaney, 1914 Balhyplotes hongraini Vaney, 1914

coatsi, Psolidium (Cucumaria), Vaney, Psolicrux coatsi (Vaney. 1908)

Synallactidae

Cucumariidae

1908

conspicua, Cucumaria, Vaney, 1908

contorla, Chirodota, Ludwig, 1875

denticulata, Cucumaria, Ekman, 1927

dilatorhis, Pseudostichopus

(Pseudostichopus), Imaoka, 1978

fuscivinculum, Bathyplotes, Gutt, 1990

globigerinae, Pseudostichopus, Herouard,

1923

Psolicrux coatsi {Vaney, 1908) Cucumariidae

Taeniogyrus contorlus {Ludwig. 1875) Chiridotidae

Heterocucumis denticulata (Ekman, 1927)Cucumariidae

Pseudostichopus dilatorhis Imaoka, 1978 Synallactidae

Bathyplotes hongraini Vaney, 1914 Synallactidae

Meseres globigerinae {Wemuavd, 1923) Synallactidae

godeffroyi, Cucumaria, Semper, 1868

godfroyi, Cucumaria, Vaney, 1914

gourdoni, Synallactes (?), Vaney, 1914

grandis, Cucumaria, Vaney, 1906

hassleri. Thyone, Theel, 1886

hyalegenis, Mw/w, Sluiter, 1901

ingolfi, Plicastichopus, Heding, 1942

involutus, Meseres, Sluitcr, 1901

japonensis, Pseudostichopus

(Trachostichopus), Imaoka, 1978

lechleri, Thvone (Thvonidium), Lampert,

1885

liouvillei, Cucumaria, Vaney, 1914

macdonaldi, Meseres, Ludwig, 1894

marenzelleri, Pseudostichopus, Herouard.

1923

mira, Cucumaria, Ludwig and Heding,

1935

Heterocucumis godeffroyi (Semper, 1868) Cucumariidae

Heterocucumis steineni (Ludwig, 1898) Cucumariidae

Bathyplotes gourdoni {Vaney, 1914) Synallactidae

Staurocucumis grandis {Vaney, 1906) Cucumariidae

Trachythyone lechleri {Lampen, 1885) Cucumariidae

Meseres hyalegerus Sluher, 1901 Synallactidae

Meseres peripatus Skiiler, 1901 Synallactidae

Meseres involutus Sluher, 1901 Synallactidae

Me.seres hyalegerus Sluher, 1901 Synallactidae

Trachythyone lechleri {Lamperi, 1885) Cucumariidae

Staurocucumis liouvillei (Vaney, 1914) Cucumariidae

Meseres macdonaldi Ludwig, 1894 Synallactidae

Meseres peripatus Sluher, 1901 Synallactidae

Staurocucumis liouvillei (Vaney, 1914) Cucumariidae

mollis, Pseudostichopus, Theel, 1886

molpadioides, Pseudostichopus, Ohshima,

1915

moseleyi, Stichopus, Thdel, 1886

navicula, Psolidium, Ekman, 1927

nudus, Pseudostichopus, Ohshima, 1915

occultatus, Pseudostichopus, Marenzeller,

1893

peripatus, Meseres, Sluiter, 1901

Pseudostichopus mollis Theel, 1886

Pseudostichopus molpadioides Ohshima,

1915

Bathyplotes mo.seleyi (Theel, 1886)

Psolicrux coatsi (Vaney, 1908)

Pseudostichopus nudus Ohshima, 1915

Meseres occultatus (Marenzeller, 1893)

Synallactidae

Cucumariidae

Synallactidae

Meseres peripatus Sluiter, 1901

Synallactidae

300

P. M. O'LOUGHLIN

Table I. Continued.

Original combination Current combination or senior synonym Family

pisanii, ChiroJota. Ludwig, 1886 ChiriJota pisanii Ludwig, 1886

plicatus var., occulkuus, Pseudostichopus, Meseres peripaitis Sluiter, 1901

Koehler and Vaney. 1905

propinquus, Pseudostichopus^ Fisher, 1907 Meserespropinquus (V'lsher. 1907)

rocoviizai. Rhipidollwria, Herouard, 1901

rohusta, Scoloplanes, Theel. 1882

rubipunetatus. Balhyploles, Gutt, 1990

spai/ui. Cucumorki, Cherbonnier, 1941

spiculiferus. Meseres sp. nov,

steineni. Cucumaria. Ludwig. 1898

tachimarmw. Pseudostichopus

(Trachostichopus), Imaoka, 1978

ton-us. Stichopus (?), Theel, 1886

trachus, Pseudostichopus. Sluiter, 1901

tuberculatus, Pseudostichopus

(Trachostichopus), Imaoka, 1990

turqueti, Cucumaria. Vaney. 1906

turricata, Thyonc. Vaney, 1906

unguiculatus, Pseudostichopus. Ohshima,

1915

Rhipidothuria racovitzai Herouard, 1901

Amperima rohusta (Theel, 1882)

Bathyplotes gourdoni (Vaney, 1914)

Staurocucumis grandis (Vaney, 1906)

Meseres spiculiferus sp. nov.

Heterocucumis steineni (Ludwig, 1898)

Meseres hyalegerus Sluiter, 1901

Meseres torvus (Theel. 1886)

Meseres trachus (Sluiter. 1901)

Meseres peripatus Sluiter, 1901

Staurocucumis turqueti (Vaney, 1906)

Paracucumis turricata (Vaney, 1906)

Meseres peripatus Sluiter, 1901

Chiridotidae

Synallactidae

Elpidiidae

Synallactidae

Cucumariidae

Synallactidae

Cucumariidae

Synallactidae

Cucumariidae

Paracucumidae

Synallactidae

villosus. Pseudostichopus, J\\ee\, 1886 Me.seres villosus {Thec\, 1886) Synallactidae

violaecuspidata. Achlyonice, Gutt. 1990 Rhipidothuria racovitzai WirouitrA. 1901 Elpidiidae

vitjazi, Amperima, GchxuV.. 1988 Amperima vitjazi Gehruk, 1988 Elpidiidae

wyviUethomsoni. Laetmogone,'Y\\ic\, 1879 Laetmogone wyvillethomsoni Theel, 1879 Laetmogonidae

Type locality. Straits of Magellan.

Distribution. Magellanic region (from 42°S on the

west and 43’S on the east of South America),

Straits of Magellan, Falkland Is, South Georgia,

Antarctic Peninsula, Antarctica at 88-89”E,

Kerguelen and Marion and Prince Edward Is,

0-560 m (summaries by Paw'son. 1964, 1969;

Branch et al., 1993). Western Antarctica, Weddell

Sea. 225-555 m (Gutt. 1991b). Eastern Antarc¬

tica. otT Terre Adelie, 180-220 m (Cherbonnier.

1974); otT Wilhelm II Land, 350-4()0 m (Ekman.

1927); off Wilkes and MacRobertson Lands,

101216 m (this paper).

Remarks. In terms of tentacle number (12) and

font! (6-7 pairs of digits with the distal pair

largest), ossicle distribution (wheels clustered,

hooks spread), and ossicle fonn and size

(wheels up to 0.17 mm diameter, sigmoid hooks

up to 0.28 mm long, tentacle rods up to 0.27 mm

long; lacking short rods in the longitudinal

muscles), all of the material e.xamined was

consistent diagnostically and also with the

descriptions by Theel (1886a) and ILL. Clark

(1907). There were regional variations in ossicle

size. Wheel diameter and hook length measure¬

ments of the BANZARE Kerguelen specimens

were the same as the measurements given by

Theel (1886a) for the Challenger Marion and

Falkland material, and were respectively up to

0.12 and 0.28 mm. In the type locality

material they were up to 0.17 and 0.21 mm;

in the Wilkes material up to 0.14 and 0.26 mm.

Ekman (1927) reported hooks to be 0.25 min

long in western Antarctic material; Deichmann

(1947) reported wheels up to 0.12 mm and

hooks up to 0.2 mm in Patagonia specimens;

Pawson (1964) reported wheels up to 0.13 mm

diameter and hooks up to 0.2 mm long in

southern Chile material; and O'Loughlin ct al.

(1994) reported both hooks and wheels up to

0.2 mm in length and diameter for eastern

Antarctic material (as Taeniogyrus cf. contortus).

The variations are not considered here to be

based on sufficient evidence for the establish¬

ment of separate species, but they do provide

some evidence of morphological change occur¬

ring with the reproductive isolation of these

viviparous populations. Further to the extensive

synonymy and distribution summary by Pawson

(1964, 1969), there have been reports of

T. contortus for Terre Adelie by Cherbonnier

BANZARE AND ANARE HOLOTHUROIDEA

301

Table 2. Selected BANZARE Holothuroidea species list.

Taxon, Registration, Specimens

Location

Station

APODIDA

Taeniogynts contortus (Ludwig, 1875)

SAM K1839 (1)

Kerguelen, 4-5 m

SAM K1840(l)

Kerguelen, 2-20 m

ASPIDOCHIROTIDA

Bathyplotes hongraini Vaney, 1914

SAM K1836 (2)

Antarctica, off Kemp Land, 603 m

Meseres spiculiferus sp. nov.

SAM K1850 (l)

Antarctica, off Enderby Land, 193 m

SAM KI851 (2)

Antarctica, Prydz Bay, 456 m

SAM K1852 (7)

Antarctica, off MacRobertson Land, 177 m

107

SAM K1853 (1)

Antarctica, off Wilkes Land, 695 m

Mesothuria bifurcata Herouard, 1901

SAM K1838 (1)

Antarctica, Prydz Bay, 1266 m

DENDROCHIROTIDA

Heterocucumis godeffroyi (Semper, 1868)

SAM KI844(2) '

Kerguelen, 47 m

SAM K1845 (2)

Kerguelen, 4-5 m

Heterocucumis steineni (Ludwig, 1898)

SAM K1832 (l)

Antarctica, off Enderby Land, 193-209 m

Staurocucumis liouvillei (Vaney, 1914)

SAM K1848 (4)

Antarctica, off Kemp Land, 177 m

107

Trachythvone lechleri (Lampert, 1885)

SAM kl 849(1)

Kerguelen, 47 m

Paracucumis turricata (Vaney, 1906)

SAM K1846(l)

Antarctica, off Wilkes Land, 474 m

ELASIPODIDA

Amperima robusta (Theel, 1882)

SAM K1842 (2)

North-east of Kerguelen, 3112m

Rhipidothuria racovitzai Herouard, 1901

SAM K1829 (l)

Antarctica, off Enderby Land, 300 m

SAM K1830 (24)

Antarctica, off Enderby Land, 300 m

SAM K1831 (3)

Antarctica, off Wilkes Land, 474 m

Laetmogone wyvillethomsoni Theel, 1879

SAM K1827 (1)

Antarctica, Prydz Bay, 540 m

SAMK1828 (1)

Antarctica, off Kemp Land, 603 m

(1974), for the Weddell Sea by Gutt (1991b),

and for Marion and Prince Edward Is by Branch

et al. (1993).

Order Aspidochirotida Grube

Synallactidae Ludwig

Bathyplotes hongraini Vaney

Balhyplotes hongraini Vaney, 1914: 5-8, pi. 1 fig. 4,

pi. 2 figs 4, 7-9 and 11-12.

Balhyplotes moseleyi. —Ekman, 1925: 25-28, fig.

3.—Ekman, 1927: 371 [non Bathyplotes moseleyi

(Theel, 1886a); not confirmed whether B. hongraini or

B. gourdoni].

Bathvplotes fuscivinculum GulX, 1990: 121-123, figs

4-6, tW, 11.—Gutt, 1991b: 324.—Gutt and Klages,

1991: 305 [new synonymy].

Balhyplotes sp. MoV 2018.—O'Loughlin et al.,

1994: 553-554.

Material examined. Bathyplotes hongraini Vaney,

1914. Holotype. Western side of Antarctic Peninsula,

68”00'S. 70°20'W, 250 m, 1909, MNHN [exten¬

sively dissected, disintegrating; ossicles few, eroded,

measurable],

Bathyplotes fuscivinculum Gutt, 1990. Paratype.

Weddell Sea. ■70-27'S, 8"40 W, 329-350 m, 1983

[ZIM E-7140 (1 specimen)] [no ossicles were detected].

Other material. Eastern Antarctica, off Kemp Land,

66°2rs, 58°50'E, 603 m, 1930 [BANZARE Stn 34,

302

P. M. O’LOUGHLIN

Table 3. Selected diagnostic characters for Meseres species (x indicates present; - indicates absent; ?

indicates unknown).

Species

distinct rounded

marginal longitudinal

projections muscles

branched gonad

gonad ossicles

tubules

respiratory tentacle

tree ossicles mesh

ossicles

globigerinae x

hyalegerus -

involiitus X

macdonaldi x

occuUatus X

peripatus x

propinquus x

spiciilijenis -

torvus X

trachus x

villosus —

Table 4. Summary of significant new distribution data.

Taxon Previous records (authors in text) New records (this paper)

Chiridota pisanii Ludwig

Baihyplotes bongraini Vaney

Balhyplotes goiirdoni (Vaney)

Meseres hyalegerus Sluiter

Meseres involulus Sluiter

Meseres peripatus Sluiter

Meseres villosus (Theel)

Mesothuria bifurcata Herouard

Heterocucumis godeffroyi

(Semper)

Psoticnvc coatsi (Vaney)

Staurocucumis liouviltei (Vaney)

Trachytbyone lechleri (Lampert)

Amperima robusta (Theel)

Rhipidothuria racovitzai

Herouard

Laetmogone wv\’illethomsoni

Theel

Magellanic region

western Antarctica

Indonesia

Cosmopolitan, western Antarctica

western Antarctica, North Atlantic

Magellanic region, eastern

Antarctica

western Antarctica

Antarctica

Magellanic, Heard Island

Antarctica, Kermadec Trench

western Antarctica

Southern Ocean, western

Antarctica

Heard Island

eastern Antarctica

Japan, south-eastern Australia

eastern Australia, Tasman Sea

North Atlantic, Indo-Pacific

eastern Antarctica

Heard and Kerguelen Islands

off Bouvet Island

Heard I.

Kerguelen 1.

southern Indian Ocean

eastern Antarctica

SAM KI836 (2)]; Prydz Bay, 66*46'-68'55'S,

67*33'-78*l4 E, 120-768 m [ANARE. NMV F68057

(1), F6807() (I), F68159-68160 (7), F68165 (1),

F68167 (1), F68675 (I), F76574-76575 (2),

F76577-76579 (3). F76587-76588 (2), F76599-

76600(2). F80I83 (2). F8I796-8I8(M (37),

F81818-81823 (42). F91299 (I)].

Distribution. Western Antarctica, Weddell Sea,

245^65 m (Gutt, 1991b, as B. fuscivinculum);

western side of Antarctic Peninsula, 250 m

(Vaney, 1914). Eastern Antarctica, Prydz

Bay, 120-768 m; off Kemp Land. 603 m (this

paper).

Remarks. Ekman(1925, 1927) considered B. hon-

graini to be a junior synonym of the non-Antarc¬

tic B. moseleyi (Theel). A distinctive diagnostic

feature of B. moseleyi is the presence of raised

round “warts” or “fungiform papillae” on the dor¬

solateral and ventrolateral interradii, and these are

BANZARE AND ANARE HOLOTHUROIDEA

303

not present in any of the Antarctic specimens

described or examined in this study. On this basis

tlie synonymy of B. bongraini with B. moseleyi by

Ekman (1925, 1927) is rejected. The type locality

of B. moseleyi (Theel, 1886a) is off southern

Chile. Rowe (in Rowe and Gates, 1995) regarded

B. moseleyi as a Junior synonym of B. nalaiis

(Sars).

Gutt (1990) acknowledged difficulty in diag¬

nosing difference between B. rubipunclaliis and

B. fiiscivinculiim, and depended on recognition

by live colour which is itself variable. Gutt

and Klages (1991) acknowledged difficulty in

distinguishing B. ruhiptmclatus and B. fuscmricu-

liim from B. moseleyi. Based on the original

descriptions and figures by Vaney (1914) and

Gutt (1990), and the material examined, the

distinctive features of B. bongraini are: large

body size (up to 260 mm long); distinct ventro¬

lateral brim with papillae; presence of two mid-

ventral radial series of tube feet which are

smaller and more numerous than in eaeh of the

lateroventral series; some bare ventral inter-

radial areas; absence of warts; absence of

C-shaped ossicles; table ossicle bases with pre¬

dominantly four amis, very rarely up to six, arms

up to 0.10 mm long [up to 0.07 in holotype;

Vaney (1914) illustrated 0.07 mm; Gutt (1990)

illustrated 0.12 mm and reported 0.13 mmj. The

numbers of dorsal papillae and ventral tube feet

vary with size, and the numbers given by Gutt

(1990) for various sizes af B. fiiscivinciilum are

comparable with numbers for ANARE material

and the illustration of the holotype by Vaney

(1914). The residual dark colour spots noted

by Vaney (1914) are present on some of the

ANARE material. B. fiiscivinciiliim is judged

here to be a junior synonym of B. bongraini,

which is in turn distinct from B. moseleyi and

other species of Balhyploies. Ekman (1925. 1927)

reported B. moseleyi from western and eastern

Antarctica, but descriptive detail is inadequate to

distinguish whether the species was B. bongraini

or B. goiircloni. B. bongraini occurs in eastern

and western Antarctica.

Bulhyploles gourdoni (Vaney) comb. nov.

Svmillacles (?) gourdoni Vaney, 1914: 4-5. pi. 2 figs

1 -.3. 5. 6.

Balhyploles moseleyi. —Ekman, 1925: 25-28, fig.

3.—Ekman. 1927: 371 [non Balhyploles moseleyi

(Theel. 1886a); not confirmed whether B. bongraini or

B. goiir(loni\.

Ballivploles rubipunclaliis GiM, 1990: 120-121, figs

1-3, tb’l. I.—Gutt, 1991b: 324.—Gutt and Klages, 1991:

301-305, figs 1-3 [new synonymy].

Malerial examined. Balhyploles rubipunclaliis Gutt,

1990. Paratypes. Weddell Sea, 74”57'S, 60”3rW,

646-661 m [ZIM E7138 (3)] [no ossicles were

detected].

Other material. Prydz Bay, 67”0r-68"0rS,

70”46'-77M9'E, 304-341 m [ANARE. NMV F68157

(1). F91296(2)].

Dislribiilion. Western Antarctica, Weddell Sea,

225-840 m (Gutt, 1991b, as B. rubipiinctatii.s)-,

western side of Antarctic Peninsula, 250 m

(Vaney, 1914). Eastern Antarctica, Prydz Bay,

304-341 m (this paper).

Remarks. Ekman (1925, 1927) considered the

single very damaged type specimen of 5. (?)

gourdoni, collected close to the same location and

depth as B. bongraini. to be conspccific with B.

bongraini and then considered B. bongraini to be

a junior synonym of the non-Antarctic B. mose¬

leyi (Theel). The synonymy of fi. bongraini with

B. moseleyi is rejected above, and the synonymy

of S. (?) gourdoni w'ith B. bongraini is rejected

here. Based on the original descriptions and

figures by Vaney (1914) and Gutt (1990), and the

material examined, the features which distinguish

S. (?) gourdoni from B. bongraini are: smaller

size (up to 120 mm long); residual pale pink pre¬

served colour sometimes evident; gelatinous body

wall, and consequent frequent loss of appendages;

inconspicuous brim and ventrolateral edge; mid-

ventral radial tube feet sometimes extending onto

ventral interradial areas, frequently creating a

complete ventral cover; table ossicle bases with

frequently four anus, up to eight and some

branching, arms up to 0.20 mm long (Vaney,

1914, illustrated 0.12 mm; Gutt, 1990, illustrated

0,13 mm and reported 0.11 mm). S. (?) gourdoni

is referred here to Balhyploles, and B. riibipiinc-

tatus is considered here to be a junior synonym of

B. gourdoni. B. gourdoni occurs in eastern and

western Antarctica.

Meseres Ludwig

Meseres Ludwig, 1894: 34, 36, 38.—Perrier, 1902:

359.—Rowe (in Rowe and Gates, 1995): 284-285.—

O’Loiighlin. 1998:497.

Molpadiodemas llcding 1935: 77-78.—lleding,

1940: 357.—Deichmann, 1940: 209-211.

TrachosUchopus llcding, 1940: 357, 361.—Imaoka,

1978: 380.—Imaoka, 1990: 152.

Blicaslichopiis lleding, 1940: 357.—lleding, 1942:

5-6.

Comparalive malerial examined. Pseudoslichopus mol¬

lis Theel. 1886 [type species of Pseudoslichopus Theel,

1886], Syntype. Southern Ocean, off Marion I.,

46‘48'S, 37"49'E, 92-137 m. Challenger Stn 144a

[BMNH 86.10.2.144(1)].

304

P. M. O’LOUGHLIN

Type species. Meseres macdonaldi Ludwig, 1894

[monotypy]; type locality: North Pacific Ocean,

off Costa Rica (2149 m) and Colombia (1644 m).

Emended diagnosis. Up to 300 mm long; body

wall soft, fragile; body rounded anteriorly and

posteriorly, Hat ventrally, convex dorsally, dis¬

tinct lateroventral margin defined by acute angle

or concentration of largest tube feet or series of

pyramidal projections (“warts”); posterior

vertical (pygal) furrow; mouth and anus ven¬

tral; calcareous ring solid, lacking posterior

prolongations; up to 20 peltate tentacles; lacking

tentacle ampullae; lacking retractor muscles;

gonad tubules in 2 series on opposite sides of dor¬

sal mesentery; respirator^’ trees well developed;

radial tube feet very small, most conspicuous on

ventrolateral margin; thin tubular appendages

frequently present dorsally, laterally, ventrally.

around mouth and anus, some branched, some¬

times matted and entangled; ossicles present in

tentacles and some tube feet, sometimes present

in gonads and respiratory trees, absent from body

wall except for large perforated plates or three-

dimensional mesh ossicles sometimes present in

posterior lobes; body frequently obscured by

cover of sponge spicules or globigcrines or

detritus.

Species. M. globigerinae (Herouard, 1923), M

hyalegenis Sluiter, 1901; A/, involutus Sluiter,

1901; A/, macdonaldi Ludwig, 1894; M. occulta-

liis (Marenzeller, 1893); A/, peripatus Sluiter,

1901; A'/, propinqnus (Fisher, 1907); A/, spi-

cidifenis sp. nov.; A/, tonnis (Thcel, 1886); A-/,

trac’/;)/.v (Sluiter, 1901); A/. v)7/a«w (Theel, 1886).

Disirihiiiion. Cosmopolitan, including Antarctic

coast; bathyal and abyssal.

Remarks. The diagnostic characters of the type

material are incompletely known, and are dis¬

cussed below under M. macdonaldi. Unbranched

or branched gonad tubules, rounded or broad flat

longitudinal muscles, and the pre.sence or absence

of ossicles in gonads and respiratory trees, are

considered here to be diagnostically significant

within the genus (Table 3).

Determination o(Meseres material is made dif¬

ficult by: .specimens completely obscured by an

actively created cover (distinguished from pas¬

sive adherence of substrate matter) of fur-like

sponge spicules or globigerines or other detrital

material; outer attached cover frequently lost dur¬

ing collecting, resulting in loss of outer body wall

with tube feet and ventrolateral margin features;

variable degree of development of “teeth” on

the posterior edge of calcareous ring radial plates

[observations in this work confirmed by illu.s-

trations for Pseudostichopus (Trachostichopu.s)

trachus by Heding (1940)]; gonad tubules lost,

or absent, and gonad ossicles sparse or absent

in species in which gonad ossicles occur;

ossicles in posterior lobes obscured amongst

embedded detrital material; longitudinal muscles

may or may not show evidence of longitudinal

division within a species (type specimens of

A/, involutus have both grooved and ungrooved

muscles).

A soft fragile body wall; lateroventral margin

marked by pyramidal projections or a band of

small tube feet or an acute angle; and thin tubular

appendages around the mouth, in the pygal

furrow, and over the body, distinguish Me.seres

from the closely-related genus P.seudostichopus

Theel, 1886. The syntype of P. mollis (type

species of Pseudoslicliopu.i) had a fimi leathery

body wall, was rounded in transverse section,

lacked a distinct lateroventral margin, had very

small tube feet present in sparse double series

along the dorsolateral radii only, and lacked thin

tubular appendages. Theel (1886a) was uncertain

about the lateroventral distribution of tube feet in

the syntypes of P. mollis, and refened to thetn as

“more crowded in two to five (?) rows”. This

description is not valid for the syntype from off

Marion I. There may be more than one species

among the syntypes. A complete review of Pseu¬

dostichopus and Meseres and establishment of

lectotypes for their species where nece.ssary are

beyond the scope of this work.

In a review of P.seudostichopus and related

genera Heding (1940) did not include Meseres.

and described a new genus Plieastichopus and the

subgenera P.seudostichopus and Trachostichopus

(for Pseudostichopus). Rowe (in Rowe and Gates,

1995) synonymised Trachostichopus Heding,

1940 and Plieastichopus Heding, 1940 with

Meseres, assigning P.seudostichopus (Trachosti¬

chopus) trachus Sluiter, 1901 and Plieastichopus

ingalfi Heding, 1942 to Meseres. Heding (1940)

assigned Pseudostichopus trachus Sluiter, 1901.

P. propinqnus Fisher. 1907, P. aleutianus

Ohshima. '915, P. arenosu.s Ohshima. 1915.

P. molpadioides Ohshima, 1915, and P. nudus

Ohshima, 1915 to Pseudostichopus (Trachosti¬

chopus). P. propinqnus is assigned below to

Meseres. P. aleutianus is synonymised below

with A'/, peripatus Sluiter, 1901. P. molpadioides

Ohshima. 1915 [off eastern Japan, 714 m], with

probably junior synonytn P. arenosus Ohshima,

1915 [off southern Japan, 781ml, were described

BANZARE AND ANARE HOLOTHUROIDEA

305

as being cylindrical with a brown globigerine

cover, distinct dorsolateral and ventrolateral tube

feet, and spinous perianal ossicles. In the absence

of a distinctive ventrolateral margin they are

assigned to Pseiuhsticfwpiis. A specimen taken

off eastern Australia at 3500 m. held in the Aus¬

tralian Museum (AM J23009), has similar diag¬

nostic characters. P. mi Jus Ohshima, 1915 was

described as subcylindrical, lacking a distinct

ventrolateral margin, and is assigned to Pseudos-

tichopus. Heding (1940) referred P. ingotfi l led-

ing, 1940 (ms.) and P. plicatus (Koehler and

Vaney, 1905) to Plicaslichopus fleding, 1940.

Both species arc synonymised below with

M. peripaltis.

Heding (1940) assigned Pseudostichopus

allcwlicus Perrier, 1898, P. occullalus Maren-

zeller, 1893 and P. villosus Theel, 1886 to

Molpadiodemas Heding, 1935 (type species

M. acaudum Heding. 1935). O’Loughlin (1998)

synonymised Molpadiodemas with Meseres, and

assigned A/, acaudum, P. utlanticus, P. occullalus

and P. villosus to Meseres. The assignment of

P. allaulicus Perrier and M. acaudum Heding to

Meseres is not in accord with the emended

diagnosis of Meseres above, and these two

species are synonymised and assigned below to

Pseudoslichopus.

Imaoka (1978, 1990) referred five new species

to Pseudoslichopus: P. (Pseudoslichopus) dila-

lorbis Imaoka, 1978; P. (Trachoslichopu.s) lachi-

maniac Imaoka, 1978; P. (Trachoslichopus)

japouensis Imaoka, 1978; P. (Pseudoslichopus)

alaltis Imaoka, 1990; P. (Trachoslichopus) luber-

ciilalus Imaoka, 1990. P. (P.seudosUchopus) dila-

lorbis [off south-western Japan, 400-450 m]

lacked a distinctive ventrolateral margin and is

appropriately referred to Pseudoslichopus.

P. (Trachoslichopus) lachimaruae and P. (Tra¬

choslichopus) japouensis are synonymised below

with M. hyalegerus. P. (Pseudoslichopus) alalus

[off southem Japan, 350 m] lacked a distinctive

latcroventral margin and is confirmed as Pseu¬

doslichopus. This species is similar to P. arenosus

and P. molpadioides, also reported off Japan by

Ohshima (1915), but Imaoka (1990) did not find

spiny perianal ossicles in P. alalus. P. (Trachosli¬

chopus) luberculaius is synonymised below with

M. peripaltis.

The new combinations Meseres globigerinae

and M. lon’us are discussed below. M. spi-

culiferus sp. nov. and M. villosus occur on the

eastern Antarctic coast. M. irachus and M. villo¬

sus have been reported by a number of authors

(see below), but frequently with insufficient detail

to confinn which species of Meseres was being

reported. Ludwig (1894) detennined North

Atlantic material (2-6’N, 81-86°W, 245-3336 m)

as P. mollis, but illustrated peripaliis-Vike gonad

ossicles which P. mollis lacks.

Meseres globigerinae

(Herouard) comb. nov.

Pseudoslichopus globigerinae Herouard, 1923:

23-25, pi. 4 fig. 6.—Mortensen. 1927: 386-388.—

Deichmann, 1930: 87,90.

Pseudoslichopus (Pseudoslichopus) globigerinae .—

Heding, 1940: 353-360.—Imaoka. 1978: tbl. 1-1.—

Thandar, 1992: 167.

Pseudoslichopus villosus. —Hansen, 1956: 47-48

(part) [non Meseres villosus (Theel. 1886)].

Tvpe localilv. North Atlantic, Bay of Biscay,

4380 m.

Remarks. Features of P. globigerinae described

by Herouard (1923) were; up to 30 mm long; thin

soft translucent body wall, covered with globiger-

ines; pygal furrow; mouth and anus ventral; vil¬

lous-like cover of unequal tube feet, except mid-

ventrally; posterior margin with tufts of tube feet

on papillae; absence of ossicles. The description

of the posterior margin indicates that diagnosti¬

cally P. globigerinae is a Meseres species, and it

is so assigned here. Hansen (1956) synonymised

P. globigerinae with P. villosus, but the tufts of

tube feet grouped on papillae along the posterior

margin, cover of unequal tube feel, globigerine

cover, and translucent body wall indicate that

P. globigerinae is not M. villosus and it is

removed from the synonymy here. The absence of

descriptions of the fonn of the gonad tubules and

longitudinal muscles, and presence or absence of

ossicles in gonads and respiratory trees, leave any

synonymy unresolved.

Meseres hyalegerus Sluiter

Meseres hvalegerus Sluiter, 1901a; 12.—Sluiter,

1901b: 50-51, pi. 5 figs 2^.—Perrier, 1902: 359.

P.seudosUchopus Irachus. —Mitsukuri. 1912: 3-9. pi.

1 figs 1-5.—Ohshima. 1915: 227-228.—Mortensen,

1918: 80-81. fig. 16 [non Me.seres Irachus (Sluiter.

1901)].

Pseudoslichopus (Trachoslichopus) lachimaruae

Imaoka. 1978: 380-382, fig. 2A-E. tbl. 1-2 [new syn¬

onymy).

Pseudoslichopus (Trachoslichopus) japouensis

Imaoka. 1978: 382-384, fig. 3A-D, tbl. 1-2.—Imaoka,

1990: 148 [new synonymy].

Maierial e.xainined. Syntypes. Indonesia, Banda Sea,

5'’28'S. 132°00'E, 204 m, Sihoga Stn 251 [ZMA 2177

(2 specimens)].

306

P. M. O’LOUGHLIN

Other material. Sagami Sea, south of Honshu.

128-553 m [Alhalross Stn 5092. USNM E17147 (47);

Stns 4968. 5069, 5093. 5055. 5094, USNM

1001645-1001649 (15)]. Eastern Australia. Tasman

Sea and Bass Strait, 293-530 m [NMV F80171 (2),

F80172 (12). F80I73 (I). F80I74 (5). F80177 (1).

F80I78 (3). F80I79(3). F80I80(2), F80181 (19)].

Distribution. Indo-Pacific. Indonesia, Banda Sea,

204 m (Sluiter, 1901a); Japan, Sagami Sea,

141 564 m (Mit-sukiiri. 1912, as P. trachus)\

south of Honshu, 128-553 m (Ohshima, 1915, as

P. trachus)\ eastern Australia, Tasman Sea and

Bass Strait, 293-530 m (this paper).

Rewarhi. The syntypes of A/, hyalegerus were

collected at 204 m. Based on Sluiter (1901a,

1901b) and the syntypes studied, the features of

M. hyalegerus are: largest was 100 rntn long; all

were covered with sponge spicules and other

detrital matter, not predominantly globigerines;

gonad tubules were unbranched, rarely bifid dis-

tally; lateroventral margin was subacute, and

lacked small projections; ossicles comprising

large open and close mesh bodies were present in

the posterior lobes; tentaele ossicles were spinous

rods up to 0.34 mm long, lacking a large central

swelling and rarely branched. In contrast the syn¬

types of P. trachus (see below) were collected at

798-883 tn; largest was 180 mm long: all were

covered with globigerines; gonad tubules were

long and thin; lateroventral margin was acute,

with small projections ("warts"); ossicles were

not detected in the posterior lobes; tentacle ossi¬

cles were thorny rods up to 0.20 mm long (0.13

mm in Sluiter. 1901b). frequently branched and

with a central swelling. Material from the Sagami

Sea, determined as Pseudostichopus trachus and

described by Mitsukuri (1912), was collected at

141-564 m; largest was 120 mm long; ail were

covered with sponge spicules and other detrital

material; gonad tubules were short, if branched

only at the ends; no reference was made to lat¬

eroventral projections; close and open-mesh lat¬

ticework ossicles were present posteriorly; tenta¬

cle ossicles (illustrated) were spinous rods, up to

0.30 mm long, lacking central swellings. The

Sagami Bay specimens of Mitsukuri (1912) and

Mortensen (1918), and the specimens from south

of Honshu of Ohshima (1915), are considered

here to be A/, hyalegerus. Two species described

by Imaoka (1978), P. (Tracho.stichopus) tachi-

maruae [off south-western Japan, 400-450 m]

and P. (Trachostichopus) Japonensis [off western

Japan, 200-300 m], were both charaeterised by

small size, dense cover of sponge spicules, dorso¬

lateral and ventrolateral series of small tube feet.

unbranched gonad tubules lacking ossicles, and

mostly unbranched tentacle rods more than 0.2

mm long and lacking a distinct central swelling.

Both species are synonymised here with Af.

hyalegerus. Sometimes tube foot rods were

detected during this study in M. hyalegerus

[reported for japonensis by Imaoka (1978)] and

posterior ossicles [fragments reported for tachi-

maruae by Imaoka (1978)].

Meseres involutus Sluiter

Figure 2e

Meseres involutus Sluiter, 1 90 la: 11-12.—Sluiter,

1901b: 49-50, pi. 8 fig. 6.—Perrier, 1902: 359 (incor¬

rectly as convolulu.'i).

Material e.xamined. Syntypes. Indonesia, Seram Sea,

3’27 S, nrOTE, 567 m. Sihoga Stn 173: Sawu Sea,

10‘49 S. 123’23’E. 918 m. Siboga Stn 300 [ZMA (2)|.

Other material. Eastern Australia. Tasman Sea. Lord

Howe Rise. 1423 m [AM J23326 (1)]; off Newcastle,

2984-3058 m [AM JI6833 (I)[; off Nowra, 1650-1750

m [NMV F80451 (I)].

Distribution. Indonesia. Seram and Sawn Seas,

567-918 m (Sluiter. 1901a); eastern Australia,

Ta.sman Sea, 1423 3058 m (this paper).

Remarks. All live specimens were covered with

globigerines, not sponge spicules. Most W'cre cov¬

ered with thin tubular appendages, most conspic¬

uous laterally, smallest ventrally. The grey body

wall with surface ridges and pockets and lumps,

and finely serrated laterovetitral tnargin created

by transverse ridges, was similar to M. peripatus,

but the marginal projections were not as pro¬

nounced as in some M. peripatus specimens. M.

involutus is distinguished from A/, peripatus by

the font! of the tentacle ossicles (include elongate,

curved, distally tapered rods with mesh-like per¬

forations); by branched gonad tubules; by broad

fiat longitudinal muscles; and by the absence of

ossicles in the gonads. Posterior lobe ossicles

were not detected. The longitudinal muscles were

fiat, some with two to three longitudinal grooves.

Meseres inacdonaldi Ludwig

Meseres macdonahii Ludwig, 1894: 34-36.— Perrier,

1902: 359.—Rowe (in Rowe and Gates, 1995); 284.—

O'Loughlin. 1998:497.

Material examined. Meseres macJonaldi Ludwig,

1894. Synlype. Northern Pacific Ocean, off Costa Rica,

5‘56'N. 85*10 W, 2149 m. Albatross Stn 3362. 1891

[USNM 30501 (I)].

Type localit}'. North Pacific Ocean, off Costa Rica

(2149 m) and Colombia (1644 m).

BANZARE AND ANARE HOLOTHUROIDEA

307

Remarks. The six syntypes of the type species

M macdamildi are in an advanced stage of

decomposition (C. Aheam, USNM, pers. com.).

Ludwig (1894) described maximum length as 30

mm; some covered with globigerincs; body flat¬

tened dorsoventrally. with distinct edge: mouth

and anus ventral; body wall thin, transparent; very

small tube feet distributed thickly over whole

body; 15 tentacles; tentacle ampullae absent; tufts

of unbranched gonad tubules on both sides of dor¬

sal mesentery; respiratory trees well-developed;

longitudinal muscles undivided; ossicles not

found in body wall or tube feet or tentacles. Lud¬

wig (1894) did not report on ossicles in gonads or

respiratory trees or posteriorly, and did not report

a pygal furrow. In the syntype examined in this

study no ossicles were found, and a pygal furrow

was not evident in the disintegrating material.

Two series of lateroventral pyramidal projections

were present, each with about three very small

digitate projections closely resembling the illus¬

tration of S. lorvus by Theel (1886a) which was

referred to by Ludwig (1894) in describing

M. macdomddi. There were very small tube feet

and thin tubular appendages over the body. The

longitudinal muscles were cylindrical. Gonad

tubules were not detected.

M. occtillaliis. M peripaliis and M propmqiius

share with A/, macdomddi having unbranched

gonad tubules, rounded longitudinal muscles, and

prominent lateroventral projections, but it is not

possible to conllnn any synonymy in the absence

of observations on gonad and respiratory tree

ossicles in M. macdomddi.

Mescrcs occidtatus (Marcnzeller)

P.seiido.siicliopii.s occtdialu.s Marcnzeller, 1893a;

15-17, pi, 4 fig, 9.—Maren/cllcr, 1893b: 10, pi. 2 fig.

3.—Perrier, 1902 ; 337-338,—Hcrouard, 1902; 14-15,

pi. 2 figs 4-14.—Mortensen, 1918; 81.— Mortensen,

1927; 387-388.—Deichmann. 1930: 89-90.

Molpadiodcimi.s occuhatiis. —lleding, 1940: 353-359.

Mesvres uccultatus. —O'Loughlin. 1998: 497.

Material examined. Syntypes. Mediterranean Sea,

24"I6'N, 35"03'E. 1445 m [MNllN Ecllh 3658 (2)].

Distribution. Mediterranean Sea, 415-1445 m

(Marcnzeller, 1893a); North Atlantic, Spain, off

Cape Finisterre, 363-510 m (Marcnzeller,

1893b); balhyal.

Remarks. Based on the description and illustra¬

tions by Marenz.ellcr (1893a, 1893b), and exam¬

ination of two syntypes, two unique diagnostic

characters for M. occidtatus are the presence of

branched ossicles in the respiratory trees, and not

the gonads, and the presence of large posterior

ossieles which are large partly double-layered

perforated plates and not multilayered mesh.

Otherwise the active attachment of detrital mate¬

rial; grey parchment-like body wall; reticulate

outer body wall with lateroventral marginal pro¬

jections; complete cover of thin tubular

appendages; double series of small tube feet lat-

erovenlrally; rounded longitudinal muscles;

unbranched gonad tubules; and fonn of ossicles in

lube feel, are similar to M. peripatus. flerouard

(1902) detemiined material as P. occidtatus, but

subsequently (1923) referred the material to two

new species (P. marenzeUeri and P. lapidus).

P. marenzeUeri is synonymised below with M.

peripatus. Deichmann (1930) determined an

Atlantic specimen from Cuba (232 m) as

P. occidtatus, but described complex branched

gonad ossicles. This character indicates that it

w'as not M. occidtatus, but data are insufficient to

confimi any synonymy.

Meseres peripatus Sluiter

Figure 2f

Meseres peripaliis Sluiter, 1901a; 10-11.—Sluiter.

1901b: 48-49, pi. 5 fig. 5. pi. 8 fig. 7.—Perrier, 1902:

359.—Rowe (in Rowe and Gates, 1995): 285.

Pseiidosliehopiis occullatus. —Herouard, 1902:

14-15, pi. 2 figs 4-14 (part, illustrated) [non Meseres

ocetillalti.s (Marenzeller. 1893)].

Pseiidosliehopiis oceiillaliis var. pliealiis Koehler and

Vaney, 1905; 9 -10. pi. 3 fig. 8, pi. 9 figs 1-3.—Heding,

1940: 353 [non kieseres occiillaliis (Marcnzeller,

1893)].

Pseiidosliehopiis aleiiliaiiiis Ohshima. 1915: 228, pi.

8 figs 5a- c.—Imaoka. 1978: 380.

Pseiidosliehopiis iingiiieiilaltis Ohshima, 1915:

230-231. pi. 8 fig 7a-c.—Imaoka. 1978: 384.—Rowe

(in Rowe and Gates. 1995): 285 [synonymy by Rowe

(in Rowe and Gates. 1995)].

Pseiidosliehopiis marenzeUeri Herouard, 1923: 25.—

Mortensen, 1927: 287-288.—Deichmann, 1930:90.

Pseiidosliehopiis (Pseiidosliehopiis) marenzeUeri .—

Heding, 1940: 353-359.—Imaoka. 1978; tbl. 1-L—

Thandar. 1992: 167 [new synonymy],

Pseiidosliehopiis (Pseiidosliehopiis) iinguieiilaliis. —

Heding. 1940: 35.3-360.--Imaoka. 1978: tbl. 1-1.—

Imaoka. 1990: 152.—Thandar. 1992: 167.

Pseiidosliehopiis (Traehosliehopiis) aleiilianiis .—

Heding. 1940: 353-359.—Imaoka. 1978: tbl. 1-2 [new

synonymy].

Plieasliehopiis pliealiis. —Heding, 1940: 354-359.—

Heding, 1942: 6 [new synonymy],

Plieasliehopiis ingolfi Heding. 1942: 5-6, figs 4-5,

pi. 1 figs 4-5.

Pseiidosliehopiis (Traehosliehopiis) liihereiilaliis

Imaoka, 1990: 149-152, pi. p. 149, fig. p. 151 [new

synonymy].

308

P. M. O’LOUGHLIN

Mcserex ingolfi. —Rowe (in Rowe and Gates, 1995):

285 [new synonymy].

Material exammed. Meseres peripalus Sluiter, 1901.

Syntypes. Indonesia, Flores Sea, 7”24'S, 118’15 E, 794

m, Sihoga Stn 45: Maluka Sea, r59'N, I25"0I E. 1200

m, Siboga Stn 122 [ZMA (2)|.

Other material. Eastern Au.stralia. Tasman Sea. off

New South Wales. 823-1750 m [AM J20026 (2),

J20027 (1). J22980 (9). J232I9 (I). J23267 (2); NMV

F80449 (1), F80450 (3). F90()70 (2)].

Pseudostichopus tmgiiiciilaliis Ohshima. 1915. Syn¬

types. Off southern Japan. 1058-1680 m, [Albatross

Stns 4960. 5083. 5084. USNM E34151(2), E24543 (I).

E24544 (I )J.

Pseudostichopus marenzelleri tlerouard. 1923. Nortli

Atlantic, 36'58' N. 26”20’W. 2871-2917 m [MNHN

EcHh 6073 (I)].

Distribution. Indo-Paciftc. Indonesia, Flores and

Maluka Seas, 794-1200 m (Sluiter, 1901, as

M. pehpatus)\ Bay of Bengal, 3009 m (Koehler

and Vaney, 1905. as P. plicatus): olT southern

Japan. 1058-1680 m (Ohshima. 1915, as P.

ungidculitlus), 660-700 m (Imaoka. 1990. as P.

tuberculatus)’, Aleutian Is, 1019-1398 m

(Ohshima, 1915, as P. aletiiicintt.s): eastern Aus¬

tralia, Tasman Sea. 823-1750 m (this paper).

North Atlantic, off Greenland. 2137-3192 m

(Fleding, 1942. as P. ingolfi): between Portugal

and the Azores, 4400 m (Herouard, 1923, as

P. marenzelleri).

Remarks. The features of M. peripalus shared by

the material e.xamined, and descriptions and illus¬

trations of PUcastichopus plicatus (Koehler and

Vaney, 1905), Pseiidosiicltopus aleulianus

Ohshima. 1915, Pseudostichopus unguiculatus

Ohshima, 1915, Pseudostichopus marenzelleri

Herouard, 1923, PUcastichopus ingolfi lleding,

1942 and P. (Trachostichopus) tuberculatus

Imaoka. 1990 are: up to 135 mm long; body nor¬

mally covered with globigerines; body Hat ven-

trally, domed dorsally, acute lateroventral margin

with prominent serrations; body wall soft, thin,

grey, scmi-translueent; reticulate body ridges and

large to small pockets, ridges with lumps and

flaps with very small digitate projections, ridges

projecting at margin to create prominent irregular

pyramidal projections (conical W'arts); ver}' small

thread-like appendages variably present over

whole body, prominent around mouth and anus,

frequently massed, some branched; small radial

tube feet, in irregular paired series dorsolaterally

and lateroventrally (ventral to marginal projec¬

tions), not detected midvcntrally; calcareous ring

radial plates with minute teeth on posterior inden¬

tation; longitudinal muscles narrow, rounded,

lacking divisions; up to 20 tentacles; gonad

tubules unbranched; tentacle ossicles rods, fre¬

quently bluntly spinous, frequently with central

swelling, rarely branched, up to 0.28 mm long;

lateroventral tube feet with endplates up to 0.12

mm wide, comprising central irregular perforated

plate intergrading with surrounding mesh of short

nodular rods, sometimes with support rods up to

O. 14 mm long: dorsolateral tube feet lacking end-

plates, with support rods up to 0.10 mm long;

gonads with ossicles, commonly rods or X- or

Y-shaped, smooth to spinous, up to 0.18 mm

long, rarely irregularly branched with branches

sometimes joined; ossicles absent from body

wall, posterior lateral lobes, respiratory trees and

thread-like appendages.

The diagnostic characteristic of M peripalus is

a combination of distinctive lateroventral body

margin, and presence of o.ssicles in unbranched

gonad tubules. The distinctive margin was illus¬

trated by Sluiter (1901b. for M. peripalus), by

Koehler and Vaney (1905. for P. plicatus), by

lleding (1942, for P. ingolfi) and by Imaoka

(1990, for P. tuberculatus). Ludw'ig (1894)

referred to the illustration of 5. tonus by Theel

(1886a) in the description of A/, macdonaldi. The

type of M. macdonaldi has this distinctive margin,

but data are insufficient to confinn a synonymy

with A/, peripalus. M. involutus has this fomi of

margin to a lesser degree, but has branched gonad

tubules lacking ossicles, and fiat longitudinal

muscles. M. lon-us has this form of margin, but

has branched gonad tubules and other data are

insutlicient to confinn a synonymy. The gonad

ossicles are illustrated by Herouard (1902, for

P. marenzelleri as P. occultatus), by Koehler and

Vaney (1905, for P. plicaiit.s), by Ohshima (1915,

for P. aleutianus and P. unguiculatus), and by

Heding (1942, for P. ingolfi).

Herouard (1902) incorrectly detemiincd mater¬

ial from the North Atlantic as P. occultatus

Marenzeller, but subsequently by his own admis¬

sion (1923) described the material as the two

species P. marenzelleri and P. lapidiis. Herouard

(1902. as P. occultatus) illustrated P. marenzel¬

leri, and showed branched ossicles in the gonads.

The description of .V/. viUosus (Theel) by

Herouard (1902) from the North Atlantic

(3745-4360 m) refers to gonad ossicles in

unbranched tubules, indicating that the material

was not A/. viUo.sus [see below] and was possibly

M. peripalus. But the size (up to 170 mm) and

absence of the distinctive lateroventral margin

make a synonymy uncertain. The North Atlantic

specimen detemiincd as P. marenzelleri (EcHh

6073) and examined here is identical in body

BANZARE AND ANARE HOLOTHUROIDEA

309

forni with Indo-Pacific specimens of M peripa-

tus, but there were no gonad ossicles and the ten¬

tacle ossicles were significantly more irregularly

branched than in M peripatus. The determination

is uncertain. Imaoka (1990) reported and illus¬

trated tube foot plates and anal body wall ossicles

for P. tiiberculaltis which are considered here

to be not holothurian ossicles, and did not find

ossicles in the gonad of the single specimen

described (absence in a single specimen is not

diagnostically reliable). The globigerine-

covered grey translucent body with distinctive

serrated margin, distribution of tube feet,

fonn of gonad tubules and colour photo of the

holotype are diagnostically identical with

M. peripatus.

Mitsukuri (1912) conducted a detailed study of

the holothurians of the Sagami Sea. and reported

Meseres hycilegerus (as P. Irachus) up to depths

of 564 m and probably did not sample depths at

which M. peripatus occurs. M peripatus is

reported olY southern Japan at 1058-1680 m by

Ohshima (1915, as P. unguiculatus) and at

660-700 by Imaoka (1990, as P. tuherculatus).

M. peripatus is a deep bathyal to upper abyssal

cosmopolitan species.

Meseres propinquus (Fisher)

comb. nov.

Pseudostichopiis propinquus Fisher. 1907: 691-693,

pi. 71 fig. 3, pi. 72 fig. 2, pi. 73 fig. 3. pi. 74 fig. 1. pi.

76 fig. 3.—Imaoka, 1978; 382.—Rowe (in Rowe and

Gates. 1995): 285.

Pseudosliciwpus (Tracliostichopus) propinquus .—

Heding, 1940: 357.—Imaoka, 1978: tbl. 1-1.—Imaoka,

1990: 148. 152.

Material examined. Holotype. Hawaiian Is, 21°! 1 N,

156‘35’W, 518-519 m [USNM 21217).

Remarks. The holotype is in a very poor state of

preservation. Based on the description and illus¬

trations by Fisher (1907), this species has the

characteristics of Meseres including a thin

translucent body wall, acute laterovcntral margin

with mammiform tubercles, small radial tube feet,

scattered very small thread-like appendages, and

body cover of sponge spicules and foramini-

ferans. A unique diagnostic character is the pres¬

ence of ossicles in both gonads and respiratory

trees. Rowe (in Rowe and Gates, 1995) syn-

onymised P. propiueptus with Pseudostichopus

pustulosus Sluiter, 1901. P. pustulosus lacks res¬

piratory tree and gonad ossicles, and has multiple¬

branching gonad tubules (Sluiter, 1901a), and the

synonymy is rejected here.

Meseres spiculiferus sp. nov.

Figures la-f, 2a-d

Pseudostichopus sp. MoV 2068.—O’Loughlin et al.,

1994: 253-255.

Material examined. Holotype. Prydz Bay. 67°10’S,

74°28'E, 428 m. T. Bardsley, R. Ickeringill and C. Hay¬

ward, 6 Mar 1997, NMV F81857,

Paratypes (8). Prydz Bay, 66°46'-67’34'S,

70’42'-77“32'E, 298-540 m, 1991, 1997, NMV

F68054 (1), F68156 (1), F72542 (1), F81805 (4),

F81806(l).

Other material. Eastern Antarctica, off Wilkes Land,

65°07’S, 107"29'E, 695 m, 1931 [BANZARE Stn 98.

SAM K1853(l)]; Prydz Bay, 66'48 S. 7r24'E, 456 m,

1929 [BANZARE Stn 30, SAM K1851 (2)]; olTMac-

Robertson Land, 66"45'S, 62'03'E. 177 m, 1931 [BAN¬

ZARE Stn 107, SAM K1852 (7)]; off Endcrby Land,

65°48'S, 53"16 E, 193 m, 1930 [BANZARE Stn 41,

SAM K1850 (1)]; Prydz Bay, Vincennes Bay,

65’33'-68°32'S, 70"20’-108'48'E, 290-600 m

[ANARE, NMV F68153 (2), F68163 (1), F76598 (1),

F81807 (4). F81813-81815 (3), F81824 (1),

F81838-81840 (6), F81860 (1), F81862 (1), F82701

(1), F90071 (1)].

Description. Up to 105 mm long, 27 mm broad,

21 mm high; body wall soft, off-white; body nor¬

mally covered with sponge spicules, sometimes

globigcrines; body Oat ventrally, domed dorsally,

rounded anteriorly and posteriorly, posterior

pygal vertical furrow; mouth, anus ventral; later-

oventral margin semi-acute, rounded, some retic¬

ulate ridges with very small digitate projections,

lacking prominent ventrolateral projections; up to

20 reddish-brown tentacles; very small thread¬

like appendages variably present over whole

body, typically 0.2 mm diameter, prominent

around mouth and furrow, frequently entangled;

small radial tube feet, up to 1.0 mm diameter,

largest along laterovcntral margin in band up to 5

very irregular rows wide continuous around ante¬

rior body, irregular double rows dorsolaterally,

sparse to absent midventrally; longitudinal

muscles rounded, undivided; single ventral polian

vesicle; calcareous ring plates solid, more wide

than high, lacking posterior prolongations, radials

frequently with 4 posterior teeth on edge of inden¬

tation; ossicles in tentacles, tube feet, gonads,

absent from body wall (including posterior lobes),

respiratory trees; tentacle ossicles curved to

straight rods up to 0.28 mm long, frequently with

central rarely distal swellings, very rarely

branched, rarely bluntly spinous; tube feet distal

support rods up to 0.20 mm long, same form as

tentacles; “endplates” up to 0.13 mm wide, open

mesh of irregular thin knobbed branched rods;

310

P. M. O’LOUGHLIN

Figure I. a-f, Meseres spiailifcnis sp, nov. a. dorsal (above) and ventral (below) views of holotype (102 mm long);

b, sponge cover, dorsal (paratype F68054, 30 mm long); c, tentaeles (paralypc F72542, specimen 83 mm long);

d, pygal posterior furrow (paratype F81805, specimen 95 mm long); e, unbranched gonad tubules and rounded

longitudinal muscles (paratype F8I806, specimen 100 mm long); f. laterovenlral tube feet (paratype F68I56,

specimen 54 mm long).

BANZARE AND ANARE HOLOTHUROIDEA

311

Figure 2. a-<i, Meseres spkuHferus sp. nov. a, radial (right) and interradial (left) plates of calcareous ring;

b, midbody section with tube feet distribution; c, tentacle ossicles (largest 0.22 mm across; BANZARE specimens);

d, gonad ossicles (0.16 mm across; paratype F81806), e, Meseres involutus Sluiter. Tentacle ossicles (longest 0.30

mm long; F80451) f, Meseres peripatus Sluiter. Gonad ossicles (longest 0.20 mm long; F80450).

312

P. M. O’LOUGHLIN

gonad tubules with abundant veiy irregular ossi¬

cles, frequently with large central swellings, rod

to X- to Y-shaped to complex branching,

branches sometimes joined to create large

perforations, up to 0.30 mm long.

Etymology. From spiculum (Latin, sharp point)

referring to sponge spicules, and ferre (Latin, to

bear), describing the attached body cover of

sponge spicules.

Distribution. Eastern Antarctica, off Wilkes,

MacRobertson, Enderby Lands, 177-695 m.

Remarks. The distinguishing diagnostic charac¬

ters of A/, spiculiferus are very irregular ossicles

in unbranched gonad tubules, absence of con¬

spicuous lateroventral body wall projections, and

narrow rounded and undivided longitudinal

muscles. Pseudosticbopus atlanticus Perrier.

1898 has similar gonad ossicles, but gonad

tubules are branched and longitudinal muscles

broad and Hat. M. peripatus has gonad ossicles

which overlap in form but are smaller (up to 0.18

mm long), and less branched; has smaller tube

foot support rods (up to 0.14 mm long): and has

prominent lateroventral marginal projections.

In the light of a general circumpolar occurrence

of Antarctic holothurian species, and inadequate

diagnostic detail recorded, material reported from

western Antarctica as Pseudosticbopus mollis

Theel and P. vUlosus Theel was possibly

M. spiculiferus. Gutt (1991a) reported that

“sponge spicules adhere to the body wall of

Pseudosticbopus villosus giving it a furry appear¬

ance”. This is not tnie of A/, villosus in eastern

Antarctica, and the description accurately fits

M. spiculiferus.

Meseres torviis (Theel) comb. nov.

Stichopus (?) lor\’us Theel. 1886a: 164-165, pi. 10

figs 2-4.—Ludwig, 1894; 34.

Type locality. Southern Pacific Ocean, off Chile,

33'42'S, 78"18'W, 2516 m.

Remarks. This species is known from a single

damaged specimen. Theel (1886a) described a

wrinkled body covered with foreign matter, lat¬

eroventral conical projections, 20 tentacles, small

crowded tube feet, solid calcfireous ring lacking

posterior prolongations, and absence of body wall

ossicles. All are features of Meseres, to which

S. tonms is assigned here. Gonad tubules were

branched, and longitudinal muscles not divided.

Reference was not made to ossicles in gonad

tubules or respiratory trees or perianal ly, or to a

pygal furrow which is presumed here to have

been obscured by damage or the cover of foreign

matter. Data are insufficient to confirm any

synonymy.

Meseres traebus (Sluiter)

Pseudosticbopus traebus Sluiter. 1901a: 15-16.—

Sluiter, 1901b; 52-53. pi. 5 fig. 1, pi. 8 fig. 8.—Perrier.

1902; 337-338.—Fisher, 1907:693.—Savel’eva, 1941:

74.—Djakonov. 1952: 127. 129.—Baranova, 1957:

239.—Djakonov et al., 1958; 366.—Imaoka, 1978:

384.—Cherbonnier and Feral. 1981: 383, 385, fig. 16.

Pseudosticbopus (Tracbosticbopus) traebus .—

Heding, 1940: 353-362. fig. 17.—Imaoka, 1978; tbl.

1-2,—Thandar, 1992: 166.

Meseres traebus. —Rowe (in Rowe and Gates, 1995):

285.—O’Loughlin, 1998: 497.

Material e.xamined. Syniype. Indonesia, Arafura Sea,

8’50 S, 127’02’E, 883 m. Sihoga Stn 286 fZMA 2496.1

(I)]-

Other material. Eastern Australia, Tasman Sea,

882-1198 m [AM J16836 (1), J23220 (1), J22957 (I),

J22972 (1), J23218 (1); NMV F80175 (1), F80176 (I),

F80448(l)].

Distribution. Indo-Pacific, Indonesia, Arafura

Sea, 798-883 m (Sluiter, 1901 a); Philippines,

14°N, I20”E, 448-1125 m (Cherbonnier and

Feral, 1981): Sea of Japan, 1600 m (Savel’eva,

1941); off Kuril Is, 113-560 m (Djakonov et al.,

1958); Bering Sea, 110 m (Baranova, 1957); east¬

ern Africa, off Kenya, 638-977 m (Heding,

1940); eastern Australia, Tasman Sea, 882-1198

m (this paper).

Remarks. Rowe (in Rowe and Gates, 1995) reas¬

signed P. traebus to Meseres. Based particularly

on the presence of a series of small but distinct

lateroventral projections on a subacute ventro¬

lateral margin, which were noted by Sluiter

(1901b) and observed here on the type, the reas¬

signment is supported here. Diagnostic characters

for M. traebus are discussed under A/, hyalegerus

above. There is an absence of reference to other

Meseres species, such as M. hyalegerus. and

insufficient diagnostic data in the reports of

P. traebus by Heding (1940), Savel’eva (1941),

Baranova (1957), Djakonov et al. (1958) and

Cherbonnier and Feral (1981), to confirm

their determinations. Depths as shallow as

110 m (Baranova. 1957) and 113 m (Djakonov et

al.. 1958), complete sponge spicule cover

(Savel’eva, 1941), and specimens up to 300 mm

long (Djakonov et al., 1958), raise doubts about

the detenninations. Mitsukuri (1912) and

Ohshima (1915) reported P. traebus for Japan,

but their material is considered above to be

M. hyalegerus.

BANZARE AND ANARE HOLOTHUROIDEA

313

Meseres villosus (Tlieel)

Figures 3a-b

PseudosHchopus villosus Theel, 1886a: 170-171.—

Ludwig, 1898a: 8.—Perrier. 1902: 337-338.—

Herouard, 1902: 11-14, pi. 2 ligs 1-3, pi. 7 fig. 3.—

Vaney, 1908: 407-108.—Grieg. 1921: 4.—Herouard.

1923: 23.—Mortensen, 1927: 387-388.—Deichmann,

1930: 89.—Hansen. 1956: 47-48, 51-53.—GuU,

1991b: 324.

Pscudoslkliopiis villosus var. violaceus Theel,

1886a: 172. pi. 10 fig. 6b.

Molpacliodemcis villosus. —Heding, 1940: 353-360.

PseudosHchopus sp. MoV 2033.—O'Loughlin et al.,

1994: 253-254.

Meseres villosus .—O’Loughlin, 1998: 497.

Material e.xamined. Syntype. Southern Ocean. 46“16'S,

48’27'E, 2928 m. Challenger Stn 147 |BMNH

86.10.2.154(1)].

Other material. Eastern Antarctica, Prydz Bay,

66’46 -68'50'S. 72’I4’-77’I9’E. 333-765 m

[ANARE, NMV F68I52 (9). F68I58 (I). F68162 (5),

F72534 (2), F76583 (2), F76597 (1). F76606 (I),

F8I816-8I8I7 (3)].

Type locality. Syntypes (II) from Atlantic,

Indian, Pacific and Southern Oceans; abyssal.

Distribution. Atlantic, Indian, Pacific and South¬

ern Oceans, 2516-5307 ni (Theel, 1886a). West¬

ern Antarctica, Weddell Sea, 4575-4795 m

(Vaney, 1908); 405^65 m (Gutt, 1991b).

Keimadec Trench, 6660-7000 m (Hansen, 1956);

North Atlantic, 34"59' N, 33"01 ’W, 2615-2965 ni

(Grieg, 1921). Eastern Antarctica, Prydz Bay,

333-765 m (this paper).

Remarks. The distinctive characteristics of M. vil¬

losus are; up to 150 mm long; subcylindrical

fomi; absence of a cover of sponge spicules or

globigerincs or detrital matter; broad band of

numerous larger tube feet lateroventrally (up to 2

mm long); lacking thin tubular appendages; broad

fiat undivided longitudinal muscles; multiple

branching gonad tubules; absence of ossicles in

tube feet, gonad tubules, respiratory trees, poste¬

riorly; dark reddish-brown body colour. Deich¬

mann (1930) thought that material off Morocco

determined by Herouard (1902) as P. villosus was

probably P. atlanticus. but Herouard (1902)

described rod ossicles in unbranched gonad

tubules which suggests M. peripatus [see above).

Deichmann (1930) detennined material from the

French West Indies (896 m) as P. villosus, but

de.scribed tube feet in bundles on distinct lateral

conical warts. This feature and the shallow

bathyal depth indicate that the material was not

M. villosus. Hansen (1956) synonymised

P. glohigerinae with P. villosus, but it is removed

here from the synonymy [see above]. This cos¬

mopolitan abyssal species occurs at shallow

bathyal depths around the Antarctic coast.

Mesothiiria hifurcata Herouard

Mesothuria hifurcata Herouard, 1901: 40.—

Herouard, 1906: 4-6, pi. 2 fig. 3.—Heding, 1942: 8, fig.

7.—O'Loughlin et al., 1994: 553-4.

Material examined. Holotype. Western Antarctica,

Bellingshausen Sea, 7ri4'S, 89"I4’W. approximately

800 111 (estimated from given latitude and longitude),

1898 (IRSNB IG 10131].

Other material. Eastern Antarctica, off Princess Eliz¬

abeth Land. 66‘28 S. 72’4rE, 1266 m, 1929 [BAN¬

ZARE Stn 29, SAM K1838 (1 .specimen)]; Prydz Bay,

66”46’-68’32'S, 70-4r-77’19'E. 320-743 m

[ANARE, NMV F68050 (1), F68154 (1). F72540 (3),

F76594-76596 (3), F81808-81812 (38),

F81858-81859 (5). F81861 (1). F8I863 (2)].

Distribution. Western Antarctica, Bellingshausen

Sea, 800 m (Herouard, 1901). Eastern Antarctica,

off Princess Elizabeth Land, Prydz Bay,

320-1266 m (this paper). North Atlantic,

61'44'N, 30"29'W, 2337 m (Heding, 1942).

Remarks. The holotype has shrunken to 4.6 mm

long, and is in poor condition. No ossicles were

found. The ANARE material is consistent with

the holotype and the descriptions and figure by

Herouard (1901, 1906). The ANARE specimens

have the following characteristics: up to 70 mm

long, subcylindrical. rounded anteriorly, slightly

tapered posteriorly, mouth ventral, anus posterior;

lacking pygal furrow, anal teeth; body wall thick,

fimi, Ilexible; preserved colour grey to reddish-

brown dorsaliy, darker ventrally; up to 20 red¬

dish-brown peltate tentaeles; gonad tubules with

multiple branching; longitudinal muscles divided;

largest tube feet in single ventrolateral row,

smaller tube feet in row lateral to ventrolateral

row; very small tube feet scattered dorsaliy, later¬

ally, few ventrally, absent midventrally; calcar¬

eous ring solid, lacking posterior prolongations,

radial plates as high as wide with anterior

V-shaped notch, interradial plates at least twice as

wide as high; all of body surface with close

bristle of table spires; fonn of table ossicles vari¬

able, normally height of spire similar to width of

disc; largest discs with three large, three smaller

perforations, angular margin, up to 0,23 mm

wide; smaller discs less regular, typically nine

perforations, more rounded margin, typically 0.08

mm wide; spires with three pillars, joined prox-

imally and distally, extending into radiating

tapered spines, height of spires variable up to 0.18

314

P. M. O’LOUGHLIN

Figures, a-b, Mcseres villosiis (Theel), specimen F81816, 88 mm long. a. laterovenlral lube feet; b, branched gonad

tubules and broad flat longitudinal muscle, c-f, Psolicnix coal.ii (Vaney). c, tentacles and elongate thin tail (F68044,

39 mm long); d, sole-like ventrum (F68047, specimen 33 mm long); c above, knobbed crosses (0.07 mm long;

F726I4); e below, plate spires (F68084); f, spired plates (largest 0.51 mm long; F68084).

BANZARE AND ANARE HOLOTHUROIDEA

315

mm; radiating spines variable with single point,

or up to four terminal spinelets, or strongly bifur-

eate with or without terminal spinelets, or doubly

bifurcate.

The description and illustrations of the single

North Atlantic /ngo// specimen by lleding (1942)

are also consistent diagnostically with the

ANARE material, including table size (illustrated

disc 0.12 mm wide, spire 0.13 mm high) and the

occurrence of doubly bifurcate spines minutely

spinous distally. Gutt (1991b) reported

Mesolhiiria laclea (Thcel, 1886) from the Wed¬

dell Sea, but did not report A/, hifurcata. Ossicles

are similar, except for the frequently bifurcate

spines at the ends of the table spires. Weddell Sea

material was probably M. bijiircciia. This near

abyssal Atlantic occurrence and shallow bathyal

occurrence near the Antarctic continent are com¬

parable with the distribution of Meseres villosiis

Theel. The BANZARE and ANARE specimens

extend the distribution to eastern Antarctica.

Pseudosikliopus atlanliciis Perrier

Pseudostichopus atkmtkus Perrier, 1898: 1665.—

Perrier. 1902; 333-338, pi. 17 figs 15-20.—Mortensen,

1927: 386 - 387 .—Deichmann, 1930; 87-88.—Deich¬

mann, 1940: 209, 211.- lleding, 1942: 5.

Molpadiotkimis acaiidtiiii Weding. 1935: 78-80, pi. 6

figs 1-2.-- lleding. 1940: 354-357.—Deichmann,

1940: 209, 211.—lleding, 1942: 4-5.

MolpadioJcimis allanlicii.s.—\\cdmg. 1940: 353-359.

Mesares O'Loughlin. 1998; 497.—

Thandar. 1999: 376-379, fig. 4.

Meseres acciiuhiitt. —O’Loughlin. 1998: 497. fig. 1

g-h [new synonymy].

Material examined I’semloslicliopus allanlicus Perrier.

1898. llolotype. paratype. North Atlantic, olT the

Azores, 42‘ 19' N. 23“36'W, 4060 m. Talisman Stn 134,

1883 [MNIIN Ecllh 2772. Ecllh 658],

Molpadiadeinas acaitdum lleding, 1935. Paratype.

North Atlantie, 60" 17’ N, 54‘05'W, 3230 m, Ingolf Sin

37, 1895 IZMUCj.

Distrihutkm. North Atlantic, 3230-4060 m (Per¬

rier, 1902, as P. iilkinlicus; lleding, 1935, as M.

acaudunt)\ South Atlantic, oil South Alrica,

34"42'S, 16”54'E. 3155-3255 m (Thandar, 1999);

abyssal.

Remarks, lleding (1940) referred Psatdasticho-

pus allaiuiais. P. villosu.s and P. occiillatus to

Molpadiadeinas lleding, 1935 (type species M.

acaudiiin). O'Loughlin (1998) synonymised Mal-

padiodemas with Meseres. Both P. allanlicus and

M. acaudinn lack a distinctive ventrolateral mar¬

gin, and do not belong in the emended diagnosis

of Meseres above. The two species are reassigned

to Pseudostichopiis. Deichmann (1940) and Hed-

ing (1942) considered that P. allanlicus and M.

acaudinn might be synonymous. The types of

both species were taken at abyssal depths in the

North Atlantic; are pale brown in colour, with

very small brown spots (collapsed papillae); have

a large sac-like form, very rugose ventrally, with

some globigerines attached; have a thick body

wall lacking a ventrolateral margin; have a cover

of very small tube feel, frequently withdrawn;

have very broad Hat longitudinal muscles; have

branched gonad tubules with complex ossicles;

and lack ossicles in tube feet and respiratory trees.

Molpadiadeinas acaudinn lleding, 1935 is con¬

sidered here to be a junior synonym of Pseiidosli-

chopus allanlicus Perrier, 1898. Meseres peripa-

tus, M. propinquus and A/, spiculiferus also have

ossicles in the gonads. P. allanlicus is distin¬

guished from M. peripatus which has unbranched

gonad tubules, projections on the laterovcntral

margin, and rounded longitudinal muscles.

M. propinquus has ossicles in the respiratory

trees, marginal projections, rounded longitudinal

muscles, and unbranched gonad tubules.

P. allanlicus and M. spiculiferus have similar

complex gonad ossicles, but M. spiculiferus has

unbranched gonad tubules and rounded longitudi¬

nal muscles. Deichmann (193(J) described a spec¬

imen of P. allanlicus from the British West Indies

(2920 m) as having narrow, undivided, circular

muscle bands, which indicates probable misiden-

tification. The rounded body form and branched

gonad tubules and broad longitudinal muscles of

P. allanlicus arc similar to M. villosus which is

distinguished by a prominent band of ventrolat¬

eral tube feet, and by lacking gonad ossicles. The

“mulberry” bodies from the body wall ot

P. allanlicus, illustrated by Perrier (1902) and

Thandar (1999), and referred to by Theel (1886a)

as present in some specimens of P. villosus, are

considered here to be detrital accretions and not

holothurian ossicles and species specific.

Order Dendrochirotkia Grube

Cucumnriidac Ludwig

Heterocucumis Panning

Ekmocncumis (Helerocucnmis) Panning, 1949: 452.

Type species. Cucuinaria sleineni Ludwig, 1898

[original designation].

Emended diagnosis. Ten equal tentacles; tube

feet on larger specimens in more than single

radial rows; calcareous ring lacks posterior pro¬

longations, frequently soft or absent with age;

316

P. M. O’LOUGHLIN

significant change in ossicle form with age; plates

with narrow extension at some developmental

stage; plates in larger specimens single-layered,

perforated; some plates with one spinous edge or

spinous tapered end or marginal pointed projec¬

tions; plates with surface spines; lacking tables,

cups and crosses; lacking towered crosses at any

developmental stage.

Species. H. denticiilata (Ekman, 1927); H. godef-

yi-qi’; (Semper 1868); H. steineni (Ludwig, 1898).

Distribution. Antarctica, Magellanic region,

Kerguelen region.

Remarks. Ileding (1942) nominated Cucumaria

tiircjueti Vaney, 1906b as type species for his new

genus Ekmocucumis. Panning (1949) created two

subgenera of Ekmocucumis: Ekmocucumis (type

species Cucumaria turqueti), and Heterocucumis

(type species Cucumaria steineni). Based on very

close similarity of developmental changes in os¬

sicle fomi. Hansen (1988) supported the original

thinking of Ekman (1927) and assigned

Cucumaria liouvillei Vaney, 1914, C. ahyssorum

Theel, 1886a and C. turqueti Vaney, 1906b to

Staurocucumis Ekman, 1927 (type species C.

liouvillei). Massin (1994) followed Hansen

(1988). Titus the type species (Cucumaria tur¬

queti) of the genus and subgenus Ekmocucumis

Heding, 1942 has been assigned to Stauro¬

cucumis, the genus name Ekmocucumis becoming

a junior synonym. The subgenus Heterocucumis

Panning, 1949 is raised here to generic status.

Cucumaria grandis Vaney, 1906b was consid¬

ered by Ekman (1925. 1927) to be a junior syn¬

onym of Cucumaria turqueti [C grandis has page

precedence over C. turqueti in Vaney 1906b, but

the relegation of C. grandis to junior synonymy

by Ekman (1925) stands under the “Principle of

the First Reviser" (ICZN Article 24a)]. Cherbon-

nier( 1974) retained Ekmocucumis grandis (junior

synonym Cucumaria spatha Cherbonnier,

1941b). With a probable synonymy [adequate

comparative material not yet examined] of

E. grandis with S. turqueti, the fonner is

provisionally refeiTcd here to Staurocucumis.

Of the three species (coatsi. denticulata,

steineni) originally assigned by Panning (1949) to

his new subgenus Heterocucumis, denticulata and

steineni are retained here in Heterocucumis, and

coatsi is referred belov\ to a new genus Psolicrux.

C. godeffroyi is referred here to Heterocucumis

on the basis of; ten subeqiial tentacles; radial dou¬

ble rows of tube feet; elongate perforated plates

with spinous surface and pointed marginal projec¬

tions, some plates with one end narrow and

spinous in smaller specimens; signilicant ossicle

change during development (Ludwig, 1898b;

Pawson, 1969); and tendency to lose the calcare¬

ous ring with age. Although body wall ossicles in

large specimens of H. godeffroyi arc significantly

smaller than in large specimens of //. steineni,

they are indistinguishable from the small ossicles

present in //. steineni. Both species have a leath¬

ery non-calcarcous body wall, tend to lose the cal¬

careous ring with size, have a maximum size of

about 70 mm, and have dark brown colouration

anteriorly.

Heterocucumis godeffroyi (Semper) comb. nov.

Cucumaria godeffroyi Semper. 1868: 53, pf 15 tigs

12, 14.—Lampert. 1885: 144.— Tliccl, 1886a:

99-100.—Ludwig, 1898b: 435-437, pi. 26 (igs

15-21.—H.L. Clark. 1910: 352-353.—Ekman, 1925:

49.—Deichmann. 1941: 83-84.—Deichmann, 1947:

334.—Panning. 1955: 43-45, figs 7-9.—Pawsoa.

1969: 131-133, fig. 1.

Stereoderma godeffroyi. —Panning. 1949: 422.^

Pawson, 1964: 457.

Material e.xamined. Kersjuelen region, Kerguelen I..

Royal Sound. 49'28’S, 70-04’E. 4-5 m, 1929 [BAN-

ZARE Stn 12. SAM KI845 (2 specimens)]; 49”28'S.

70'12 E, 47 m, 1930 [BANZARE Stn 59. SAM KI844

(2)]; Heard 1., 5n6'-53M2'S, 73’05'-76”02T,.

200-379 m [ANARE, NMV F84996 (1).

F84999-85001{3)].

Type locality. Chile. Iquique.

Distribution. Magellanic region, Chile, 0-60 m

(summary by Pawson, 1969). Kerguelen region,

Kerguelen and Heard Is, 4-379 m (this paper).

Remarks. Panning (1949) assigned C godeffroyi

to .Stereoderma Ayres, but subsequently (1955)

provided a description of Cucumaria godeffroyi.

implicitly returning the species to Cucumaria.

Panning (1964) fonnally regarded Stereoderma

as monotypic. excluding C. godeffroyi.

The sparse flat perforated plates with spinous

surface and pointed marginal projections are com¬

monly 0.06 and up to 0.10 mm long in larger

Heard and Kerguelen specimens, and up to 0.14

mm long in smaller specimens. Pawson (1969)

reported plates in larger specimens from olTChile

as larger (average 0.17 mm long). The change in

ossicle form with size reported and illustrated by

Pawson (1969) is evident in the Heard and Ker¬

guelen material, and the dark brown colour

around the base of the tentacles is present in some

Heard specimens.

This species is not Cucumaria godfroyi Vaney,

1914, which Ekman (1925, 1927), Panning

(1949) and Massin (1994) regarded as a junior

BANZARE AND ANARE HOLOTHUROIDEA

317

synonym of Ekmociicumis steineni (Ludwig),

now Heterocucumis steineni (below). This paper

reports an extension of the distribution of H. god-

effroyi from the Magellanic to Kerguelen region,

at a generally greater depth. It has not been

reported from the Antarctic coast.

Heterocucumis steineni (Ludwig)

Cucumaria steineni Ludwig, 1898a: 30-32. pi. 2 figs

22-24.—Ekman, 1925; 38^5, fig. 6.—Ekman, 1927;

363,390-393, figs 12, 13.—Panning, 1936; 15-16, figs

6 , 7.—Cherbonnier. 1941a: 464-468.

Cucumaria antai'clica Vaney, 1906b: 6—10, figs 3, 8,

26.—Vaney, 1908: 427.—Vaney, 1914: 8-9.

Cucumaria godfroyi Vaney, 1914: 11-12. pi. 4 figs

1-5.

Ekmocucumis (Heterocucumis) steineni. —Panning,

1949; 452-453. fig. 51.

Heterocucumis antarctica. —Cherbonnier, 1974:

609.

Heterocucumis godfroyi.— C^iahonmer, 1974: 609.

Ekmocucumis steineni. —Gutt, 1991b: 324.—

O’Loughlin et al.. 1994: 549, 554.

Heterocucumis steineni. —Massin, 1994; 130-145,

figs 5-16.

Ekmocucumis sp. MoV 2005.—O'Loughlin et al.,

1994: 551-552, 554.

Material examined. Antarctica, off Enderby Land,

65‘48'S, 53*16'E. 193-209 m, 1930 [BANZARE Stn

41, SAM K1832 (1)]; Prydz Bay, off Enderby and Mac-

Robertson Lands, 66*53'—68"3I S, 65"26'-78"13'E,

105-743 m [ANARE. NMV F68024 (1), F68030 (2).

F68053 (1). F68100 (3), F68678 (1), F69109 (1),

F69111 (1), F72615 (4), F91297 (27)].

Type localit}’. South Georgia.

Distribution. Western Antarctica, South Orkney

Is, South Georgia, Falkland Is and Burdwood

Bank, 2-400 m (summary by F.kman, 1927); off

Graham Land. 0-110 m (Vaney, 1906b, as C.

antarctica); off Alexander L, 6-297 m (Vaney,

1914, as C. antarctica and C godfroyi); Weddell

Sea, 160-1180 m (Gutt, 1991b). Eastern Antarc¬

tica, off Terre Adelie, 6-210 m (Cherbonnier,

1974, as H. antarctica and H. godfroyi); off Wil¬

helm 11 Land, 350-400 m (Ekman, 1927); off

Enderby and Mac Robertson Lands, 105-743 m

(this paper).

Remarks. Ekman (1927) considered C. antarctica

and C. godfroyi to be junior synonyms of C.

steineni. Cherbonnier (1941a, 1974) rejected the

synonymies by Ekman (1927). and reported Het¬

erocucumis antarctica (Vaney) and Heterocu¬

cumis godfroyi (Vaney) for Terre Adelie. Massin

(1994) supported the synonymies after a study of

developmental changes in ossicle form. Determi¬

nation of H. steineni in this study is based on the

illustrations of variation in ossicle form with size

by Massin (1994). Material determined as

Ekmocucumis sp. MoV 2005 by O’Loughlin et al.

(1994) comprises mostly larger specimens of H.

steineni.

Psolicrux gen. nov.

Type species. Psolidium coatsi Vaney, 1908.

Diagnosis. Body wall thin, calcareous; mouth

anterior on short taper; anus posterior on elongate

tapered tail; 10 subequal dendritic tentacles; cal¬

careous ring lacking posterior prolongations;

mid-body ventral radial tube feet larger than those

extending to introvert and anus, creating sole-like

ventrum; scattering of dorsal and lateral tube feet

beyond dorsolateral radial series; body wall ossi¬

cles plates and knobbed crosses; plates irregularly

oval, typically 0.44 mm long in western Antarc¬

tica (0.60 mm in eastern Antarctica), perforated,

smooth surface and margin, one end tapered into

narrow, sometimes perforated, distally spinous

spire angled above plate surface; crosses small,

typically 0.08 mm long in western Antarctica

(0.06 mm in eastern Antarctica), not cupped, with

irregular short frequently distally knobbed

branches; lacking tables and cups.

Etymology’. From a family name Psolidae, refer¬

ring to the sole-likc ventrum, and crux (Latin for

cross), referring to the knobbed crosses.

Remarks. Psolicna is distinguished from other

genera of Cucumariidae by a combination of

elongate thin tail, sole-like ventrum, scattered

dorsal and dorsolateral tube feet, smooth perfo¬

rated plates with angled spinous spires, and irreg¬

ular knobbed crosses.

PsoHcrux coatsi (Vaney) comb. nov.

Figures 3c-f

Psolidium (Cucumaria) coatsi Vaney, 1908:

424-425, pi. 4 figs 47-50.

Cucumaria conspicua Vaney, 1908: 433, pi. 2 figs

15-16, pi. 5 fig. 67.—Ekman, 1927: 414.—Panning,

1949: 416 [new synonymy],

Psolidium navicuh Ekman. 1927: 408-414. fig.

18.—Panning. 1949: 455.—Gutt, 1991b: 324 [new syn¬

onymy].

Psolidium histriatum Ludwig and Heding, 1935:

165-167, figs 30-31.—Panning, 1949: 455 [new syn¬

onymy].

Ekmocucumis (Heterocucumis) coatsi. —Panning,

1949:452.

Heterocucumis coatsi. —Cherbonnier, 1974: 609.

Ekmocucumis sp. MoV 2013.—O’Loughlin et al.,

1994: 549.

318

P. M. O’LOUGHLIN

Material examined. Western Antarctiea. Weddell Sea.

115 m, 1958 [USNM E5I318 (7 specimens)]: South

Sandwich Is, 415-613 m, 1975 [USNM E51314 (1)];

South Shetland Is, 97-113 m. 1966 [USNM E5I317

(2)]; Palmer Archipelago, 55 m, 1972 [USNM E51315

(1)]. Eastern Antarctica, off Wilkes Land, 183-237 m.

1961 [USNM E51316 (3)]: olT Enderby and Mac-

Robertson Lands, Prydz Bay. 65"56'-67"4rS.

50°52 -77‘32'E, 98-400 m [ANARE. NMV F68044

(1), F68047 (2), F68084 (I). F68668 (1). F726I4 (1),

F84985(l)].

Type locality. Western Antarctica. Scotia Sea.

Distribution. Western Antarctica, Weddell Sea,

125-990 m (Gutt, 1991b, as P. navicula); Scotia

Sea. 165-183 m (Vaney, 1908); off Coats Land.

South Shetland Is. South Sandwich Is, Palmer

Archipelago. 55-613 m (this paper). Southern

Ocean, east of Bouvet 1., 567 m (Ludwig and

Heding, 1935, as P. histriatum). Eastern Antarc¬

tica, off Terre Adelic, 10-170 m (Cherbonnicr,

1974); olTWilhelm II Land, 350-385 m (Ekman,

1927, as P. navicula): Prydz Bay, 98-400 m (this

paper).

Remarks. Based on the original descriptions, and

examination of P. coalsi specimens. Psolidium

navicula Ekman, 1927 and Psolidium histriatum

Ludwig and Heding, 1935 are considered here to

be junior synonyms of Psolicncx coalsi (Vaney.

1908). A distinctive diagnostic character is the

consistent occurrence of the small, irregular,

bluntly spinous crosses illustrated by Vaney

(1908) for P. coatsi. by Ekman (1927) for

P. navicula, by Ludwig and Heding (1935) for

P. bistriatum, and referred to as “cups" by Pan¬

ning (1949) [they are not cupped]. The three

authors illustrated short spinous apophyses rising

from the end of smooth perforated oval to elon¬

gate plates, and obsen'ed that the mid-body ven¬

tral radial tube feet were larger than those extend¬

ing to the introvert and anus and referred their

species to Psolidium. All recognised the posterior

tapered conical body form. Ekman (1927) and

Ludwig and Heding (1935) illustrated the distinc¬

tive constricted base of the anterior projections of

the radial plates of the calcareous ring. Ludwig

and Heding (1935) considered that the plates with

a narrow perforated spinous end and the knobbed

crosses were similar to the ossicles of Cuciimaria

insolens Theel, 1886 from South Africa, but in the

illustrations by Theel (1886a) the plates in

C. in.sotens are knobbed, not smooth, and the

spinous end is not angled above the plate. The

knobbed crosses in C. insolens are cupped.

Clicit maria conspicua Vaney, 1908 was

described from a single small specimen (10 mm

long) taken at the same station as P. coalsi. It haq

a thin flat oval ventrurn with large radial tube feel,

small tube feet scattered beyond the dorsolateral

radial series, terminal mouth and anus, short poSv

terior taper, and single-layered perforated plat^>

ossicles with spinous taper. This specimen ij;

considered here to be a juvenile P. coatsi.

Ekman (1925) described material asi

Cucumaria coalsi which had anterior interradia|

brood pouches, ten tentacles with the two ventral

ones smaller, and ossicles not consistent with

those of P. coalsi, which does not have broocl

pouches, has ten subequal tentacles, and has small

spinous crosses and elongate perforated plaltj

ossicles with raised narrow short perforatetl

spinous ends. The material described by Ekmaq

(1925) was not P. coalsi. Ossicles of P. coatsi

illustrated by Massin (1994) are similar to those

of Ekman (1925). and do not resemble those IIIuSn

I rated by Vaney (1908) for the type of P. coatsi.

The distribution of P. coalsi is extended here to

eastern Antarctica.

Staurocucuntis UouvUlei (Vaney)

Cucumaria UouvUlei Vaney. 1914: 12-14, pi. 3 figs

1-3.—Ekman. 1925: 85-89. fig. 18.

Staurocucumis UouvUlei. —Ekman, 1927: 363

374-381, figs 7. 8.—Heding. 1942: 33.—Panning]

1949: 455.—Cherbonnicr, 1974: 609.—Hansen, 1988;

301-302, 304, 307, lig. 3.—Massin, 1994: 129-132]

fig. 1.—O'Loughlin et al., 1994: 552. 5.54.

Cucumaria mira Ludwig and Heding, 1935-

172 179. figs 39^1.

Tracliylliyoiie mira. —Panning, 1949: 426.-^

O'Loughlin et al.. 1994: 552. 554 [new synonymy].

.Ahy.v.socucumis UouvUlei .—Gull, 1991b: 324.

Material e.xamined. Eastern Antarctica, off Kemp Land,

66-45'S, 62'03 E. 177 m. 1931 [BANZARE Stn 107,

SAM K1S48 (4 specimens)]: Piy'dz Bay, 66“53''

68"58 S. 68'56 -77'33'E, 131-791 m [ANARE. NMV

F68018 (1), F68022 (1). F68025-68027 (12), F68029

(5). F68031-68034 (27). F680.36 (I). F68038 (1),

F68040 (2). F68048 (1). F68055 (1), F68059 (I),

F68061-68062 (2). F68064 (1). F68077-68078 (2).

F68080-68081 (4). F68086 (1). F68089 (1). F68096

(2), F68099 (1), F68656 (1), F69101 (2). F691()3 (4).

F69129 (1). F69133 (2). F71991 (1)]. Heard 1.,

51' I7'-53'I3 S. 71 45'-76'32 E, 60 .541 m [ANARE.

NMV F85015-85034 (105)].

Type locality. Western Antarctica, Belling¬

shausen Sea side of Antarctic Peninsula.

Disirihiiiion. Western Antarctica, west of Antarc¬

tic Peninsula, 98-297 m (Vaney, 1914); South

Georgia, 125-310 m (Ekman, 1925); Weddell

Sea, 160-745 m (Gutt, 1991b). Southern

Ocean, east of Bouvet I., 439-567 m (Ludwig

BANZARE AND ANARE HOLOTHUROIDEA

319

and Hcding, 1935, as C. mira). Eastern

Antarctica, off Terre Adelie, 385 m (Cherbon-

nier, 1974); off Wilhelm 11 Land, 350-485 m

(Ekman, 1927); Prydz Bay, 131-791 m (this

paper). Kerguelen region. Heard 1., 60 541 m

(this paper).

Remarks. The Antarctic and Heard 1. specimens

were indistinguishable in terms of: size (up to 80

mm long); thin body wall with outer gelatinous

brown layer, darkens with age; ten equal long

very branched tentacles, always extended, fre¬

quently lost; tube feet on radii only, more devel¬

oped ventrally; calcareous ring normally not

evident; ossicles frequently not evident in larger

specimens; oval bowl ossicles with denticulate

perforations. Vaney (1914) illustrated a bowl 0.22

mm long for type material [size of specimen

sampled not given). For Bouvet material Ludwig

and Heding (1935) illustrated a bowl 0.11 mm

long for a 7.5 mm specimen. For western Antarc¬

tic material Massin (1994) illustrated bowls up to

0.13 mm long in a 10 mm long specimen, and

0.22 mm long in a 51 mm specimen. Eastern

Antarctic material in this study had bowl sizes up

to 0.29 mm long in a 65 mm long contracted spec¬

imen (NMV F68031). Bowl size increases signif¬

icantly with increasing size of specimens. Eastern

Antarctic and Heard bowl sizes differed signifi¬

cantly for specimens of similar size, the largest

bowls up to 0.29 mm long in a 65 mm long con¬

tracted specimen (Antarctic. NMV F6803I) and

0.17 mm long for a 70 mm long contracted spec¬

imen (Heard, NMV F85033). Based on the

description and illustrations of C. mira Ludwig

and Heding, 1935. it is considered here to be con-

specific with S. liouvillei. The distribution of S.

liouvillei is extended here to the Kerguelen

region.

Trachythyone lechleri (Lampert)

Thvone (ThvomJium) lechleri Lampert, 1885: 253,

pi. I ’fig. 64.—Thcel. 1886a: 267.

Thvone te/i/er/.-Ludwig. 1898a: 44-49, pi. 2 figs

26-30, pi. 3 figs 31-33.—Perrier, 1905: 35-38.—

Ekman, 1925: 101 103, fig. 22.—Deichmann, 1947;

335-336.

Thvone ha.v.sleri T\\6c\. 1886b; 11-12.

Trachvthvone /ec7i/c'W.—Panning, 1949: 426, figs

12-14.-Tpanning, 1964: 166-167. figs 6, 7.—Pawson.

1964: 459-461. fig. 2(1-5).—Hernandez, 1982;

256-257; figs lb, 3b. 4; pi. 2j-m.

Material examined. Kerguelen 1., Royal Sound,

49"28'S, 70"12'E, 47 m. 1930 [BANZARE Stn 59.

SAM K1849 (1 specimen)]; Heard I., 52‘32’-53'30'S,

73”10'-73'4rE, 60-238 m [ANARE, NMV

F84992-84995 (6)].

Type locality. Straits of Magellan.

Distrihution. Magellanic region. Straits of Magel¬

lan, Tierra del Fuego, up to 30 m (summary by

Pawson, 1964). Kerguelen region, Kerguelen and

Heard Is, 47 -238 m (Pawson, 1964; this paper).

Remarks. The Heard I. material does not differ in

any recognisable way from the descriptions and

illustrations of Magellanic region material by

Ludwig (1898a), Ekman (1925) and Pawson

(1964). The largest specimen is 145 mm long

(Pawson gave 150 mm), and the multiple series of

radial tube feet create a superficial appearance of

covering the whole body. The distinctive irregu¬

larly oval plates, with sparse to absent small per¬

forations. are typically 0.16 and up to 0.28 mm

long (Ekman gave 0.12 mm. Pawson 0.2 mm).

The large perforated plates, present in small spec¬

imens only, are up to 0.38 mm long (Ludwig gave

0.43 mm, Ekman 0.21 mm). The thick cupped

spinous crosses are 0.05-0.06 mm long (Pawson

gave 0.05 mm).

The type material for T. hassleri was taken in

the Straits of Magellan, and Theel (1886b)

acknow ledged the strong resemblance to T. lech¬

leri. Ludwig (1898a) synonymised the two

species, and Pawson (1964) commented in detail

on the synonymy and agreed. Pawson (1964)

identified a single specimen from I leard 1. in the

Dominion Museum (Wellington, New Zealand)

as T. lechleri, and additional material is present in

the BANZARE collection from Kerguelen and

ANARE collections from Heard 1. This species

has not been reported for Antarctica, but occurs in

the Magellanic region and at greater depth in the

Kerguelen region.

Paracucumidae Pawson and Fell

Paracucumis turrieata (Vaney) comb. nov.

Thvone turrieata Vaney, 1906a: 401^02, fig. 3.—

Ekman. 1925: 108-1II. fig. 25.

Paracucumis antarctica Morlensen. 1925: 9-12, figs

6-8.—Panning. 1936: 11-14. figs 1-5.—Heding and

Panning, 1954: 46-47, fig. 7.—Pawson. 1982; 815.—

O’Loughlin el al., 1994: 552, 554 [new synonymy],

Paracucumis turrieata. —Panning. 1936: 14.

Ypsilocucurnis turrieata. —Panning, 1949: 455.—

Cherbonnier. 1974: 610.—Cult, 1991b; 324.

Material examined. Thvone turrieata. Vaney, 1906.

Hololype (MNHN). Western Antarctica, South Orkney

Is, Scotia Bay, littoral. 1905.

Other material, liaslem Antarctica, off Wilkes Land,

65”10 S. 108'12'E, 474 m. 1931 [BANZARE Stn 97.

SAM K1846 (I specimen)]; Prydz Bay and Mae-

Robertson Shelf, 66°54'-68”57 S, 63°06'-76°38'E,

320

P. M. O’LOUGHLIN

367-1240 m [ANARE, NMV F68101 (1), F6815I (5).

F6867I-68674 (5), F69122-69123 (48), F69I28 (2),

F69I32 (29). F71983-71984 (9), F7I986 (1). F8I430

(1), F91298 (3)].

Distribution. Western Antarctica, Weddell Sea,

620-705 m (Gutt, 1991b); off Graham Land, 125

m (Ekman, 1925); South Orkney Is (Vaney,

1906a). Ea.stem Antarctica, Ross Sea, Discovery

Inlet, 550-560 m (Mortensen, 1925, as P. antarc-

tica)\ oFT Terre Adelie, 10-15 ra (Cherbonnier,

1974, as Y. turricata); off Wilkes Land. Prydz

Bay, MacRobertson Shelf, 367-1240 m (this

paper).

Remarks. The illustrations of unique large plates

with central digitate towers for P. antarctica by

Mortensen (1925) and Heding and Panning

(1954), and for T. turricata by Vaney (1906a) and

Ekman (1925), show the same ossicle fonn, and

illustrate the plates seen in this study of the

type and BANZARE and ANARE material.

Paracucumis antarctica Mortensen. 1925 is

considered here to be a junior synonym of

Thyone turricata Vaney, 1906a. This synonymy

removes the anomaly of the previously reported

mutually exclusive occurrences of T. turricata in

Western Antarctica and off Terre Adelie. and of

P. antarctica in the Ross Sea and Prydz Bay.

Ossicle form does not vary significantly with

specimen size, but tentacle form and number (up

to 15) are variable. The description by Vaney

(1906a) of eight and two small tentacles on the

small (12 mm long) type specimen of T. turricata

does not apply to large specimens. ANARE spec¬

imens were up to 220 mm long (excluding tenta¬

cles). Tw'o specimens (more than 200 mm long.

NMV F9I298) had 14 tentacles in combinations

of ten large subequal with four adjacent small

subequal, and 12 large with two adjacent .small.

Calcareous ring form is also variable, as noted

and illustrated for P. antarctica by Mortensen

(1925) and Heding and Panning (1954). One

specimen (200 mm long, NMV F68101) had five

radial plates deeply bifurcate anteriorly and five

interradials with single narrow anterior projec¬

tions, creating 15 anterior projections. Mortensen

(1925) noted for P. antarctica that ossicles were

spaced in a thin transparent body wall in relaxed

specimens, but imbricating in contracted speci¬

mens.

Panning (1936) recognised the close similarity

of T. turricata to P. antarctica, and reassigned

T. turricata to Paracucumis. Panning (1949) sub¬

sequently reassigned T. turricata to his new genus

Ypsilocucumis. T. turricata has dendritic tentacles

and belongs in the Dendrochirotida, not the

Dactylochirotida with digitiform tentacles (Paw-

son and Fell, 1965), and cannot be retained in

Ypsilocucumis Panning, 1949. It is reassigned

here to Paracucumis Mortensen, 1925.

Order Elasipodida ThecI

Elpidiidac Theel

Amperima rohusta (Theel)

Scotoplanes rohusta Theel, 1882: 35-36, pis 6,

34(6-7), 37(9).

Amperima robustum. —Agalep. 1967a: 56-51.

Amperima rohusta. —Hansen. 1975: 161-162, figs

n-1^.

Material e.xamined North-east of Kerguelen, 47'05’S,

79°16'E, 3112 m. 1930 [BANZARE Stn 66, SAM

K1842 (2 specimens)].

Type locality Southern Ocean, 53°55’S,

108'35'E,3568 m.

Distribution. Antarctica, 2010-4240 m; Kermadec

Trench, 2640 m (summary by Hansen, 1975);

southern Indian Ocean, 3112m (this paper).

Remarks. The two specimens were small (up to

14 mm long) and very damaged, but had the char¬

acteristics of the species as described by Hansen

(1975). There were ten tentacles; the velum con¬

sisted of the two pairs of papillae with a pair of

very small lateral ones; the small specimens had

only nine pairs of tube feet, but they bordered the

posterior two-thirds of the sole with the anterior

pairs larger and more spaced and the posterior

pairs decreasing in size; the ossicles were tripar¬

tite with distal spines and spinous apophyses on

each arm. The ossicles were eroded, but two vari¬

ations from the descriptions by Hansen (1975)

were evident. Spinous apophyses were present on

ventral tripartite ossicles which were up to 0.26

mm wide, and no C-shaped ossicles were found.

These damaged specimens were similar to two

species described by Gebruk (1988). On the lim¬

ited evidence available, they differed from

Amperima helyaevi Gebruk. 1988 [Aleutian Is,

5030 m] by having slightly larger ventral ossicles,

larger os.sicle spinelets, and less evenly spaced

tube feet. They differed from A. vitjazi Gebruk,

1988 [off Chile. 4300 m; Japan Sea, 6096 m] by

being significantly smaller, having unbranched

ossicles, and unevenly spaced tube feet.

Rhipidothuria racovitzai Herouard

RhipiJothuria racovitzai Herouard, 1901: 41-42.—

Herouard . 1906: 7-8, pi. 1 figsl-S.—Hansen, 1975:

131.—Gebruk, 1990: 81, fig. 28 (3^).—Gebruk and

Shirshov, 1994: 148-149, fig. 1.

BANZARE AND ANARE HOLOTHUROIDEA

321

Achlyonice vinlaectispklata Gutt, 1990: 123-125,

Ergs 7-10.—Gutt, 1991b: 324.

Material examined Achlyonice violaecuspidata. Gutt,

1990. Paratypes. Western Antarctica. Weddell Sea.

77'08'S, 48‘36'W. 235 m, 1984 fZIM E-7144 (2 spec¬

imens)].

Other material. Eastern Antaretica, olT Enderby

Land. 66‘10'S. 49'4I 'E. 300 m, 1930 [BANZARE Stn

39, SAM K.1829 (I)]; 66‘12'S, 49-37'E, 300 m. 1930

[BANZARE Stn 40, SAM KI830 (24)]; olT Wilkes

Land, 65”ID'S, I08‘12'E.474 m. 1931 [BANZARE Stn

97, SAM KI831 (3)].

Type localit}'. Western Antarctica, Belling¬

shausen Sea.

Distribution. Western Antarctica, Weddell Sea,

225-785 m (Gutt, 1991b, as Achlyonice violae-

cutipidata); Bellingshausen Sea, 800 in

(Herouard. 1901; depth estimated from latitude

and longitude). Eastern Antarctica, off Enderby

and Wilkes Lands, 300-474 m (this paper).

Remarks. The 28 BANZARE specimens (Itted the

diagnosis by Gebruk and Shirshov (1994), except

that no crosses were found. Specimens were up to

60 mm long. There were ten tentacles, the antcr-

ionnost largest; seven pairs of dorsal papillae,

distributed evenly along the dorsum; 12 pairs of

tube feet, the posterior ones smallest and webbed;

and a velum comprising two pairs of transverse

papillae. Tentacles, mouth, papillae, tube feet and

anus were a residual violet brown colour. Rod

ossicles were found in the tentacles, tube feet and

body wall. Rods were bent or straight or curved,

unbranched, not perforated, sometimes thickened

centrally, and sometimes with small knobs. Rod

lengths were up to 0.46 mm (tentacles) and 0.37

mm (body wall). Herouard (1901) noted that ossi¬

cles were probably in the form of a cross sur¬

mounted by a single point. Crosses were not

found by Gutt (1990), Gebruk and Shirshov

(1994), and this author. “Crosses" were observed

in the BANZARE material, but were sponge

spicules. This species is not represented in the

extensive ANARE collections from eastern

Antarctica, but the BANZARE specimens do

extend the distribution of this species to eastern

Antarctica.

Lactniogoiiidac Ekman

Laetmoi’one wyvillelliomsoni Theel

Laetmogonc wwilletliomsoni Theel, 1879: 10, pi. 1

figs 12-13.—Tiiecl, 1882: 73-78, pis 11, 12,

31(14-16). 34(1), 36(3), 37(5, 7, 11), 38(9), 39(4),

42(1, 7), 43(4), 44(14), 46(2-3).—Agatep, 1967b: 63,

pis 8(1-11),9(1-10); tbl. 1.—Hansen, 1975:54-57,%

20, tbls 5-8.—Gutt, 1991b: 324.—Gebruk, 1993: 240,

fig. 6(1).

Laetmogone cf. m’villethomsoni .—O’Loughlin et al.,

1994: 553-554.

Material examined. Syntype. Pacific Ocean. 33°42'S.

78'18'W, 2514 m. Challenger Stn 300 [BMNH

83.6.1.8.45 (1 specimen)].

Other material. Eastern Antarctica, off MacRoberl-

.son Land. 66*48 S. 71‘24'E, 540 m. 1929 [BANZARE

Stn 30, SAM KI827 (I)]; olf Kemp Land, 66‘2rS,

58‘50 E, 603 m, 1930 [BANZARE SUi 34, SAM

K.1828 (1)]; Prydz Bay. 67‘10'-68-32'S,

7ri8'-73'14'E, 515-743 m [ANARE, NMV F68168

(1), F76585 (1), F76605 (1), F81826-81828 (22)].

Type locality. Southern Pacific Ocean, ofT Chile.

Distribution. Southern Ocean. Kermadec Trench.

2514^410 m (Hansen. 1975); Scotia Sea, South

Shetland Is, 2672—4136 m (Agatep, 1967b).

Western Antarctica, Weddell ,Sea, 245-1130 m

(Gutt, 1991b). Eastern Antarctica, off Kemp

Land, Prydz Bay, 515-743 m (this paper).

Remarks. One well-preserved specimen (NMV

F81825) was 55 mm long and had 14 tentacles, 20

pairs of tube feet. 14 pairs of thin papillae some as

long as the body, large tenninally spinous ventral

rods up to 0.4 mm long, an overall body range of

wheel sizes from 0.04 to 0.21 mm diameter with

8-16 outer spokes and 4-6 inner rays, and

showed no significant variations from the type

examined and the obsetx ations by Hansen (1975).

Hansen (1975) gave wheel diameters up to 0.17

mm, Theel (1879) up to 0.16 mm. Two eastern

Antarctic specimens (NMV F8I826), 80 and 140

mm long, had wheel diameters up to 0.19 and

0.18 mm respectively. Wheel size does not vary

with increase in specimen size. In the Southern

Ocean and the Kennadec Trench this species is

abyssal, but around the Antarctic continent it

occurs at bathyal depths.

New distributions summary

Four holothurian species arc conflmied here as

occurring in both the Magellanic and Kerguelen

regions: Chiridota pisanii Ludwig, 1886, Taenio-

gyrus contortus (Ludwig, 1875), Heterocucumis

godeffroyi (Semper, 1868) and Trachythyone

lechleri (Lamport, 1885). Only T. contortus

occurs also on the Antarctic coast. Staurocucumis

liouvillei (Vaney. 1914) occurs in Antarctica and

off Heard L, but not in the Magellanic region.

All of the holothurians reported here with an

Antarctic distribution occur in both western and

eastern Antarctica, with Meseres spiculifenis sp.

nov. an unconfinned exception. Significant new

distribution data are tabulated in Table 4.

322

P. M. O’LOUGHLIN

Acknowledgments

I am grateful for the assistance with materials and

resources given by Cynthia Aheam (USNM); Dr

Nadia Ameziane (MNHN); Dr Penny Berents

(AM); Joke Sleeker (ZMA); Dr Margit Jensen

and Dr Claus Nielsen (ZMUC); Dr Dietmar

Keyser (ZIM); Dr Claude Massin (IRSNB); Ann

Morgan (BMNH); Tania Bardsley, Dr Tim

O’Hara and Chris Rowley (NMV); Dr Alexei

Smirnov; and Wolfgang Zeidler (SAM). I am

grateful to llelga Griffin, Dr Igor Smirnov and

Luba Sosnin for translations; to Mark Darragh for

assistance with photography; and to Ben Boonen

for the presentation of the images. I am particu¬

larly grateful to Cynthia Aheam, Dr Claude

Massin, Dr Tim O’Hara, Dr David Pawson and

Dr Gary Poore for their most helpful reviews of

the manuscript.

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the Royal Society of Edinburgh 46:405—441,5 pis.

Vaney, C., 1914. Hololhuries. Deuxieme Expedition

Antarctique Fran 9 aise (1908-10). Sciences

Naturelles: Documents Scientifiques. 54 pp., 5 pis.

Memoirs of the Museum of Victoria 59(2): 327-331 (2002)

A NEW SPECIES OF NEANTHES (POLYCHAETA: NEREIDIDAE)

FROM SOUTHERN AUSTRALIA

Torkild Bakken

Department of Natural History, Norwegian University of Science and Technology, N-7491 Trondheim, Norway

(e-mail: torkild.bakken@vm.ntnu.no)

Abstract

Bakken, T., 2002. A new species of Nemuhes (Polychaeta: Nercididac) from southern

Australia. Memoirs of Museum Victoria 59(2): 327-331.

A new species of Neaulhes from southern Australia is described. Examination of several

species of Neatuhes has resulted in a re-interpretation and stronger emphasis of parapodial

characters than is usual in descriptions of nereidid species. These characters are used to

describe Neanihes tasnumi sp. nov. and to distinguishing the new species from the closely

related Neatuhes bassi Wilson. 1984.

Introduction

In an earlier study of Neatuhes from Victoria,

Wilson (1984) found four specimens closely

allied to Neatuhes bassi Wilson, 1984 that po.s-

sessed characlers indicating they belonged to a

different species. Re-examination of these four

specimens and additional material, as well as

more material of N. bassi, confirms that this

material belongs to a new species.

This study has also resulted in a re-evaluation

of parapodial characters used to describe nerei-

dids. The use of detailed descriptions of mor¬

phology of parapodia in nereidids was presented

by Hylleberg ct al. (1986) and Hylleberg and

Nateewathana (1988). The importance of these

characters is evident in several apparently closely

related species of Neatuhes from southern Aus¬

tralian waters. Parapodial characters tend to vary

with specimen size. Wilson (1984) showed that

parapodial ligules and lobes tend to occur over a

smaller range of chaetigers in specimens with

body width less than 1.5 mm than in larger spec¬

imens. Parapodial ligules and lobes may be miss¬

ing altogether in specimens with body width less

than 1 mm, especially pre- and postchaetal lobes.

In the present study a di.stinction is made

between a prechaetal or postchaetal lobe and an

acicular process. These features are used in the

present description to distinguish N. tasmani sp.

nov. from closely related taxa. My own examina¬

tion of several nereidine taxa has proved these

features to be important characters over a wide

range of taxa within the subfamily. In the

notopodium a prechaetal lobe is clearly present as

a lobe (Fig. IB) but might be of any size from a

small lobe barely evident beyond the dorsal collar

to a lobe as long as the dorsal and ventral ligules.

An acicular process on the other hand is fused to

the ventral notopodial ligule as a ridge on the

ligule itself (Fig. ID). In the ncuropodium a

clearly identifiable postchaetal lobe is found in

many species. This lobe might be digitiform or

llattened, a distinction that should be explicit in

descriptions. The postchaetal lobe may be

reduced in posterior chaetigers, present through¬

out the body or be absent. I f the lobe is absent an

acicular process might be present instead, most

often seen as an oval process protruding beyond

the lip of the aciculum. Pre- and postchaetal lobes

and acicular processes can be seen in figures in

earlier studies (e.g. Hutchings and Turvey, 1982;

Wilson, 1984), but it is important that they are

clearly outlined in species descriptions so that

otherwise similar species can be distinguished

using these characters. To give parapodial char¬

acters a stronger emphasise an end-view of

parapodia arc drawn for N. tasmani following

the example of Hylleberg and Nateewathana

(1986).

The material presented in this work is

deposited in Museum Victoria. Melbourne

(NMV), Australian Museum, Sydney (AM) and

the South Australian Museum, Adelaide (SAM).

Measurements of body width are measured with¬

out parapodia at about chaeliger 10. This is con¬

sistent with earlier studies (e.g. Wilson, 1984)

and will be used in future studies as a standard

measurement.

327

328

T. BAKKEN

Neanthes Kinberg, 1866

Type species. Neanthes vaalii Kinberg, 1866.

Diagnosis. Eversible pharynx with conical parag-

naths on both rings, bar-shaped paragnaths in

Area IV may be present. Four pairs of tentacular

cirri. Parapodia biramous. Notochaetae

homogomph spinigers; neurochaetae including

homogomph and heterogomph spinigers (after

Wilson, 1988).

Neanthes bassi Wilson

Neanthes bassi Wilson, 1984: 210-212, fig. 1.

Material examined. Holotype, NMV F50005.

Paratypes. NMV F50006—FSOOl I (fully location data

given in Wilson, 1984).

Additional material. Victoria. NMV

F50()I2-F50017; NMV F50I23 (2 specimens); NMV

F4I866 (referred to as GI866 with full data in Wilson,

1984).

Tasmania. Coles Bay, near boatramp (42‘7.0'S

148"I7.0'E), 0.5 m, Zostera sediment, airlift. 21 Apr

1985. R.S. Wilson (stn TAS 18), NMV F88282.

South Australia. Upper Spencer Gulf (33’16'S

137"51 'E), 16 km SW of First Creek, subtidal sand. 12.1

m, T.J. Ward and P.C. Young (stn 795 CG/7), AM

W2I787; Spencer Gulf, Sir Joseph Banks, Group

Reevesby Is, Home Bay South, seagrass bed. upper sub-

tidal, 22 Jan 1986, S.A. Parker. SAM T.E5831 (32

specimens); Pt Lincoln. SAM T.E58.59 (I specimen,

epitoke).

Western Australia. Stuart Head, Yacht Club, Princess

Royal Harbour (35"04'S, 117"55' E), Posidonia aus¬

tralis, 1 m, handheld corer on SCUBA, P. Hutchings,

Jan 1988 (site 7), AM W268II; Bramble Point.

Princess Royal Harbour (35”02'S, 117'55'E), Posidonia

sinuosa, 2.5 m. handheld corer on SCUBA, P. Hutch¬

ings, Jan 1988 (site 11), AM W268I2.

Description. Size range of material examined

from 3 mm long, 15 chaetigers and le.ss than 1

mm wide to 22 mm, 70 chaetigers. 1.5 mm wide

(complete specimens); from 10 mm, 20

chaetigers, 2 mm wide to 38 mm, 58 chaetigers. 4

mm wide (anterior fragments). Pharynx with con¬

ical paragnaths and bars in Area IV, paragnath

counts for 29 specimens includes: I = 0-4; II =

6-27; HI = 1-14; IV = 1-18, in addition 2-7 bars

on each side; V = 0-1; VI = 2-16, usually less

than 10; VII-VIII = 5-30. Glandular patches in

notopodia present from midbody chaetigers.

Prechaetal notopodial lobe (as described by Wil¬

son, 1984) absent but notopodial acicular process

present in chaetigers 5-25.

Habitat. Some of the material reported here was

collected in the intertidal and upper subtidal,

extending the depth-range for this species from

intertidal to 51 m. The shallower records are

from seagrass-beds including Posidonia and

Zostera.

Distribution. Neanthes bassi is recorded for the

first time in Western Australia at Albany and in

South Australia from Spencer Gulf. This species

is only known from southern Australia, Albany,

WA, to off Lakes Entrance. Vic., including north

and east coasts of Tasmania.

Remarks. The material examined agrees well with

the original description, although parapodial char¬

acters are reinterpreted. Numbers of paragnaths

are extended for some Areas. Dorsal pigment

spots described from the material in the original

description are absent in some specimens from

Spencer Gulf (SAM T.E5831), and tend to vary in

prominence in other specimens.

Neanthes tasmani sp. nov.

Figure 1

Neanthes cf. bassi Wilson. 1984: 212.

Material examined. Holotype. Eastern Bass Strait, 100

km off North Point, Flinders 1.. (3r51.8'S. 148°26.5'E).

130 m, tine sand, Smith-Macintyre grab, R. Wilson, 1 s'

Nov 1981. RV Tangaroa (stn BSS 170-G), NMV

F500I8.

Paratypes. Eastern Bass Strait, 100 km off North

Point. Flinders 1. (3r52.6'S, 148°25.2'E), 140 m,

WHOI epibenthic sled, R. Wilson, 15 Nov 1981, RV

Tanproa (stn BSS 170-S), NMV F500I9-F50021 (3

specimens). Tasmania, E of Maria I. (42'36.0'S,

I48'I0.0'E), 75 m, fine bryozoa and shell, WHOI

epibenthic sled, R.S. Wilson, 23 Apr 1985 (stn TAS

30). AM W27491 (2 specimens).

Description. Holotype, complete specimen 9 mm

long for 44 chaetigers, 1.5 mm wide. Body robust,

flattened, tapering posteriorly. Colour in alcohol

creamy yellow. Prostomium slightly wider than

long. Two pairs of dark red to black eyes. One

pair of antennae 1.5 times longer than palps. Palps

stout with conical palpostyles. wider than long.

Four pairs of tentacular cirri, faintly annulated,

longest (posterodorsal) pair reaching to chaetiger

9. The first (apodous) segment broadened, enclos¬

ing the posterior part of prostomium. Pharynx

with translucent yellow to light brown jaws with

7 teeth. Conical paragnaths present on both rings

and short bars also present in Area IV. arranged as

follows: I = 3 in a longitudinal row; II = 18 (left),

20 (right), 2 rows in an arc; III ^ 5 in a diamond¬

shaped group; IV = 15 (left), 7 (right), bars

missing on left side, 4 bars on right side; V = 0;

VI = 3 (left), 3 (right), in one row; VII-VIII = 8,

in single row.

NEW SPECIES OF A?£^;vr//£5 (POLYCHAETA)

329

Figure 1. Necmthes tasmani sp. nov. llolotype, NMV F5()0I8: a, anterior view of chaetiger 3; b, anterior view of

chaetiger 10; c, anterior view of chaetiger 21; d. anterior view of chaetiger 30; g, hetcrogomph falciger from

chaetiger 10.

Paratype, AM W27491: e, hetcrogomph spiniger with short blade from chaetiger 3; f, hetcrogomph falciger from

chaetiger 30.

Abbreviations: postch = postchactal lobe, prech = prechaetal lobe, aciepro = acicular process. Scale bars A-D,

0.1 mm; E-G, 0.01 mm. Drawings of the end-view of parapodia are not to scale. End-views are drawn with the ante¬

rior end of the specimens to the right.

330

T. BAKKEN

Dorsal cirrus slightly shorter than ventral

notopodial ligule in anterior chaetigers, becoming

longer posteriorly up to 1.5 times longer in the

few posteriormost chaetigers, basally attached

throughout. Dorsal notopodial ligule conical,

rounded anteriorly becoming pointed posteriorly;

not more than 1.5 times as long as ventral ligule

anteriorly, becoming smaller than ventral ligule in

posterior chaetigers (Fig. ID). Ventral ligule con-

ieal anteriorly slender and pointed in midbody

and posterior chaetigers. Distinct digililbrm

prechaetal lobe present in chaetigers 5-15 (Fig.

IB). Glandular patches in mid-body and posterior

chaetigers.

Neuropodia with prominent inferior lobe in

anterior and midbody chaetigers, redueed posteri¬

orly. A distinct digitiform postchaetal lobe pre¬

sent in chaetigers 1-22 (Figs lA-C). Ventral

neuropodial ligule conieal, rounded, as long as

aeicular ligule in anterior chaetigers. smaller,

pointed and reduced in posterior chaetigers. Ven¬

tral cirri approximately 0.5 times as long as

neuropodial aeicular ligule, basally attaehed

throughout.

Notochaetae homogomph spinigers, in single

straight row between dorsal and ventral ligules.

Neuroehaetae dorsal fascicle homogomph

spinigers and heterogomph falcigers, falcigers

with long blades in anterior chaetigers, with short

blades in posterior ones (from approximately

chaetiger 10). Ventral fascieic heterogomph

spinigers with both short and long blades in ante¬

rior chaetigers (Fig. IE), including falcigers (Fig.

IG) from approximately chaetiger 10. in posterior

ehaetigers heterogomph spinigers (long blades)

and heterogomph falcigers (Fig. 1F). Paired cirri-

form pygidial cirri reaching back 6 chaetigers.

Variation. Variations of 5 paratypes, size range 3

mm long for 14 chaetigers, less than 1 mm wide

to 12 mm long for 30 chaetigers. 2 mm wide

(anterior fragments), one complete specimen

(NMV F88283) 9 mm long for 41 chaetigers, 1

mm wide. Eyes black in paratypes (NMV

F88283). Paragnath counts for 5 specimens as fol¬

lows: 1 =2-4; II = 10-25; 111 =3-13, in transver¬

sal rows; IV = 8-23, also short bars 0-3 on either

side; V = 0; VI = 2-8, in circular groups;

VII Vlll = 4-7 in a single row. Paragnaths often

pale and llattened with large base, including short

bars present in Area IV. These might de difficult

to distinguish from cones, as the cones are some¬

times placed clo.sely together.

Notopodial prechaetal lobe present from

chaetiger 5-15. Neuropodial postchaetal lobe pre¬

sent from chaetiger I to 18-20.

Remarks. Neaiilhes tusmani closely resembles

N. bassi but can be distinguished by the absence

of a prechaetal notopodial lobe. Other distin¬

guishing characteristics are given in Table 1.

Neaiuhes lasmani also resembles N. flindersi Wil¬

son, 1984 as this species has notopodial

prechaetal lobes and neuropodial postchcatal

lobes in the same range as N. tasmani. The two

species can be distinguished by length of dorsal

parapodial cirri which in A. flinJersi is 1.5 2.5

times the dorsal notopodial ligule, and by the

absence of bars in Area IV in the latter species.

Neanihes kerguelensis McIntosh, 1885 and N.

nanhaiensis Wu, Sun and Yang. 1985 both have

Table I. Comparison of Neanthes hassi and N. tasmani.

Neanthes tasmani

Neanthes bassi

Area 1 paragnaths

2-4

Area 11 paragnaths

10-25

5-27

Area 111 paragnaths

3-13

1-14

Area IV (bars) paragnaths

7-23 (0^)

1-18 (2-7)

Area V paragnaths

0-1

Area VI paragnaths

2-8

2-16

Area Vll-VIII paragnaths

4-8

5-30

Length of dorsal cirri (times length

of dorsal notopodial ligule)

<1-1.5

Notopodial dorsal ligule

reduced in posterior chaetigers

as long as notopodial ventral

ligule throughout

Notopodial prechatal lobe

present, in chaetigers 3-15

absent, aeicular process in

chaetigers 5-25

Neuropodial postchaetal lobe

present, in chaetigers 1 -21

present, in chaetigers 1-12

Habitat

fine biogenic sand, 75-140 m

sand, shell and mud, seagrass

beds, intertidal to 51 m

NEW SPECIES OF NEANTHES (POLYCHAETA)

331

notopodial prechaetal and neuropodial

postchaetal lobes but can be distinguished from

N. tasmani by the distribution of paragnath num¬

bers and the length and fonn of the dorsal cirri

and dorsal notopodial ligule (Wu et al., 1985;

Wilson. 1984).

The reduced size of the dorsal notopodial ligule

in the posteriormost chaetigers in N. tasmani

(observed in the two complete specimens only) is

also found in N. isolala Hutchings and Tur\'cy,

1982 and N. uniseriata Hutchings and Turvey,

1982, but these species can be distinguished from

N. tasmani by the absence of notopodial

prechaetal and neuropodial postchaetal lobes, and

by the number of paragnaths (Hutchings and

Turvey, 1982; Wilson, 1984).

The faint short bars found in Area IV in N. tas¬

mani are different from those present in N. hassi,

which are very distinct and well developed, even

in small specimens with body width less than I

mm. In contrast, even in the largest specimens of

N. tasmani the bars are faint and poorly devel¬

oped. although clearly present. More material is

needed to judge if there are further differences

between the two species in this respect.

N. tasmani has a deeper depth range than the

closely related N. bassi. Specimens were found

from 75 m to 140 m. while N. bassi has a depth

range from intertidal to 51 m.

Etymology^ The species is named after Abel Tas¬

man who was among the first Europeans to

explore the southern seas of Australia and to set

foot on Tasmania.

Distribution. Eastern Tasmania (east of Maria

Island) and eastern Bass Strait; 75-140 m, fine

biogenic sand.

Acknowledgements

1 am grateful to Robin Wilson (Museum Victoria)

for providing laboratory space and access to

material, and for comments on the manuscript.

Pat Hutchings and Rate Attwood (Australian

Museum) provided access to Australian Museum

and South Australian Museum material. Com¬

ments from two anonymous referees improved

the quality of the manuscript. This work was sup¬

ported by a grant from The Research Council of

Norway (grant no 138717/410).

References

Hutchings, P.A. and Turvey, S.P., 1982. The Nereidi-

dae of South Australia. Transactions of the Royal

Society of South Australia 106: 93-144.

Hyllebcrg. J. and Natcewalhana A., 1988. Polychaetes

of Thailand. Nereididae (part 2); Ceratocephale

and Gyinnonereis, with description of two new

species and notes on the subfamily Gymnonereidi-

nac. Phuket Marine Biological Center Research

Bulletin 49: 1-20.

Hylleberg, J., Nateewathana A. and Bussarawit S.,

1986. Polychaetes ofThailand. Nereididae (part 1);

Perinereis and Pseudonercis with notes on species

of commercial value. Phuket Marine Biological

Center Research Bulletin 43: 1-22.

Wilson. R.S.. 1984. Neanthes (Polychaeta: Nereididae)

from Victoria with descriptions of two new

species. Proceedings of the Royal Society of Victo¬

ria 96; 209-226.

Wilson. R.S.. 1988. Synonymy of the genus Nec-

toneanthes Imajima. 1972, with Neanthes Kinberg.

1866 (Polychaeta; Nereididae). Proceedings of the

Biological Society of Washington 101 (1): 4-10.

Wu. B., Sun. R. and Yang. D., 1985. Nereidae (Poly-

chaetous Annelids) of the Chinese Coast. Springer

Verlag: Berlin. 234 pp.

Memoirs of the Museum of Victoria 59(2); 333-336 (2002)

CORYSTOLONA, A NEW HYDROID GENUS (LEPTOLIDA: LEPTOTHECATAE)

FROM SOUTHERN AUSTRALIA

Jeanette E. Watson

Honorary Associate, Invertebrate Zoology, Museum Victoria, P.O. Box 666E, Melbourne, Vic. 3001, Australia

(e-mail; jewatson@bigpond.com) ^

Abstract

Watson, J.E., 2002. Corystolona, a new hydroid genus (Leptolida; Leptothecatae) from south¬

ern Australia. Memoirs of Museum Victoria 59(2); 333-336.

A hydroid first described as Reticularia annulala Watson, 1973 from the Great Australian

Bight is the type species of a new genus Corystolona in the family Clavidae, subfamily Cory-

dendriinae Calder, 1988. The simple, stolonal colonies of Corystolona lack nematophores and

have internal gonophores.

Introduction

Abundant colonies of a hydroid previously

described as Reticularia annulala Watson, 1973

were collected in a survey of the Port of Launce¬

ston, Tasmania, Australia, in May, 2001. The

colonies were found on wharf pilings at depths of

1-4 m below low water mark. The hydroid

assemblage on the piles included the common

southern Australian species, Pennaria wilsoni

Bale, 1913, Bimeria australis Blackburn, 1937

and Aglaophenia pluinosa Bale, 1882. The find¬

ing of several colonies with intact, extended

hydranths and gonophores enclosed within the

hydrocaulus showed R. annulala to be referrable

to the family Clavidae McCrady, 1859 and to

belong, with slight emendation, to the subfamily

Corydendriinae Calder, 1988. Type and other

material is held in the Museum of Victoria,

Melbourne, Australia (NMV).

Clavidae McCrady, 1859

Corydendriinae Calder, 1988

Diagnosis (emended). Clavid hydroids with

stolonal or erect colonies; branches of erect

colonies adnate to hydrocaulus for a varying dis¬

tance basally. Hydranths elongate, more or less

cylindrical, not polymorphic; tentacles filifonn,

scattered over much of hydranth. Nematophores

absent. Gonophores fixed, external or internal

sporosacs or free medusae, arising from

hydrorhiza, hydrocaulus, branches or pedicels but

not from body of hydranths.

Remarks. Calder (1988) discussed the status

of the family Clavidae, considering it to be

sufficiently disparate to warrant separation into

several subfamilies additional to the nominal sub¬

family Clavinae McCrady, 1859 and proposed

the subfamily Corydendriinae Calder, 1988. With

one difference the present specimens accord with

the diagnosis of the Corydendriinae, this being

that it is not clearly stated whether gonophores

may be external or internal. The diagnosis of

the Corydendriinae is therefore emended to

accommodate both external and internal

gonophores.

Corystolona gen. nov.

Diagnosis. Clavid hydroids with stolonal

colonies, hydrocaulus simple, unbranched, peris-

arc finn, terminating at hydranth base; hydranths

elongate, tubular, hypostome dome-shaped, ten-

taeles filifonn, scattered over hydranth body;

gonophores fixed sporosacs, arising as blind sacs

beside hydranths within perisarc of hydrocaulus.

Type species. Reticularia annulala Watson, 1973.

Elyinolog}’. Refers to relationship with the Cory-

dendriidae and strictly stolonal habit of the

colonies.

Remarks. Corystolona is close to a group of four

clavid genera: Coiydendrium van Beneden, 1844;

Tuhiclava Allman. 1864; Rhizodendrium Calder,

1988; and Merona Norman, 1865. These genera

were rejeeted from consideration for Coiyslolona

annulala because the gonophores of Tuhiclava

are bonie on the hydranth body, those of Rhizo-

dendrittm and Merona are hydrorhizal, and Cor}’-

dendritim as defined by van Beneden (1844),

Millard (1975) and Bouillon (1985) includes only

333

334

J. E. WATSON

those species with erect, ramified colonies, w'ith

either internal or external gonophores. The

strictly stolonal colonies together with internal

gonophores sets Coryslolona apart from these

genera. Coiydendrium brevicaulis Hirohito, 1988

from Japan is a sparsely branched species with

internal gonophores which should be included in

the Corydendriinae. While C. brevicaulis is close

to Coryslolona, the strictly simple, stolonal

hydrocauli of Coryslolona distinguishes it from

Hirohito's species.

Coryslolona anniilala (Watson, 1973)

Figure la-e

Reticularia anniilala Watson, 1973: 164, figs 5, 6.

Material examined. Holotype. NMV GI922,

microslide. NMV G209I, preserved material, remain¬

der of holotype eolony. Pearson Island. South Australia.

17 m, on calcareous bryozoan. NMV F91279 (malinol

mounted microslide, infertile colony), NMV F91280

(malinol mounted microslide, female colony), NMV

F91281 (malinol mounted microslide, male colony),

NMV F91282 (alcohol pre.served material). Port ot

Launceston. Tasmania, colonics from bryozoans on

wharf pilings 1-4 m, Aquenal Pty Ltd. May 2001.

Description (of Tasmanian material). Colonies

comprising a few to many hydrocauli given oil

from poorly adherent tubular stolons; stolons

unbranched or very sparsely branched, occasion¬

ally produced into a loose tangle where growing

end becomes free of substrate; perisarc smooth to

rather crumpled.

Single hydrocauli given off at various angles

and at irregular intervals along stolon.

Perisarcal tube hom-shaped, proximally narrow

at junction with stolon, but without true pedicel,

thereafter widening to become cylindrical; length

very variable, younger tubes olfen quite short and

narrow, mature ones wider, straight or with 1 or 2

bends; sometimes recurved into a loop.

Perisarc of younger hydrocauli thin but finn,

becoming thicker with age; perisarc evenly and

deeply transversely annulated in basal third, cor¬

rugations usually continuing throughout length of

tube, occasionally becoming smoother dislally.

Margin circular, perisarc of rim usually thinner

and more fragile than rest of tube, sometimes tom

by eruption of hydranth.

Hydranth long, cylindrical, with tall, narrowly

dome-shaped hypostome with deep central slit;

body with 26-30 scattered filifomi tentacles, an

ineipient ring of 4-5 tentacles below hypostome;

tentacles longest in mid-region, becoming shorter

proximally; tentacles armed with prominent rings

of nematocysts. Base of hydranth enclosed in a

long funnel-shaped sheath of tissue inside tube;

when hydranth retracted sheath becomes a

muscular supporting girdle.

Colonies dioecious, gonophores laying beside

hydranth deep within perisarcal tube; gonophore

digitate with blunt distal end, no spadix, female

containing 10-12 small ova, male of same

shape as female but with undifferentiated

spermatogenic mass.

Two kinds of nematocysts in hydranths: (I)

desmoncmes, capsule broadly droplet-shaped,

6.4-f>.8 X 3.6-4.4 pm; and (2) euryteles, capsule

elongate droplet-shaped, 8.4-10.4 x 4.0 4.8 //m,

set in rings in tentacles. Both types of nemato¬

cysts very abundant; none found discharged.

Colour. Perisarc of younger hydrocaulinc tubes

translucent white, darkening to hom-coloured

with thickening of perisarc. Young stolons

opaque white. Hydranth and tentacles (fomiol

preserved) white, a trace of pink in some

liydranths.

Measurements of mature specimens (/vm):

Hydrorhiza

Diameter of stolon

140-176

Hydrocaulus

Length of mature tube

640^,000

Proximal width

152-176

Diameter at margin

320-560

(mature tube)

Hydranth

Length extended

540-900

(preserved material)

Remarks. The small sample of Reticularia annul-

ala from Pearson 1. in the eastern Great Aus¬

tralian Bight had deeply withdrawn, partly

decomposed, sterile hydranths and disposition of

the specimen on the bryozoan host was such that

the bases of the hydrocauli were obscured; the

specimen was thus mistakenly referred to Relicu-

laria Thomson, 1853 (junior synonytn, FUellunu

Hincks, 1868). The smaller dimensions of the

type compared with those of the present speci¬

mens is almost certainly due its being a young,

infertile colony while the Tasmanian material

ranges from young to aged. The difference in size

of immature hydrocauli compared with older ones

is a striking feature of the present material.

In 1839 d'Orbigny described "Tubidaria

nigosa" collected from a jetty south of the Rio

Negro in Patagonia. He considered the specimen

remarkable by its long annulated tubes but did not

mention any sol) parts. Bedot (1905) considered

the material indetenninablc but Leloup (1937)

referred it to Stephano.scyphus. Notwithstanding

A NEW HYDROID GENUS

335

Figure 1. a, hydrorhiza, hydrocaulus and hydranlh with young male gonophore. b, mature female gonophore with

ova. c, rings of nematoeysts in tentacle, d, desmoneme, undischarged, e, cuytele, undischarged, from tentacles. Scale

bar: a, b, 0.5 mm; c, 0.2 mm; d, e, 10 mm.

the opinion of these authors the similarities

between d'Orbigny’s speeimen and Corystolom

anmdeUa suggests that it may be the same or sim¬

ilar speeies. As the type material of T. rugosa

appears to be lost the question cannot be settled

until the finding of live material.

Little information could be gleaned about sub¬

strate preferences of Coryslolona annukita from

the present samples since most colonics had been

detached frotn their substrate in the laboratory.

However, like the Pearson 1. specimens several

colonies were attached to a bryozoan, in this case.

Amalhia sp. Because of their poor adherence long

strands of stolons could be easily freed from the

substrate. The tightly contracted tentacles with

many rings of nematoeysts suggest that the ten¬

tacles are probably very extensible in life. The

deeply slit hypostome indicates a capacity for

wide expansion to engulf quite large prey.

Because of retraction during preservation it could

not be unequivocally established whether the

mature hydranth is fully retractable into the tube,

but the strong muscular band below the hydranth

suggests that it is capable of doing so.

336

J. E. WATSON

Acknowledgments

I thank the management of Aquenal Pty Ltd

Environmental Consultants, Tasmania, for pro¬

viding material for examination. Thanks are also

due to Dr W. Vervoort of the Natural History

Museum, Leiden, The Netherlands, and Dr D.R.

Calder of the Royal Ontario Museum, Canada for

helpful advice.

References

Allman, G.J., 1864. On the construction and limitation

of genera among the Hydroida. Annals and

Magazine of Natural History (3) 13(77): 345-380.

Bale. W.M.. 1882. On the Hydroida of south-eastern

Australia, with descriptions of supposed new

species, and notes on the genus Aglaophenia.

Journal of the Microscopical Societv of Victoria 2:

15-48.

Bale. W.M., 1913. Further notes on Australian

hy droids. 11. Proceedings of the Royal Society of

Victoria (new series) 26: 114-147.

Bedot, M., 1905. Materiaux pour scrvir a Thistoire des

hydroides. 2me Periode. Revue Suisse de Zoologie

13: 1-183.

Blackburn. M., 1937b. Notes on Australian Hydrozoa,

with descriptions of two new species. Proceedings

of the Royal Societ)’ of Victoria (new series) 50:

170-181.

Bouillon, J., 1985. Essai de elassification des

Hydropolypes — Hydromeduses (Hydrozoa-

Cnidaria). Indo-Malayan Zoology 2: 29-243.

Calder, D.R.. 1988. Shallow-water hydroids of

Bermuda. The Athecatae. Life Science Con¬

tributions Royal Ontario Museum 148: 1-107.

D’Orbigny, A., 1839. Zoophytes. Pp. 7-28 in: Voyage

dans I 'Amerique meridionale, execute pendant les

annees 1826-1833. Vol. 5.

Hincks, T., 1868. A history of the British hydroid

zoophytes. John Van Voorst: London. 338 pp.

Hirohito, Emperor of Japan, 1988. The hydroids of

Sagami Bay. Publications of the Biological

Laboratory of the Imperial Household: Tokyo.

179 pp.

Leloup, E.. 1937. Hydropolypes et Scyphopolypes

recueillis par C. Dawydoff sur les cotes de I’lndo-

chine frangaise. Memoires du Museum Royal

d'Hisloire Naturelle de Belgique 12: 1-73.

McCrady, J., 1859. Gymnopthalmata of Charleston

Harbor. Proceedings of the Elliott Society of

Natural History of Charleston, South Carolina 1:

103-121.

Millard, N.A.H., 1975. Monograph of the Hydroida of

South Africa. Annals of the South African Museum

68: 1-513.

Nonnan, A.M., 1865. On Merona, an undcscribcd

genus of British Hydrozoa. Annals and Magazine

of Natural History (3) 15: 261-262.

Thomson, W.T.C., 1853. Notes on some British

zoophytes. Annals and Magazine of Natural His¬

tory! (2) II; 442-444.

Van Beneden, P.J., 1844. Memoire sur les campanu-

laires de la cote d'Ostende, consideres sous le

rapport physiologique, embryogenique et

zoologique. Nouveaux Memoires de I'Academie

Royal des Sciences et Belles-Lettres de Bruxelles

17: 1^2.

Watson, J.E., 1973. Pearson Island expedition 1969-9.

Hydroids. Transactions of the Royal Society of

South Australia 91: 153-200.

Memoirs of the Museum of Victoria 59(2): 337-354 (2002)

HYDROIDS (CNIDARIA: HYDROZOA) FROM SOUTHERN QUEENSLAND

Jeanette E. Watson

Honorary Associate, Museum Victoria, P.O. Box 666E, Melbourne, Vic. 3001, Australia

e-mail: jewatson(aJbigpond.com

Abstract

Watson, J.E., 2002. Hydroids (Cnidaria: Hydrozoa) from southern Queensland. Memoirs of

Museum Victoria 59(2): 337-354.

Fifteen species of hydroids were collected from a reef off Palm Beach, southern Queensland.

Six records are new for Queensland; several species are circumtropical and two are

cosmopolitan. Symplectoscyphus sihogae is

locality of Indonesia.

Introduction

A collection of hydroids was made on 9 May

2001 by the author using SCUBA from a reef

approximately I km offshore from Palm Beach,

southern Queensland (28°5'52"S, I53°26'34"E).

In this area, a reef system extends 1-2 km off¬

shore along the coast at a depth of 5-15 m, sev¬

eral higher parts of the reef being emergent as

small islets. The water regime in the area is sub¬

tropical, the temperature at the time of the survey

being 26°C. While not subject to .strong currents

the reefs arc exposed to swells of the South

Pacific Ocean.

The reef surrounding Cook Island off Burleigh

Heads was inspected but yielded no hydroids.

This island is a marine sanctuary, the surrounding

waters being closed to recreational fishing; the

invertebrate reef community is heavily grazed by

fishes. In contrast, the reef off Palm Beach, out¬

side the protected area and subject to consider¬

able recreational fishing pressure, has fewer fish

and the invertebrate community is much more

diverse and abundant.

With the exception of Macrorhynchia philip-

piiui and Aglaoplienia sinuosa which grow on

exposed reef surfaces all species w'ere small and

cryptic, mostly occurring in sheltered crevices.

Colonies of M. pliilippina, a tropical species,

were quite small, indicating that it may be near

the southern end of its range. Colonies of

A. sinuosa, the other visually dominant hydroid,

showed evidence of grazing by fish.

The collection, made in one dive over an area

of several hundred square metres of reef, yielded

recorded for the first time outside the type

15 species of hydroids. Six species, Eudendriitm

glomeratum, Hydrodendron daidalum. Salacia

desmoides, Symplectoscyphus sihogae. Agla-

ophenia sinuosa and Ohelia angulo.sa and are

new records for Queensland, having been previ¬

ously recorded from temperate southern Australia

and tropical northern Australia. Pcnnycuik (1959)

recorded 46 species from the Queensland coast

between the New South Wales border and

north of Moreton Bay (latitudes 26-28°S). As

only four species in the present collection (Anten-

nella secundaria, Synlhecium campyhearpum (=

S. patulum), Hincksella cylindrica, Macror¬

hynchia philippina) were recorded from this

region by Pennycuik, the number of known

species from southern Queensland is thus

increased to 57.

Several species in the present collection are cir-

cumtropical in distribution, and two (Halecium

sessile and Antennella secundaria) arc cos¬

mopolitan. Symplectoscyphus sihogae is recorded

for the first time outside the type locality of

Indonesia. The collection also contains a small

colony of a species of Halecium that could not be

matched with any known species and may there¬

fore be new. In the following descriptions only

those synonymies relevant to Australia or the

.southern hemisphere are given. Material is

deposited in Museum Victoria (NMV).

List of Species.

Eudendrium 'Iglomeratum Picard, 1951

Hehellopsis scandens (Bale, 1888)

Halecium sessile Norman, 1867

Halecium sp.

337

338

J. E. WATSON

Hydrodendron daidalum (Watson, 1969).

Salacia desmoides (Torrey, 1902)

Dvnamena qiiadridentala (Ellis and Solander,

1786)

Dynamena qiiadridentala (Ellis and Solander,

1786)

Symplectoscyphus sibogae (Billard, 1924)

SytUhecitim campylocarpum Allman, 1888

Hincksella cylindrica (Bale, 1888)

Antennella secundaria (Gmelin, 1791)

Lyiocarpia brevirostris (Busk, 1852)

Aglaophenia sinuosa Bale, 1888

Macrorhvnchiaphilippina (Kirchenpauer,

1872)

Obelia angulosa Bale 1888

Anthoathccatae

Eudendriidae L. Agassiz, 1862

Eiidendriiiin Ehrenbcrg, 1834

Eudendrium Iglomeratum Picard

Eudendrium glomeratum Picard, 1951: 338.—Picard.

1955: 183.—Teissier, 1965: 14.—Watson, 1985:

213.—Boero and Cornelius. 1987: 244.—Watson,

1996: 78.—Watson, 1999: 7.

Eudendrium genendis von Lcndenfeld, 1885: 351.—

Thornely. 1904: 110.—Jaderholm, 1916:3.

Eudendrium indopacificum Stechow, 1924: 59.

Material examined. Palm Beach, Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91283, several infertile

stems to 4 mm high on barnacle, alcohol preserved.

Description. Stolon tubular, entwining substrate.

Stems unfascicled, unbranched or sparingly

branched in one plane; hydranths terminal on

short pedicels or on single pedicels arising

directly from stolon. Cnidome comprising small

microbasic curyteles in tentacles and macrobasic

euryteles on hydranth body. Colour, stems brown,

hydranths white.

Distribution. Previously known in Australia from

the tropical north and north-western coasts

(Watson. 1999).

Remarks. The material is doubtfully assigned

to E. glomeratum based on the presence of

macrobasic euryteles distributed over the

hydranth body. Although the nematocysts w'ere

not aggregated in pads on the hydranth as is

normal in this species, reduction or loss of dis¬

crete pads and spreading of nematocysts over the

entire hydranth body sometimes occurs (Watson,

pers. obs.). If indeed E. glomeratum it is the most

southerly record of the species in Australian

waters.

Leptothecatae

Lafoeidae A. Agassiz, 1865

Hehellopsis Hadzi, 1913

Hehcllopsis scandens (Bale)

Figure lA

Lafoea scandens Bale. 1888: 758.—Billard, 1904:

481.—Billard. 1906: 174.—Warren. 1908: 272, 341,

349.

Hchella scandens. —Marktanner-Tumeretscher, 1890:

214.—Millard. 1975: 182.—Migotto, 1996: 26.—

Watson, 1996: 78.— Boero et al., 1997: 8.

Hehellopsis scandens. —Vannucci-Mendes 1949:

237.—Vannucci-Mendes, 1950: 85.—Calder, 1991:43,

45. 95.

Material examined. Palm Beach, Queensland, 2 km

offshore, depth 5-15 m. NMV F91284, infertile

colonies epizoic on Dynamena qiiadridentala, alcohol

preserved.

Description. Stolons tubular, very thin, wrinkled,

closely adpressed to the hydrorhiza of Dynamena

quadridentata, then ninning up stem of host.

Hydrotheca tubular, issuing beside or between

hydrothecae of host. Pedicel short, thin, tubular,

wall of hydrotheca obtusely flexed about halfway

along length, creased al bend; rim of hydrotheca

circular, transverse to hydrothecal axis, slightly

thickened and very W'cakly outrolled. Perisarc

rather thin, fragile at hydrothecal margin.

Hydranth with about 12 tentacles deeply with¬

drawn into hydrotheca (preserved material).

Colourless.

Distribution. Cosmopolitan.

Remarks. The narrow stolon of //. scandens can

only be distinguished from that of D. quadriden¬

tata as a thin, almost invisible tube lying closely

beside the broad hydrorhiza of the host. The

colonies of H. .scandens contain neither straight

nor slightly asymmetrical hydrolhecae conform¬

ing to the accepted concept of H. scandens. Vari¬

ous authors (Vervoort, 1968; VervooiT and

Vasseur, 1977; Boero et al.. 1997) have reported

straight and contorted hydrothcca in the same

colony of //. scandens W'hile hydrothecae of

H. scandens from norlhcni Australia were all con-

rorted (Watson, 2000). While it is possible that

//. .scandens may undergo torsion as a response to

ecological conditions it is nevertheless possible

that there may be two closely related species

involved, one with almost symmetrical hydrothe¬

cae and one with contorted hydrothecae. This cati

only be resolved with further collecting of an ade¬

quate live material over a wide geographical

HYDROIDS FROM SOUTHERN QUEENSLAND

339

Figure 1. A, HebeUopsis scandens (Bale, 1888). Three hydrothecac on Dynamem qiiadridentala. B, C, Halecmm

sessile Norman, 1867. B, part of branch; C, stem inlernodes with hydrothecae. D, E. Halecmm sp. D, part of colony

with stolonal hydrothecae and erect stem; E, hydrophores and hydrothecae. F, Hydrodendnm daidaliim (Watson,

1969). Hydrorhiza, stem, hydrotheca and nematophore. Scale bar: A, D, C, 6.5 mm; B, 1 mm; E, 0.1 mm;

F, 0.2 mm.

340

J. E. WATSON

range and detailed investigation of reproductive

strategies.

Haleciidae Hincks,1868

Haled lint Oken, 1815

Haleciiiiii sessile Norman

Figures 1B, C

Halecitim sessile Norman. 1867: 205.—Ritchie,

1911: 812.—Stechow, 1913: 9.—Billard. 1927: 329.—

Picard. 1958: 192.—Ralph, 1958: 331.—Pennycuik,

1959: 174.—Teissier, 1965: 21. -Vervoort. 1966:

100.—Redicr, 1967: 386.—Vervoort, 1968: 95.—

Schmidt. 1972: 42.—Cornelius, 1975: 406.—Millard.

1975: 154.—Watson. 1979:234.—Ramil and Vervoort.

1992: 85.—Watson, 1994: 66.—Cornelius. 1995:

292.—Hirohito, 1995: 27.

Material examined. Palm Beach. Queensland, 2 km off¬

shore, depth 5-15 m, NMV F 91285, sparingly

branched infertile colonies on various invertebrate sub¬

strates in crevices, alcohol preserved.

Description. Flydrorhiza reptant on substrate,

stolons tubular. Stems monosiphonic, to 4 mm

high, straight to slightly zig-zag, simple or

sparsely branched near base. Stem intemodes

widening distally to hydrophore, nodes trans¬

verse, deep. Hydrophore short, primary

hydrotheca very shallow, sessile or very closely

adpressed to intemode at level of node, shallow

dish-shaped, sloping gently downwards away

from internode, diaphragm distinct with a row of

desmocytes above. Margin circular, not everted.

Secondary hydrophorcs rare, on lower parts of

stems, issuing from diaphragm of preceding

hydrotheca, adcauline wall deeply indented at

level of diaphragm. Perisarc smooth throughout,

thick on proximal parts of stems, becoming very

thin on hydrotheca. I lydranths robust, very exten¬

sible with about 24 stubby tentacles. Colour yel¬

lowish-brown.

Disiribiilion. Cosmopolitan. Recorded in Aus¬

tralia from southern Queensland to Bass Strait.

Remarks. Secondary intemodes are sometimes

present between the main stem intemodes. The

material is assigned to Haleciitm sessile because

of the almost straight stems and rather shallow,

sessile primary hydrothccae. Although H. sessile

was originally described from the cool temperate

northern Atlantic there are now many records

from other parts of the world, although some may

be incorrect. Until a full world review of the

species is undertaken, it is best to refer the

Queensland material to H. sessile.

Halediiiii sp.

Figures 1D, E

Material examined. Palm Beach. Queensland, 2 km off¬

shore, depth 5-15 m, small infertile colonies on Ohelia

angulosa (NMV F9I294) and Salacia desmoides

(NMV F91287), alcohol preserved.

Description. Stolon straggling, tubular, faintly to

strongly rugose. Stems simple, very short, arising

at inter\als from hydrorhiza; basal stem intem¬

odes long, tubular, smooth, nodes deep, trans¬

verse. intemode tumescent above node. Primary

hydrophorcs given off singly from stolons and

di.stally on stem intemodes, outwardly bent, suc¬

ceeding intemode arising from a short apophysis

below hydrophore. Primary hydrophore fairly

short, cylindrical, surmounted by a deep, bowl¬

shaped hydrotheca; rim of hydrotheca strongly

outrolled; diaphragm distinct, a clearly visible

ring of peg-shaped desmocytes above. Secondary

and tertiary hydrophorcs arising from diaphragm

of primary hydrotheca, usually several proximal

constrictions in perisarc of hydrophore just above

diaphragm. Perisarc of colony rather thin

throughout. Hydranth too contracted for tentacle

count. Colour, white (preserved material).

Remarks. The colony is so minute that it was

noticed only during microscopical examination of

Obelkt angulosa. The rather tangled stolon is eas¬

ily detached from the substrate, the few stems

being difficult to dilTerentiate from the stolons.

The specimen bears no resemblance to any of the

smaller species of Halecitim known from Aus¬

tralian waters, In size of colony and morphology

of the hydrotheca it somewhat resembles H. pyg-

maetim Fraser, 1911; but dimensions are greater

than those given by Hirohito (1995) for that

species. Since many haleciids cannot be accu¬

rately identified in the absence of the gonosome,

the material is not presently assigned to species.

Hydrodendron Hincks, 1874

Hydrodeitdroii daidaliiiii (Watson)

Figure IF

Scoreshia daidaia Watson, 1969: 111.—Watson,

1979: 234.—Watson. 1982: 92. -Stranks, 1993: 6.

Hydrodendron daidalum. —Rees and Vervoort.

1987: 22.

Material e.xamined Palm Beach, Queensland, 2 km olT-

shore. depth 5-15 m. NMV F91286. infertile colony on

the brown alga Zonaria crenata, alcohol preserved.

Description. Hydrorhiza a reticulating network

covering algal surface; stolons very wide, flat and

HYDROIDS FROM SOUTHERN QUEENSLAND

341

ribbon-like, outer flanges with dark excavations.

Hydrotheca distal on a short, thick, deeply rugose

pedicel, deep bowl-shaped, rim circular, slightly

everted. Nematothecae small, elongate urn¬

shaped, inserted on hydrorhiza beside hydrothecal

pedicel, margin circular with everted rim.

Hydranth with thick dome-shaped hypostome and

20-24 tentacles too large to retract into

hydrotheca. Nematophore long, extensile, armed

with a distal cluster of nematocysts. Colour, pale

honey-yellow.

Dislrihulion. This record extends the range of H.

daiJalum from southern Australian coastline to

warmer subtropical waters of the central east

coast. Endemic to Australia.

Remarks. H. daidalum is an obligate epiphyte of

the small brown alga Zonaria crenata, the

hydrorhiza and widely spaced pedicels fomiing

an easily recognisable network on the algal thal-

lus (Watson, 1969). Although Rees and Vervoort

(1987) synonymised Scoreshia with Hydroden-

dron there arc nevertheless several points of dif¬

ference such as the simple, pedicellate stems and

stolonal nematothecae that set H. daidalum apart

from the typically branched colonies and cauline

nematothecae of the accepted concept of

Uydrodendron.

Sertulariidae Lamouroux, 1812

Salacia Lamouroux, 1816

Salacia desmoides (Torrey)

Figure 2A, B

Seiiularia desmoides Torrey, 1902: 65.—Billard,

1924a: 66.—Fraser, 1938: 110.—Fraser, 1948: 247.—

Stechow, 1923: 213.—Millard. 1975: 274.—Watson.

1996: 78.—Watson, 1997: 518.—Medel and Vervoort,

1998: 30.

Material examined. Palm Beach, Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91287, infertile colony on

bryozoan, alcohol presen ed.

Description. Hydrorhiza loosely replant on sub¬

strate, stolon tubular. Stems short, unbranched,

proximal stem of same diameter as stolon, athe-

cate, ending in a strong V-shaped joint; stem

thereafter thecate. Internodes slender, conical,

nodes usually V-shaped, sometimes transverse.

Hydrothecae biscriate, paired, tubular, situated

about two-thirds distance up inlemode, adcauline

walls conjoined, parallel to intcrnodal axis,

straight to slightly convex, free adcauline wall

directed outwards in convex curve, abcauline wall

sinuous to flatly concave, floor of hydrotheca

transverse to intemode, straight or downwardly

convex. Hydrotheca narrowing somewhat to mar¬

gin; margin oblique to both intemodal and

hydrothecal axes, operculum of one valve

attached inside abcauline margin. Perisarc

smooth, rather thick in lower stem region, thinner

but firm on hydrothecac. Hydranth with about 20

tentacles. Colour, deep yellow.

Distribution. California, south-west Indian

Ocean, Mediterranean Sea. Recorded previously

in Australia from Houtman Abrolhos Is, Western

Australia. A new record for the Australian east

coast.

Remarks. The intemodes of the present material

are much shorter and the hydrothecae thus closer

together than in specimens described from the

Houtman Abrolhos Is by Watson (1997). The

oblique margin with abcauline opercular valve is

characteristic of the species.

Dynamena Lamouroux, 1812

Dynamena qiiadridentata (Ellis and Solander)

Figure 2C-F

Seiiularia qiiadridentata Ellis and Solander, 1786:

57.—Lamarck. 1816:2, 121.

Dvnamena quadridentata. —Billard, 1925: 194,

222!—Trebilcock, 1928: 23.—Blackburn. 1938:320.—

Blackburn, 1942: 113.—Pcnnycuik, 1959: 193.—

Ralph, 1961:790.—Mammen,l965: 49.—Ralph, 1966:

159.—Shepherd and Watson, 1970: 140.—Millard,

1975: 266.—Gibbons and Ryland, 1989: 411.—Calder,

1993: 68.—Vervoort, 1993: 108.—Calder, 1995:

543.—Hirohito. 1995: 176.—Watson. 1996: 78.—Wat¬

son, 1997: 520.—Watson. 2000: 15

Pasya qiiadridentata. —Stechow, 1922: 148.—Ste¬

chow, 1923: 166.—Fraser, 1948: 239.

Pasythea (Seiiularia) quadridentata. —Lamouroux,

1812: 183.—Lamouroux. 1816: 156.—Whitelegge,

1889: 193.—Nutting, 1927: 226.—Yamada, 1959: 58.

Dynamena lhankasseriensis Mammen, 1965: 48.

Materiale.xamined. Palm Beach, Queensland, 2 km ofl-

shore, depth 5-15 m. NMV F9I288. sparingly fertile

colony on the bryozoan Fhisira sp. and infertile colony

on red alga, alcohol preserved.

Description. Hydrorhiza replant on substrate,

stolon flat, wide and ribbon-like, edged by a

llange and canal with irregular, cusp-like indenta¬

tions. Stems short, 2-4 mm long, widely spaced

along hydrorhiza, proximal stem intemode athe-

cate with a strongly twisted V-shaped distal node,

internodes thereafter hydrothecate, vv'ith 1—4 pairs

of hydrothecae, nodes between sets of hydro¬

thecae obscure, usually represented by constric¬

tion of perisarc. Hydrothecae biscriate, opposite.

342

J. E. WATSON

Figure 2. A, B, Salacia desmoides (Torrey, 1902). A, part of stem; B, intemodes with hydrothecae. 2C- F, Dyna~

menu quadridentata (Ellis and Solander, 1786). C, hydrorhiza and hydrothecate stem internode; D, anterior view of

hydrotheca; E, lateral view of hydrotheca with operculum; F, gonothecae. Scale bar; A, F, I mm; B, C, 0.5 mm;

D, E, 0.1 mm.

elongate saccate, adcauline walls in partial con¬

tact across intemode, conjoined walls straight to

convex, remaining adnate and free adcauline wall

bent outwards to margin, floor of hydrotheca

transverse to decidedly rounded. In stems with

single pairs of hydrothecae the pairs separated by

a narrow intemodal neck; when more than one

pair on intemode, these closely adpressed,

slightly overlapping, adcauline hydrothccal wall

overlapping base of that above; free adcauline

wall short, convex; abcauline wall complexly

curved to almost straight; basal abcauline wall of

first pair of group with pronounced lateral

swelling, base of higher pairs acutely rounded,

fitting into intemode.

Floor of basal group transverse, thick, a basal

abcauline swelling forming a wide tunnel through

perisarc; base of adcauline wall with large knob,

floor penetrated by a central funnel-shaped

hydropore, directed upwards. Margin with a pair

of broad opposite lateral lobes, adcauline wall

thickened and recurved into a cusp; operculum

delicate, of 2 valves. Perisarc of stems and

hydrotheca thick, thinning to hydrothccal margin.

llydranth with about 16 tentacles on a short,

wrinkled column, no caecum. Gonothcca large,

arising from hydrorhiza on a moderately long

cylindrical pedicel; barrel-shaped, widest about

middle; body with 6-8 deep annular corrugations,

apex transverse, truncate, with thickened rim.

HYDROIDS FROM SOUTHERN QUEENSLAND

343

Gonophore slender, cigar-shaped containing

numerous small ova. Colour, honey-brown.

Distribution. Circumglobal in tropical waters.

Previously recorded from tropical Australia.

Remarks. The annular corrugations on the

gonothecae are usually not all the same depth

around the body, tending to become shallower on

one side. In the present material there is no sub¬

marginal abcauline intrathecal perisarcal thicken¬

ing as described in specimens from Darwin (Wat¬

son, 2000), the colonies being more like those

reported from the Houtman Abrolhos Is (Watson,

1997). In most hydrothecae a strand of tissue

infested with numerous large ovoid diatoms

extends from the hydranth along the inner

abcauline wall of the hydrotheca towards the mar¬

gin. Hehellopsis scandens is a common epizoite

of the colonies.

Sertularella Gray, 1848

Sertularella minuscula Billard

Figure 3A-E

Sertiiiarella minuscula Billard, 1924b; 648.—

Billard, 1925: 139.—Leloup, 1932: 161.—Pennycuik,

1959: 195.—Hirohito, 1974: 18.—Cooke, 1975:98.

Material examined. Palm Beach, Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91289, small fertile

colony on test of a solitary ascidian, alcohol preserved.

Description. Stolon tubular, rugose; stems to 5

mm high, unbranched, lower stem of same

diameter as stolon, proximal intemode athecate,

stem thereafter thecate, intemodes with 2

alternate hydrothecae, nodes distinct, oblique,

sloping alternately left and right, usually a small

tumescence above and below node.

Hydrothecae on both stolon and caulus, all of

Figure 3. A-E, Sertularella minuscula Billard, 1924. A, stolon with single hydrothecae and erect thecate stem;

B, hydrothecate stem intemodes; C, anterior view of hydrotheca with operculum; D, lateral view of hydrotheca and

operculum; E, female gonotheca. Scale bar: A, 1 mm; C, D, 0.1 mm; B, E, 0.5 mm.

344

J. E. WATSON

same shape. Stolonal hydrothecae widely spaced,

on a short pedicel, cauline hydrothecae sessile,

displaced slightly towards front of stem;

hydrotheca tubular, narrowing slightly to margin,

adcauline wall convex, about half free of inter¬

node, abcauline wall contiguous with border of

intemode, almost straight to weakly concave;

floor of hydrotheca transverse to intemodal axis;

margin quadrangular (anterior view), rim with 4

cusps, lateral cusps sharp with deep semicircular

embayment between, operculum of four fragile

triangular valves. Hydranth with 10-12 tentacles;

no evidence of an abcauline caecum in contracted

or extended hydranths. Gonotheca irregularly

obovoid, usually widening distally, body deeply

crumpled, some gonothecae with 1 or 2 deep

flanges, body narrowing into a rather wide,

smooth pedicel, some arising from within stolonal

hydrothecae, apex of gonotheca a short inverted

funnel, orifice circular. An oval body in 1

gonotheca may be an ovum. Perisarc rather thin,

fragile at hydrothecal margin, operculum and

gonotheca. Colour, colony pale yellowish-white.

Distribution. Indonesia, Caribbean, Microne¬

sia, Japan and Queensland.

Remarks. The species is remarkable for its

small size and delicacy and the crumpled appear¬

ance of the gonotheca.

Symplectoscyplius

Marktanner-Tumeretscher, 1890

Syniplectoscyphiis sibogae (Billard)

Figure 4A, B

Symplecloscyphii.'! sibogae Billard, 1924; 69.—

Billard. 1925; 166.—Vervoort. 1993; 241.

Material examined Palm Beach, Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91295, infertile stem

detached from substrate, malinol mounted microslide

preparation.

Description. Stolon thick-walled, tubular. Stem 4

mm high, straight, with 5 alternate hydrothecae,

an indistinct transverse basal node between

hydrorhiza and first hydrotheca; no other obvious

nodes. Hydrothecac large, tubular, arching out¬

wards from stem, narrowing to margin, adnate

adcauline wall short, parallel with intemode, free

adcauline wall weakly convex to almost straight,

abcauline wall contiguous with border of inter¬

node, faintly concave; floor of hydrotheca nar¬

row, transverse to intemodal axis. Margin everted

with 3 fairly sharp cusps with shallow embay-

ments between; a large internal submarginal cusp

at base of each embayment; operculum of 3

delicate triangular valves. Perisarc thick and

Figure 4. A, B, Symplecloscyphus sibogae (Billard,

1924). A, part of stem; B, hydrotheca. Scale bar; A, 1

mm, B, 0.5 mm.

smooth throughout; opercular valves thick.

Colour white.

Distribution. This is the second record of the

species, the original record being from the type

locality in Indonesia.

Remarks. Although the small colony is obviously

juvenile, it nevertheless conforms to descriptions

of the species. Because of their position below the

embayments the marginal cusps are rather diffi¬

cult to see. The species is notable for its smooth

perisarc, and large, elegant hydrothecae with

great length free of the intemode,

Syntheciidae Marktanner-Tumeretscher, 1890

Synthecium Allman, 1872

Synthecium campylocarpum Allman

Figure 5A-C

Synthecium campylocarpum Allman, 1888; 78.—

Marktanner-Tumeretscher, 1890; 248.—Farquhar,

1896; 466.—Stechow, 1913; 127.—Jaderholm, 1919;

14.—Totton, 1930; 169.—Ralph, 1958; 347.—

Yamada, 1959; 52.—Hirohito, 1969; 18.—Watson,

1996; 78.—Watson, 2000; 40.

HYDROIDS FROM SOUTHERN QUEENSLAND

345

Figure 5. A-C, Synthecium campylocarptim Allman, 1888. A, whole stem: B, hydrothecate hydrocladial inter-

nodes; C. hydrotheca showing regrowth of margin from within hydrotheca. D, Hincksella cylindrica (Bale, 1888).

Part of stem. E-F, AnIennelUi secundaria (Gmelin, 1791). E, fertile hydrocladium; F, female gonotheea (after

Watson. 2000). Scale bar: A, 2.5 mm; E. I mm; B, D, F, 0.5 mm; C, 0.1 mm.

Material examined. Palm Beach, Queensland, 2 km off¬

shore, depth 5-15, NMV F91900, three short, infertile

stems on calcareous bryozoan.

Description. Hydrorhiza thick, tubular. Longest

stem 12 mm high, straight, hydrocladia opposite,

three widely separated pairs almost perpendicular

to axis of stem; stem inlernodes long, cylindrical,

smooth, nodes transverse, just above hydrocladia,

a subpposite pair ofhydrotheca between, halfway

along some intemodes. Hydrocladia inserted on

short, tubular apophyses narrowing distally but

without obvious distal node.

Hydrothecae paired, opposite, tubular, of same

diameter throughout length, supported on a short

conical section of intemode between pairs;

adcauline walls in contact or slightly separated on

intemode, adnate walls parallel to intemode,

straight, or slightly convex, free wall convex or

continuing bend of adnate wall, abcauline wall

sinuous, basally convex, concave behind margin;

floor of hydrotheca convex, thickened, a hydro¬

pore at base of abcauline wall. Margin faintly sin¬

uous (lateral view), tilted upwards to intemode,

rim slightly but sharply outrolled, many margins

346

J. E. WATSON

regenerated by growth of new hydrotheea from

deep within old, abcauline wall of new

hydrotheca free, visible, adcauline wall attached

to wall of primary hydrotheca. Hydranth with

about 12 long tentacles. Perisarc of stem, hydro-

cladia and hydrothecae thick and smooth, thinner

on hydrothecal margin. Colour, pale yellow.

Distribution. Type locality. New South Wales

(Allman, 1888). Tropical northern Australia and

New South Wales. Previously recorded from

southern Queensland as S. patiilum by Pennycuik

(1959).

Remarks. The distinction between Synthecium

campylocarpum and S. patulum (Busk, 1852) vvas

discussed by Watson (2000). The present material

is assigned to S. campylocarpum because of the

shorter hydrothecae, less pronounced marginal

sinuosity of the hydrotheca and yellow colour of

the colony.

Hincksella Billard, 1918

Hincksella cylindrica (Bale)

Figure 5D

Serliilarclla cylindrica Bale, 1888; 765.

Synthecium cylindricum. —Ritchie, 1911: 847.—

Stechow. 1923: 150.—Fraser. 1948:234.

Hincksella cylindrica. —Blackburn, 1937: 173.—

Pennycuik. 1959: 189.—Vervoort. 1959: 245.—

Vervoort.1968: 101.—Millard, 1975; 232.—Watson,

1979; 234.—Calder, 1993; 68.—Veivoort, 1993:

193.—Watson and Meinnes. 1999: 108.

Material e.xamined. Palm Beach, Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91901, several infertile

stems detached from substrate, malinol mounted

microslide preparation.

Description. Stems fragile, to 4 mm high, proxi¬

mal part of stem athecate, upper stem with up to 6

hydrothecae, stem intemodes wide, cylindrical,

curved, smooth, an indistinct slightly oblique

node below some hydrothecae. Hydrothecae sub¬

opposite to alternate, cylindrical, adnate

adcauline wall convex, free adcauline wall about

one quarter length of adnate wall, continuing

curve of fused wall, abcauline wall concave; floor

of hydrotheca transverse to intemode, narrow.

Margin circular, rim weakly outrolled. Perisarc of

lower stem thick, thinning markedly on hydrothe-

eae, most of which have collapsed in mounting.

Colour, white.

Distribution. Type locality. Port Jackson, New

South Wales (Bale, 1888). Southern Australia,

Pacific coast of North America, Caribbean.

Previously recorded from southern Queensland.

Remarks. Hincksella cylindrica is an uncommon

species, usually found as single stems or colonies

consisting of only a few stems.

Halopterididae Millard, 1962

Antennella Allman, 1877

Antennella secundaria (Gmelin, 1791)

Figure 5E-F

Sertidaria secundaria GmcVm. 1791: 3856.

Aglaophenia .secundaria. —Lamouroiix, 1824: 19.

Antennella .ver(/;i(/on(7.—Pennycuik. 1959: 176.—

Watson. 1973: 183.—Millard, 1975: 332.—Ryland and

Gibbons, 1991: 525.—Ramil and Vervoort, 1992: 143.

—Mcdel and Vervoort, 1995; 35.—Watson. 1996;

78.—Schuchert, 1997: 14.—Calder. 1997: 29 (full syn¬

onymy).—Watson. 1997: 522.—Watson, 2()0(): 45.

Anienella secundaria. —Stechow and Muller, 1923:

473.

Material examined. Palm Beach. Queensland, 2 km off¬

shore. depth 5-15 m, NMV F91290, fertile colonics on

barnacles and solitary ascidians in crevices, alcohol

preserved.

Description. Stems (hydrocladia) simple,

unbranched, to 6 mm high, arising from a tubular

stolon; proximal part of hydrocladium with one to

three athecate intemodes, nodes transverse, each

intemode bearing one or two nematothecae, dis-

talmost node strongly oblique. Intemodes there¬

after alternately thecate and athecate, proximal

node of thccate intemode strongly oblique, pass¬

ing beneath hydrotheca, distal node transverse

below hydrothecal margin.

Hydrotheca occupying almost entire intemode,

facing forward, deep cup-shaped, walls almost

parallel, free adcauline wall straight to slightly

concave, adnate part convex, curving back to

small hydropore at base of abcauline wall; margin

circular, slightly sinuous. Nematothecae bithala-

mic, small, 2 on athecate intemode, basal cham¬

ber stout, cup small, foreshortened on adcauline

side; mesial inferior stout, adcauline side of cup

foreshortened, just reaching base of hydrotheca,

lateral nematotheca borne on a long, slender pedi¬

cel where adcauline wall of hydrotheca becomes

free, cup broad and shallow, excavated on

adcauline side, not quite reaching hydrothecal

margin; a small mesial superior nematotheca

inserted beneath hydrotheea. Female gonothecac

obovoid, somewhat flattened, facing forward,

borne on a short, thick pedicel beside mesial infe¬

rior nematotheca; 2 large nematothecae at base;

basal chamber of nematotheca long, cup wide,

shallow, excavated on side facing gonolheca. Ori¬

fice of gonolheca distal, subcircular, closed by a

HYDROIDS FROM SOUTHERN QUEENSLAND

347

thin operculum, gonotheca containing a single

planula larva almost filling gonotheca at maturity.

Colour, golden yellow, planula shining white.

Distribution. A cosmopolitan species with tem¬

perate and tropical distribution in Australian

waters.

Remarks. A tew male gonothecae are present on

the stems. The species conforms in all respects

with descriptions of A. .^ecioularia from other

Australian localities (Watson, 2000).

Aelaopheniidae Marktanner-Turneretscher,

1890

Lytoairpia Kirchenpauer, 1872

Lytocarpia hrevirostris (Busk)

Figure 6A-D

Phimuiaria hrevirostris Busk, 1852; 397.

Aglaophenia hrevirostris. —Bale, 1913: 135.

Thecocarpiis hrevirostris. —Billard, 1913; 89.—

Briggs, 1918 : 34, 45.—Stechow, 1919: 137.—Bedot,

1922; 157.—Jarvis. 1922; 350.—Pcnnycuik, 1959:

187.—Vasseur, 1974: 158.—Millard, 1975:454.

Lvtocarpia hrevirostris. —Stcchow, 1922: 151.—Ste¬

chow, 1923: 245.—Rylaiid and Gibbons. 1991: 545.

Material examined Palm Beach. Queensland, 2 km off¬

shore, depth 5-15 m, NMV F9I291, infertile colony on

polychaete tube, alcohol preserved.

Description. I lydrorhiza a stout tube from which

stems arise at interv'als. Stems short, pinnate, to

10-15 mm high, proximal part of stem ahydro-

cladiate, some with a supplementary fascicular

tube running part way up stem. Stem internodes

short, nodes poorly defined, transverse to slightly

oblique; hydrocladia alternate, about 2 mm long,

1 per stem inteniode. inserted on a short apophy¬

sis on front of stem; a small open pore (mamelon)

on base of apophysis.

llydrocladial intemodes long, base of each

strongly displaced with respect to next one, node

present, indistinct or absent at point of displace¬

ment, internode with I complete septum beneath

posterior quarter of hydrotheca and another, par¬

tial, below saddle in hydrotheca. Hydrotheca

elongate, parallel with internode, margin separ¬

ated from mesial nematotheca by deep saddle;

margin sloping forward with a long, sinuously

curved, blunt anterior cusp, followed by 3 pairs of

deeply incised, sharply pointed cusps, the third

pair usually obscured by lateral nematothccae;

base of hydrotheca with a short, fonvardly point¬

ing intrathecal septum. Mesial hydrothecal nema¬

totheca rather short, digitate, at an angle of about

50° to internode, apex about level with anterior

hydrothecal cusp, orifice small, shallowly

canaliculate; lateral nematotheca flask-shaped,

laying almost parallel to intemode, orifice small,

canaliculate down to intemode. Two cauline

nematothccae, one above and one below hydro-

cladial apophysis; same shape as laterals, but base

a little more inflated, orifice of upper facing away

from apophysis, orifice of lower facing up and

outwards. Colour, yellow.

Distribution. A common Indo-Pacific species pre¬

viously recorded from Queensland.

Remarks. The colony was found on the tube of a

womi of the genus Eunice. The tough, mucilagi¬

nous tube of this polychaete genus is a favoured

substrate of hydroids in tropical and temperate

waters (Watson, 2000).

Aglaophenia Lamouroux, 1812

Aglaophenia siniiosa Bale

Figure 6E-H

Aglaophenia sinuosa Bale, 1888: 790.—Whitelegge,

1889: 194.—Jaderholm, 1916: 19.—Watson, 1994; 67.

Material examined. Palm Beach. Queensland, 2 km oil-

shore, depth 5-15 m, NMV F91292, infertile colonies

of many stems on exposed rock surfaces, alcohol pre¬

served.

Description. Hydrorhizal stolons thick, reptant on

substrate; stems monosiphonic, up to 70 mm long,

internodes short, wide, nodes transverse to

slightly oblique, sloping in different directions, a

deep shoulder at node. Hydrocladia to 10 mm

long, alternate, one per cauline intemode, inserted

on a short apophysis. Hydrocladial intemodes

deep, nodes slightly oblique, distinct, a weak

intranodal septum sometimes below posterior of

hydrotheca.

Hydrotheca saccate, base convex, posterior

wall deeply indented, a thick septum passing

down into body from junction ofabcauline wall

with mesial nematotheca, edge of septum ragged

in anterior view. Margin sloping sharply down¬

wards to intemode; margin (lateral view) with a

thick rostnim overtopping median anterior

tongue-shaped cusp, flanked by 2 pairs of tongue¬

shaped cusps and a pair of sharp, upwardly

pointed cusps behind nematothccae. In anterior

view, first pair of cusps outwardly directed, sec¬

ond pair produced inward over hydrothecal mar¬

gin, third pair slightly outwardly directed; the

entire margin having a fluted appearance. Mesial

nematotheca short, free end blunt, reaching level

of intrathecal septum, margin widely canaliculate,

curving down to junction with hydrotheca.

348

J. E. WATSON

Figure 6. A-D, Lytocarpia brevirosiris (Busk, 1852). A, whole stem; B, hydrothecate hydrocladial internodes:

C, hydrotheca, anterior view; D, stem intemode with cauline nematothccac and mamelon. E-Il, Aglaophenia

sinuosa Bale, 1888. E, single stem from colony; F. hydrocladial hydrothecae; G. hydrotheca, anterior view; H, front

of stem showing oblique internodcs and cauline nematothecae (hydrocladia not shown). Scale bar; A, E, 20 mm;

B, C. D, F, G, I I, 0.3 mm.

Lateral nematothecae conical, widening from

base to margin, base parallel with internode, ori¬

fice circular, level with node, excavated down to

intemode. Three cauline nematothecae: i) on front

of internode at base of apophysis, facing inwards,

ii) between apophysis and proximal node, facing

outwards, iii) one behind apophysis; all nenia-

tothecae with 2 opposite orifices. Colour, dark

brown.

Distribution. Tasman Sea, New South Wales. Not

previously recorded from the Queensland coast;

possibly at the northern end of its range.

Remarks. Aglaophenia sinuosa is easily recognis¬

able among Australian aglaopheniid hydroids by

the sinuous bends at intervals along the stem,

each bend marking a reversal of the direction in

which the hydrocladia face so that succeeding

groups of hydrocladia alternately face frontwards

and backwards. The marginal hydrothecal cusps

are quite difficult to describe as their apparent

position and morphology changes according to

the angle of view; in lateral view the cusps may

appear as sharply pointed whereas in anterior

View some are tongue-shaped and inw'ardly and

outwardly fluted. There is some discrepancy

between the morphology of the hydrotheca of the

present material and Bale’s description and figure

of the species, his specimens having tw'o intrathe¬

cal septa and more upright lateral nematothecae.

This may be a rather variable species, but not

enough is yet known of its range of variation.

HYDROIDS FROM SOUTHERN QUEENSLAND

349

Macrorhynchia Kirchenpauer, 1872

Macrorhynchia philippina (Kirchenpauer)

Figure 7A, B

Macrorhvnchia philippina Kirchenpauer, 1872:

19 .—Stechow, 1923: 241.—Slechow and Muller, 1923:

475.—Stechow. 1924: 69.—Stechow, 1925: 258.—

Hirohito, 1983: 78.—Rees and Vervoorl, 1987: 177.—

Watson, 1996: 79.—Migotto, 1996: 40.—Watson.

1997: 538.—Calder, 1997: 62.—Watson. 2000: 67.

Aglaopbenia philippina. —Kirchenpauer, 1872: 29,

45.

Lytocarpus philippinus. —Kirkpatrick. 1890: 604.—

Bale. 1888: 786.—Billard. 1913: 78.—Bale, 1914a:

6.—Bale, 1915: 293.—Jiiderholm, 1916: 7.—Briggs

and Gardner, 1931: 193.—Millard, 1958:220.—Penny-

cuik. 1959: 186.—Millard and Bouillon, 1973: 93.—

Millard, 1975:449.

Material and record. Palm Beach. Queensland. 2 km

offshore, depth 5-15 m, NMV F91293, many infertile

colonies on rock faces, alcohol preserved.

Description. Hydrorhiza reptant on substrate,

stems plumose, to 30 mm high, fascicled, sub-

pinnately branched, stem internodes indistinct.

Hydrocladia frontal on branch, alternate, hydro-

cladial intemodes short, deep, nodes slightly

oblique; two partial intranodal septa, one passing

down from base of hydrotheca, the other From

base of lateral nematotheca.

Hydrotheca saccate, adcauline wall flat or

slightly convex in contaet with intcmode, free

part attached to lateral nematotheca, a deep prox¬

imal indentation in base of adcauline wall,

abcauline sinuate, rising abruptly behind margin,

a deep V-shaped septum penetrating hydrotheca

between mesial nematotheca and margin. Margin

circular with a pair of opposite, low' lobes. Mesial

nematotheca erect, tubular, parallel with

hydrothecal abcauline wall, reaching to or beyond

hydrothecal margin, tapering a little distally, ter¬

minal orifice circular, a little excavated on

adcauline side, a secondary orifice above junction

with hydrotheca and another, internal, connecting

with hydrotheca. Lateral nematotheca long, tubu¬

lar, inclined forward beyond hydrothecal margin,

orifice circular or slightly excavated. A cauline

nematotheca on branch at base of hydrocladium.

short, body inflated, orifice circular, on a short

neck. Colour, colonies glowing white in situ,

stems of preserved material pale brown,

hydrocladia greyish-white.

Distribution. Macrorhynchia philippina is a very

common circumglobal tropical species. It occurs

throughout the Australian tropics (Watson 2000).

Remarks. None of the numerous colonies of

M. philippina observed off Palm Beach were

more than a few centimetres high, growing close

to the rocky substrate in exposed situations.

Although the colonies were very small for this

species they showed no evidence of grazing by

fish, thus suggesting that dwarfism may be due to

the species being close to the southern end of its

temperature range.

Campanulariidae Johnston, 1836

Ohelia Peron and Lesueur, 1810

Obelia angulosa Bale

Figure 1C

Obelia angulosa Bale, 1888: 752.—Whitelegge,

1889: 195.—Stechow, 1914: 129.—Stechow, 1919:

49.—Vannucci-Mendes, 1951b: 115.—Blanco, 1968:

210.—Stranks, 1993: 4.—Blanco. 1994: 193.—Mig¬

otto, 1996: 123.

Material e.xamined. Palm Beach. Queensland, 2 km off¬

shore, depth 5-15 m, NMV F91294, many infertile

colonies on barnacles, sponges and ascidians. alcohol

preserved.

Description. Hydrorhiza tubular, reptant on sub¬

strate. Stems to 6 mm high, monosiphonic, low'er

stem smooth, unbranched, a series of annulations

above base, upper stem region slightly sympodial,

simple, dichotomously or trichotomously

branched, intemodes moderately long, tubular,

straight, widening a little distally, nodes oblique,

faint to absent, proximal part of intemode deeply

annulated. Hydrothecal pedicels usually alternate,

given off at an acute angle distal on internode,

pedicels variable in length, annulated throughout

or with a central smooth section. Hydrotheca

gracefully campanulate, diaphragm transverse to

hydrothecal axis, margin faintly everted.

Hydranth robust, number of tentacles could not be

counted. Perisarc smooth throughout, fairly thick

on intemodes, pedicels, and around upper body of

hydrotheca. Colour, lower stems shining brown,

upper stems fading to white.

Distribution. Not previously recorded from

Queensland; known from New South Wales

(Bale, 1888) and Victorian waters (Watson,

unpubl.) and South America.

Remarks. This very delicate species is easily dis¬

tinguished from its more common southern

Australian congener, Obelia dicbotoma (Lin¬

naeus, 1758), by its smaller size and almost

invariably transverse diaphragm. The coenosarc

of the hydrorhiza and stems and the tentacles

of the specimens are packed with zooxanthellae.

350 J. E. WATSON

Figure 7. A, B, Macrorhynchiaphilippina (Kirchenpauer. 1872). A. whole stem; B, hydrocladial hydrothecac (after

Watson, 2000). C, Obelia angidosa Bale, 1888. Stem intemodes and hydrotheca. Scale bar: A, 50 mm; B, 0.3 mm;

C, 0.5 mm.

O. angidosa is the most abundant species in the

collection although it is probably seasonal and

likely to be absent at other times of year.

Acknowledgements

1 sincerely thank Dr Rex Neale of Palm Beach,

Queensland, and his family for providing a boat,

diving companionship and ho.spitality during field

work on the Gold Coast.

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Memoirs of the Museum of Victoria 59(2): 355-392 (2002)

A REVISION OF THE AUSTRALIAN GENUS UMBILIA (GASTROPODA: CYPRAEIDAE)

Thomas A. Darragh

Museum Victoria, PO Box 666E, Melbourne, Vic. 3001, Australia

(tdarragh(3)museum. vic.gov.au)

Abstract

Darragh. T.A., 2002. A revision of the Australian genus Umbilia (Gastropoda: Cypraeidae).

Memoirs of Museum Vidoria 59(2): 355-392.

Umbilia, an endemic Australian genus of eool-water cowry, comprises 11 species ranging

in age from Late Oligocene to Recent, of which four species are known only in the living

fauna. Fossil species occur in the Eucla. St Vincent, Muiray. Otway, Bass and Gippsland

Basins, and living species range from We,stem Au.stralia to central Queensland. Species of the

genus probably have no free-swimming larval stage, so that there is considerable morpho¬

logical variability which has led to the creation of many synonyms. Two subgenera are recog¬

nised here. Umbilici (Umbilia). with ten species, U. prosila sp. nov., U. lepiorhyncha.

U. pelilirosiris sp. nov., U. planrliyuclia. U. cmgustior. U. eximia (= brevis, maccoyi, frank-

sloneusis. monlismarlhae and sphaerodoma), U. hesilala (= beddomei, eera, liowelli and

laleU). U. siplionaUi (= breviplicata), U. armeniaca and U. capricornica, and U. (Pallio-

cvpraea) with one species, U.(P.) gaslroplax. A possible ancestor of Umbilia, Palaeocypraeal

eripnides, from the Upper Paleocene-Lower Eocene, of the Chatham Is, New Zealand, is

newly described. Rhynchocypraea Cossmann, 1898 (type species Cypraea lepiorhyncha

McCoy) is synonymised vs ilh Umbilia.

Introduction

Umhilla is an endemic Australian genus of

cowry. Living species are found in deep water

from the southern coast of Western Australia and

along the east coast to the Capricorn Channel area

of Queensland. Fossil species arc known from the

Eucla, St Vincent, Murray, Otway, Bass, and

Gippsland Basins in rocks ranging in age from

Late Oligocene to Late Pliocene. In general,

fossil specimens are not common except in the

Middle Miocene Fyansford and Gellibrand For¬

mations of the Otway Basin. One of the extant

species (U. hesituta) also has fossil records

(Miocene-Pliocenc).

Umhilla is a member of an endemic group of

Australian cowries that includes Notocypraea,

Austrocypraea and ZoUa (Wilson, 1985). The

present paper reports results of part of a wider,

comprehensive study of all the Australian fossil

cowries.

Previous work. The first fossil species of

Umhilla to be described, Cypraea eximia G. B.

Sowerby I, 1845, was based on a specimen col¬

lected by P. E. Strzelecki allegedly from Tas¬

mania. McCoy (1867a, 1875, 1876) described

another species, Cypraea (Aricia) plalyrhyncha

from Torquay as well as illustrating Victorian

examples of Sowerby’s species and describing a

new variety of it, Cypraea (Aricia) eximia var.

hrevis. He also described Cypraea gaslroplax

(McCoy, 1867a), now included in the subgenus

Umhilia (Palliocypraea) and Cypraea (Luponia)

lepiorhyncha (McCoy, 1877) herein regarded as

an Umhilia. Tate (1890) listed previously

described species, recorded new occurrences and

described two new taxa, Cypraea sphaerodoma

and C. amygdalina, the latter renamed C. lalei by

Cossmann (1903) owing to preoccupation of

Tate’s original name. Tate also noted a general

resemblance of Cypraea eximia to the living

C. timhilicala G. B. Sowerby I (= hesilala

Iredale), type species of Umhilia Jousseaume,

1884.

Pritchard (1896) described a new variety of

McCoy's species, Cypraea plalyrhyncha var.

anguslior from northern Tasmania, that is here

elevated to specific rank. Harris (1897) placed C.

eximia and C. sphaerodoma in Umhilia, which he

regarded as a subgenus of Cypraea. Cossmann

(1898) erected Rhynchocypraea foi' Cypraea lep¬

iorhyncha. This is synonymised with Umhilia

(Umhilia) in this paper. In 1906 Cossmann

erected Palliocypraea as a siibgenus of his genus

Rhynchocypraea, with type species Cypraea

355

356

THOMAS A. DARRAGH

gastroplax McCoy. Chapman (1922) described

Cypraea siphonata from near Waikerie, South

Australia, a species now included in Umbilia. In

1926, Schilder named Gisortia breviplicata based

on Tate’s (1890) description of C sphaerodoma

var. In 1932, Schilder listed all previously

described taxa under the genus Umbilia and

erected U. (U.) eximia maccoyi based on McCoy's

figures (McCoy, 1876) of Cypraea eximia.

In 1935, Schilder revised all the Tertiary

cowries from Australia, describing many new

taxa based on material (much with poor locality

data) in European collections. U. ( U.) brevis mon-

tismarthae and U. (U.) brevisfraiikslonensis were

the new taxa of Umbilia so erected. He had little

idea of the variability of the taxa owing to the rel¬

atively small number of specimens available to

him (18 specimens of Umbilia) and he had no

idea of their stratigraphic positions. Schilder

regarded both Rbynchocypraea and Pallio-

cypraea as subgenera of Umbilia. maintaining

Rhynchocyprea as a montotypic subgenus and

assigning Gisortia breviplicata and Cypraea

siphonata to Palliocypraea. The last fossil

species to be erected, U. cera, was described by

Cotton (1947) from the Dry Creek Sands of the

St. Vincent Basin, South Australia.

There are four known living species of

Umbilia-. U. (U.) armeniaca (Verco, 1912) from

southern Western Australia and South Australia;

U. {U.)hesitata (Iredale, 1916) widely distributed

from South Australia to southern Queensland; and

U. (U.) capricornica Lorenz, 1989 and U. (U.)

petilirostris sp. nov. from south-central Queens¬

land. Of these, only U. (U.) hesitata is known

from the fossil record.

Generic status. There has been considerable

debate as to the worth of many of the genera

within the family that were erected on the basis of

shell morphology, particularly as anatomical

studies of several species show that the anatom¬

ical features of many of these “genera” are very

similar to one another and do not support their

separation on shell characters (Kay, 1960). This

applies particularly to the Indo-Pacific tropical

cowry' genera. In the case of Umbilia. however,

there does seem to be sufficient grounds for

recognising the group, either at generic or sub¬

generic level. These grounds are essentially its

unique geological history, its geographic and

stratigraphic isolation from other cowry groups,

and its paucispiral protoconch. Most of the tropi¬

cal cowries have pelagic larval stages, which

accounts for their widespread distribution. These

tropical species have multispiral protoconchs

(Ranson, 1967).

Molluscs with paucispiral protoconchs usually

do not have a pelagic larval stage but hatch

directly from the egg as crawling juveniles.

Though nothing is known yet of the mode of

reproduction of species of Umbilia, the distribu¬

tion and fossil history of the genus also suggest

that there was no pelagic larval stage. Ranson

(1967) illustrated the protoconchs of specimens

of Umbilia hesitata and showed these as being

paucispiral. Sections made by me of two fossil

species, U. (U.) (Pritchard) and U. (U.)

eximia (Sowerby) (Figs lA-B), show that these

also have paucispiral protoconchs, so it seems

likely that all species of Umbilia. living and

extinct, feature direct development rather than

pelagic larval stages. Wilson (1985) has shown

that species of the other endemic genera, Zoila.

Austrocypraea and Notocypraea. feature direct

development.

Direct development in molluscs leads to exten¬

sive infraspecific variation (Wilson, 1993).

Species of Umbilia. both living and fossils show

considerable infraspecillc variation in shell mor¬

phology and colour. This variation has led

inevitably to the erection of a considerable num¬

ber of specific and subspecific names for both

colour and morphological varieties which the

present work seeks to reetify.

Fossil history. The earliest fossil records of

Umbilia are from the Late Oligocene, by which

time it seems that direct development in the genus

had been established. It is suggested here that

Umbilia may have been derived from an ancestor

such as Palaeocypraedl eripnides sp. nov.. from

the Paleocene of New Zealand, which has a mul¬

tispiral protoconch suggesting a pelagic larval

stage in its development. Another possible ances¬

tor occurs in the Eocene of Westeni Australia,

Cypraeorbis sp., which could have given rise to

the Late Oligocene U. prosila sp. nov. The fossil

record of Umbilia shows two possible species

lineages, both beginning in the Late Qligoccne

and continuing through to the Recent: U. platy-

rhyncha - U. angustior - U. eximia leading to

the living U. hesitata and U. prosila sp. nov. -

U. leptorhyncha leading to the the living

U. petilirostris sp. nov. There are also two other

species in the Middle Miocene which on present

knowledge do not seem closely related to these

lineages and have no known descendants. One of

these is sufficiently distinct to justify subgeneric

status, Palliocypraea. It is possible that the genus

Umbilia as recognised here is polyphyletic, but

until mollecular studies have been undertaken on

living representatives, the question must remain

unresolved.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

357

Rhynchocypraea Cossniann, 1898 (type species

Cypraea leplorhyncha McCoy) is synonymised

with Umhilia (Umhilia) in this paper.

Species are described here in groups according

to their affinity with one another, and within these

groups in stratigraphic order.

Terminology. Measurements in millimetres are

given as follows: L, total length of shell; W, width

of shell; H, height of shell. Tooth counts are cited

as LT, labial teeth and CT, columellar teeth. The

terms left and right refer to the animal’s true left

and right sides respectively.

All material used in this study, unless otherwise

stated, is held in the collections of Museum

Victoria, registration numbers with prefixes

P (Invertebrate Palaeontology collection) and

F (living mollusca collection). Localities are cited

where possible using the Museum Victoria fossil

locality register with prefix PL. The full locality

data for these numbers has been published in

Darragh (1989, 1991) and Beu and Darragh

(2001). Museum acronyms arc as follows:

BMNH, Natural History Museum, London; TM,

Type mollusca collection. New Zealand Institute

of Geological and Nuclear Sciences, Wellington;

SAM T, Tate Collection, South Australian

Museum, Adelaide; AM, Australian Museum,

Sydney, New South Wales.

All specimens figured were coated with ammo¬

nium chloride for photography, unless stated

otherwise.

Class Gastropoda

Cypraeidae

Palaeocypraea SchWitr, 1928

Fa/ac’ociprat’« Schilder, 1928: 17.

Type species. Cypraeacites spircUa Schlotheim,

1820 by original designation (Danian, Denmark).

“IPalaeocypraea eripnides sp. nov.

Figure 2A-D

Cypraea (Zoih) n. sp. Beu and Maxwell, 1990; 96,

pi. 4j, k.

Type material. Holotype TM8I24, paralype TM8125,

P.A. Ma.xwell, 1977.

Type locality. CH/f471, Wave-cut platform below ‘The

Bluff homeslead. Pitt I., Chatham Is, New Zealand. Pitt

1. 713236. Red Bluff Tuff, Late Teurian-late

Waipawan, Late Paleocenc-Early Eocene.

Occurrence. Type locality only (2 specimens).

Description. Shell of large size for genus, smooth,

elongate, pyriform, tapering gently anteriorly.

Protoconch multispiral, not distinguished from

teleoconch whorls. Spire conical, projecting,

covered with thin glaze.

Posterior canal short, inner lip sharp, curved

towards and attached to spire. Anterior canal

short. Aperture slightly sinuous; outer lip with

12-14 teeth present on anterior two-thirds to half

of lip; columella with 8-11 teeth present on ante¬

rior two-thirds to half. No terminal ridge. Fossula

wide and shallow.

Dimensions L W H LT CT

Holotype TM8124 44 22 18 12 8

ParatypeTM8125 51 27 27 14 11

Time range. Late Teurian-late Waipawan, Late

Paleocene-Early Eocene.

Remarks. Beu and Maxwell (1990) placed this

taxon in Zoi/a; however, the fossula is wide and

shallow, quite different from the well developed

spoon shaped fossula which in all species of Zoila

is bounded anteriorly by a ridge. Of the Southern

Hemisphere cowries, it comes closest to Umhilia,

but possesses a conical spire and multispiral pro-

toeonch in contrast to the umbilicate spire and

paucispiral protoconch of Umhilia. However, it

may well be ancestral to it as discussed below.

This species is referred to Palaeocypraea with

considerable doubt, since the .shell is very elon¬

gate when compared with other taxa referred to

the genus and the fossula is relatively poorly

developed, whereas in Palaeocypraea spirata the

fossula is broad and concave with a distinct notch

on the smooth inner border. Owing to the preser¬

vation of the specimens, it has not been possible

to determine whether this species has fenestrate

sculpture on the early whorls of the spire as in

other species of the genus.

This is one of the oldest cowries known from

the Southern Hemisphere and is included here as

a possible ancestor of the southern Australian

Umhilia, the earliest species of which is known

from the Late Oligocene.

Umhilia Jousseaume, 1884

Umhilia Jousseaume. 1884a: 414.—Jousseaume.

1884b: 90; Schilder, 1935: 342.—Schilder, 1939:

186.—Wenz, 1941: 995.

Type species. Cypraea timhilicala Sowerby, 1825

(non Dillwyn, 1823 = C. hesilala Iredale, 1916)

by monotypy. Recent, south-eastern Australia.

Diagnosis. Shell highly glazed, of medium to

large size for the family, pyriform with umbilicate

358

THOMAS A. DARRAGH

spire. Anteriorly and posteriorly rostrate, the ante¬

rior rostrum generally larger and bearing 2 dorsal

tubercles. Rostra supported by Ranges that may

be weakly or strongly developed or extend around

the base of the last whorl. Aperture sinuous with

weakly or strongly developed teeth. Fossula

weakly developed or absent.

Remarks. The monotypic subgenus Palliocypraea

from the Miocene is characterised by the devel¬

opment of a thin, wide, flat flange extending

around the base. Species of Umbilia lacking this

feature are all placed in the subgenus Umbilia

sensu stricto.

Umbilia (Umbilia) Jousseaume, 1884

Umbilia Jousseaume, 1884a: 414.—Jousseaume,

1884b: 90.—Schilder, 1926: 378.—Thiele, 1929:

275.—Schilder and Schilder, 1939: 145.

Cypraca (Umbilia). —Harris, 1897: 209.—

Cossmann, 1903: 160.—Wilson, 1993: 191.

Rhynchocypraea Cossmann, 1898: 17, type species

(original designation) C\'praea (Luponia) leptorhvncha

McCoy, 1877.—Wenz, 1941: 994.

Rhynchocypruea,—Cossmann, 1903: 174. in part;

not of Cossmann. 1898.

Umbilia (Umbilia). —Schilder, 1935: 342.—

Schilder. 1939: 186.—Wenz, 1941: 995.

Umbilia (Rhvnchocvpraea). —Schilder, 1935: 342.—

Schilder. 1939: 186. ‘

Diagnosis. Shell solid, highly glazed, of medium

to large size for the family, elongate to sub-

globosely pyriform, generally somewhat more

ventricose posteriorly. Anteriorly and posteriorly

rostrate; anterior rostnim elongate to very elon¬

gate in some species, flattened ventrally, bearing

2 weakly to well developed dorsal tubercles

separated by sinuous groove. Posterior rostnim

deeply notched. Rostra supported by triangular

extensions of base (extensions poorly developed

on species with short rostra). Anterior and pos¬

terior canals incised into anterior and posterior

rostra. Dorsal surface of shell generally smooth,

somewhat granular on some specimens.

Spire depressed below last whorl, early teleo-

conch and protoconch covered with thin glaze, in

most species spire whorls and protoconch visible

on slightly decorticated specimens. Protoconch of

3 or 4 smooth dome-shaped whorls, tilted at about

10° to axis of teleoconch. Tcleoconch of 3-4

whorls.

Aperture sinuous. Labial teeth well to poorly

developed, on some specimens covering whole

length of lip, on others fading on posterior half;

columella teeth variable in expression, on one

species developed into a series of prominent

elongate ridges separated by deep grooves; teeth

and ridges variable in number, on some species

fading on posterior half of columella. Terminal

ridges of aperture, variable in strength and num¬

ber, from one to three. Fossula on most species

weakly developed, absent on some specimens.

Time range. Janjukian (Late Oligocene) to

Recent.

Distribution. Southern Western Australia

(Pliocene, Recent); South Australia (Middle

Miocene-Late Miocene. Recent); Victoria (Late

Oligocene-Late Miocene, Recent); Tasmania

(Early Miocene, Late Pliocene, Recent); New

South Wales (Recent); southern Queensland

(Recent).

Remarks. The well developed anterior rostrum is

the most characteristic feature of this subgenus.

The posterior rostrum is generally much less

developed and varies between species from being

greatly extended to a short stump.

Cossmann erected Rhynchocypraea in 1898,

with type species, Cypraea (Luponia) leptorhyn-

cha McCoy for Australian fossil species having

rostral extensions of the aperture. In 1903, he

either overlooked or ignored his previous desig¬

nation and cited C. loxorhyncba Tate (an error for

to.xorhyncba) as type species instead. I le placed

C leptorhyncha in his newly erected Austro-

cypraea (type species C contusa McCoy).

C. toxorhyncha is a junior synonym of Zoila

platypyga (McCoy). Vredenberg (1927) accepted

Cossmann’s 1903 interpretation, unaware of

Cossmann's original designation of a type species

for Rhynchocypraea but synonymised both

Umbilia and Rhynchocypraea with Gisortia from

the Paleogene of Europe.

In 1930, Schilder regarded C. leptorhyncha as

an Umbilia. stating that numerous identical spec¬

imens seemed to exclude their being juveniles of

U. eximia and that the species connected the ros¬

trate Umbilia with Austrocypraeapyrulata (Tate).

In 1935, however. Schilder maintained Rhvn-

chocypraea as a siibgenus ol'Umbilia. stating that

it was related to Umbilia (Palliocypraea) and also

that it was closely related to some species of his

new genus, Notoluponia. (presumably pyrulata).

Schilder separated Rhynchocypraea from

Umbilia by the presence of a shallow rather than

deeply notched posterior outlet and by the fact

that it was not rostrate. In 1939, Schilder stated

that Rhynchocypraea was more pear shaped,

more gibbous, with a more slender anterior end.

wider mouth and narrower siphonal canal. These

dilTerences are quite insignificant, leptorhyncha

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

359

is just as anteriorly rostrate as hesitata, if not

more so and the posterior outlet and width of

aperture and canals vary in degree of develop¬

ment from species to species. As 1 can find no

consistent character which can Justify separation

of leptorhyncha from hesitata at the generic level,

1 regard Rhynchocypraea as a synonym of

Uinhilia (Umbilia).

Wilson (1993) suggested that Umbilia (U.) hes¬

itata fed on bryozoans. Bryozoans are a common

element in the fossil assemblages of the fonna-

tions in which fossil species of Umbilia have been

found. Lorenz (1988) reported finding the

remains of sponges in the stomachs of specimens

of U. (U.) capricornica and Wilson (1998)

reported divers taking U. (U.) armeniaca crawl¬

ing on sponges. Species of Zoila are known to

feed on sponges (Wilson and McComb, 1967).

The anatomy of only one living species of

Umbilia is known. Lorenz (1988) briefly

described and illustrated the broad features of U.

capricornica, comparing it to Cypraeovtila fus-

corubra and Cypraea cruikshanki from southern

Africa. .ludging from Lorenz’s figures the

anatomy of U. capricornica is somewhat similar

to that of Zoila friendii described and illustrated

by Wilson and McComb (1967), particularly in

the pre.sence of a bursa copulatrix in the female

genital system.

The origin of the genus is not known. There is

nothing quite like it known from the Southern

Hemisphere, e.xcept possibly 'IPalaeocypraea

eripnides sp. nov.. Late Paleocene-Early Eocene,

Chatham Islands, New Zealand and an unde¬

scribed species of Cypraeorbis from the Late

Eocene of Western Australia. The New Zealand

species seems to have a multispiral protoconch, as

in other species of Palaeocypraea. suggesting a

planktonic larval stage, whereas species of

Umbilia have paucispiral protoconchs more

consistant with direct development. If Umbilia

is descended from this or some similar taxon,

there is adequate time between the Paleocene

and Late Oligocene for a switch to direct devel¬

opment to have taken place. Other changes

would involve an elongation of the anterior

canal. Otherwise the overall morphology of

’IPalaeocypraea eripnides sp. nov. is not dissimi¬

lar from that of small specimens of the Late

Oligocene, Umbilia plalyrhyncba. and from spec¬

imens of the Early Miocene, U. angusiior. In con¬

trast to this suggestion, the Western Australian

species of Cypraeorbis is closer to the Late

Oligocene U. prosila sp. nov. in overall shell

shape but has a much shorter anterior canal and

deeper fossula.

Umbilia {Umbilia) platyrhyncha (McCoy, 1876)

Figures 2E-F, H-K; 3A-F

Cypraea (Aricia) platyrhyncha McCoy, 1876: 40: 30,

figs 2-2c.

Umhilia (Umbilia) platvrhviicha. —Schilder, 1935:

343.

Type and figured material. Hololype P12138.

Figured specimens. P59I69, Richard Daintree, Geo¬

logical Survey of Victoria. Jul-Aug 1861. P302697,

C.W. Mallett, 15 Oct 1972. P302784, G.B. Pritchard

Collection.

Type locality. Geological Survey of Victoria locality

Ad24 (PL3024 Cliff section opposite Bird Rock, below

Bird Rock cap, Torquay, Victoria, Torquay 642518).

Jan Juc Fonnation, Late Oligocene, Janjukian.

Occurrence and material. Type locality only (17

specimens).

Description. Shell of medium size for genus, pyri-

fonn, maximum cotivexity close to posterior,

tapering anteriorly and produced into broad, flat¬

tened rostrum with 2 weak dorsal tubercles pre¬

sent on some specimens. Transverse section of

last whorl subcircular. Spire barely if at all

visible, on most specimens covered with thick

glaze. Protoconch of 4 smooth dome-shaped

whorls, tilted at about 10° to axis of teleoconch

whorls.

Posterior canal very short, twisted dorsally

toward spire; margins thickened, outer margin

more so and produced beyond inner margin.

Anterior canal elongate, deeply incised into ros¬

trum. Aperture only very slightly sinuous; outer

lip with 15-20 teeth present on about half lip

to midpoint of aperture. Columella with about

12 weak teeth, fading in strength posteriorly;

slight trace of ridge extending from posterior

teeth along columella to left side of posterior

canal. Temiinal ridge absent. Fossula broad,

very gently concave, merging imperceptibly

into columella margin. Dorsal surface smooth,

shining.

Dimensions

Molotype PI2138

Figured specimen

P302697

Figured specimen

P302784

very

faint

Figured specimen

12+

20+

P59I69

Time range. Janjukian, Late Oligocene.

Remarks. This is the one of the two oldest known

species of the genus recorded, and the rarest of

the fossil species. Most specimens are slightly

decorticated, slightly distorted, or broken. The

360

THOMAS A. DARRAGH

species is characterised by weakly developed

teeth, an elongate shape, lack of an umbilicus,

concealed spire, weakly developed dorsal tuber¬

cles and massive flat anterior rostrum and heavy

posterior canal.

This species bears some resemblance to

7Palaeocypraea eripnides sp. nov., Pitt I., New

Zealand (Fig. 2A-D). but it is much more mas¬

sive. about twice the size, has a relatively longer

anterior rostrum and a paucispiral rather than a

multispiral protoconch. P. eripnides has slightly

stronger developed teeth that e,xtend almost to the

end of the anterior canal. The fossulae of both are

similar. P. eripnides has a prominent conical exert

spire, whereas the spire in Umhilia (U.)

platyrhyncha is depressed and covered with a

thick callus deposit.

Unibilia {Umhilia) anfiustior (Pritchard, 1896)

Figures lA; 2G; 3G-1; 4A-I

Cvpraea plalvrhynclui angnstior Pritchard. 1896:

107; 4, figs 8, 9;

Umhilia (Umhilia) plalvrhvncha angnstior .—

Schilder, 1935; 343.

Umhilia sp.—Burgess, 1989; II, fig D.—Lorenz,

1989: fig. 5.

Type and figured material. Holotypc P267I, E.D.

Atkinson Collection.

Figured specimens. P302785, F.A. Cudmore Collec¬

tion. P302688, T.A. Darragh, 15 Nov 1984. P302703.

T.A. Darragh 30 Nov 1972.

Type locality. Table Cape. The matrix indicates that the

holotypc came from the lower bed, i.e. PL3028 Lower

bed in clilT between Fossil BlulT and 1.5 km NW

towards Table Cape, Wynyard. Tasmania. Table Cape

930630. Freestone Cove Sandstone. Early Miocene,

early Longfordian.

Occurrence and material. Freestone Cove Sandstone:

PL3028 Lower bed. Table Cape (25 specimens). Fossil

BlulT Sandstone: PL3029 Upper bed. Table Cape (5).

Puebla Formation; PL3032 Jan Juc Beach (2). Fishing

Point Marl: PL3037 Camping Reserve, Hordern Vale

(2); “Picnic bed”. Red Hill district. Hordern Vale (2);

Fischers Point, Hordern Vale (2); Cutting on road to

Hordern Vale camping reserve (I ).Gellibrand Fonna-

tion: PL3033 Birregurra (2).

Description. Shell of small size for genus, pyri¬

form, maximum convexity close to posterior,

tapering rapidly to anterior, produced into a

rapidly narrowing rostrum bearing 2 low tuber¬

cles. Transverse section of last whorl subcircular.

Spire covered with thick glaze, slight trace of

umbilicus on some specimens. Posterior canal rel¬

atively short, twisted back dorsally towards spire,

margins produced, thin. Anterior canal long,

deeply immersed in rostrum.

Aperture somewhat sinuous, outer lip with

8-22 teeth, on most specimens fading about mid¬

point of aperture, inner lip with 8-20 teeth on

most specimens, fading about midpoint of

aperture; slight trace of ridge extending from

teeth to side of posterior canal margin. Tenninal

ridge absent. Fossula narrow, flattened, merging

imperceptibly into columella. Dorsal surface

smooth.

Dimensions

Holotypc P2671

Figured specimen

P302785

Figured specimen

P302688

PL3032

Figured specimen

P302703

PL3037

Time range. Longfordian, Early Miocene.

Remarks. As pointed out by Pritchard (1896), this

species differs from Umhilia {U.) platyrhyncha

(McCoy) by its smaller size, more rapidly taper¬

ing anterior rostrum, more strongly developed

dorsal tubercles and the relatively thin sides of the

posterior canal. Although Pritchard considered

angnstior merely to be a variety of U. (U.)

platyrhyncha, the material before me, though

showing some variation, does not in facl

overlap in morphology with the latter and there¬

fore 1 accept angnstior as a full species. It seems

most likely to have evolved from U. ((/.)

platyrhyncha and to have probably given rise

to U. {U.) e.ximia. Some specimens are rather

tumid, approaching U. (U.) eximia in whorl shape

(Figs 2G-H), but lacking the prominent rostra of

U. (U.) eximia.

Umhilia (Umhilia) e.ximia

(G.B. Sowerby I, 1845)

Figures IB; 5A-H; 6A-H; 7B-I; 8A-H

Cvpraea e.ximia G.B. Sowerby I, 1845: 296, pi. 19

figs'I-3.—Tate. 1890: 209.

Cvpraea (Aricia) e.ximia. —McCoy, 1876: 35, pis.

27-28 figs 2-2b. All figures reversed in litliogruphing.

Cvpraea (Aricia) eximia var. brevis McCoy, 1876:

36.'

Cvpraea sphaerodoma Tale, 1890: 209.—Tate.

I89'2: pi. 8 fig. 5.—Harris: 1897; 211.

Cypraea (Umhilia) e.ximia. —Harris, 1897: 210.

Rhynchocypraea eximia. —Cossmann, 1903; pi. 6

fig. 11.

Rhynchocypraea lo.xorhynclia Tate.—Coss-

niann. 1903: pi. 8 fig. 5 (error for toxorliyncha) not

toxorhyncha ol'Taie, 1890.

Gisortia eximia. — Vredenburg, 1927: 50, pi. 3 fig. 5

(copy oTCossmann, 1903: pi. 6 fig. 11).

THE AUSTRALIAN GASTROPOD GENUS UMBIUA

361

UmhiUct (Umhilia) eximia maccoyi Schildcr. 1932:

183.—Schilder. 1935:344.

Umhilia (Umhilia) montis-marthae Schilder, 1935:

344, ng. 29.

Umhilia (Umhilia) brevis hrevis. —Schilder. 1935:

344.

Umhilia (Umhilia) hrevis frankslonensis Schilder,

1935: 344.

Umhilia (Umhilia) eximia eximia. —Schilder. 1935:

345.

Umhilia .spliaerocloma (Tate). —Lorenz. 1989. fig. 5.

Type and fiyiured material, llolotype BMNII G9694.

P.E. Strzeiecki Collection. The specimen bears a paper

label inside the shell bearing the number 215.

Syntypes of Cypraea (Aricia) eximia var. hrevis

NMV P5296, Geological Survey of Victoria. Geo¬

logical Survey of Victoria locality Aw 9. 3 miles W of

the mouth of the Gellibrand River; NMV PI2I36, Geo¬

logical Survey of Victoria, between Mount Eliza and

Mount Martha. The colour and preservation of PI 2136

suggests that the specimen probably came from the

Fyansford Formation at Fossil Beach, Balcombe Bay.

This specimen is chosen as Icctotype of hrevis.

llolotype of Umhilia (U.) brevis frankstonensis

BMNII G4()069, E.O Tealc Collection presented Jan

1926. This specimen is labelled ‘Grices Creek.

Frankston, Mornington’ on a printed label. Another

handwritten label bears the locality ‘Mornington’. The

light colour of the specimen and matrix (Fyansford

Formation) suggest that Grices Creek is correct.

llolotype of U. (U.) hrevis montismarihae BMNII

70406. purchased Bryce M. Wright. 90 Great Russell

St., London. This specimen bears a printed dealer’s

label with the handwritten number 17 and locality

‘Mount Martha. Victoria’. This specimen is abraded

and has fragments of hard dried clay inside the aperture.

It is typical in matrix, colour and condition of those

specimens washed out from the clay (Fyansford Forma¬

tion) and picked up on the surface at Fossil Beach. S of

Mornington.

Syntypes of Umhilia (U.) eximia maccoyi NMV

PI2135 (McCoy. 1876. figs 2, 2b). P16I7I (McCoy,

1876, fig. 2a) Geological Survey of Victoria, between

Mount Eliza and Mount Martha. The original labels

with these specimens are missing. The original draw¬

ings for McCoy's plate were undertaken by Arthur

Bartholomew in Jun and Jul 1861. which indicates that

the specimens were collected before then and so almost

certainly by Alfred Selwyn during his survey of the

Mornington Peninsula in 1854. The complete specimen.

PI2135, is chosen as lectotype of Umhilia (U.) eximia

maecoyi. The colour of the matrix, light grey, and the

light colour of the shells suggest that they were eol-

leeted from outcrops of the Fyansford Formation at

Grices Creek or nearby Manyung Rocks.

Lectotype of Cypraea sphaerodoma Tate SAM

T821, R. Tate Collection. ‘Eocene, River Murray

ClilTs, near Morgan’ i.c.. PL 3084 Small gully 4.8 km S

of Morgan Fcrry-Gadell road on left bank of Murray

River opposite Brenda Park Homestead, South Aus¬

tralia. Morgan 790280. Morgan Limestone, Cadell Marl

Lens, Middle Miocene, Balcombian. This is the original

of Tate. 1892, pi 8 fig. 5. which is chosen as lectotype.

Fieured specimens. P302779. F.A. Cudmore Collec-

tion.>3027l2, E.D. Gill. 27 Apr 1956. P3027I6. pre¬

sented H.J. llauschildt, 30 Jul 1906. P302782, T.A.

Darragh, 4 May 1996.

Type locality. The holotype described by Sowerby was

obtained by P.E. Strezelccki during his visit to Australia

in 1839-1843. According to the original description

given in Strezelecki (1845). the holotype was ‘Found in

a muddy sand, in sinking a well to 140 feet in depth, at

Franklin’s Village, Van Diemen’s Land, about 15 miles

from the sea’. There are two Franklin’s Villages in Tas¬

mania. The better known of the two is now an outer

suburb of Launceston on the Hobart Road about 5 km

south of the centre of the city and thus nowhere near ’ 15

miles from the sea’. It is situated in the Tamar Graben

which is filled with non-marine sediments. The possi¬

bility that the specimen could have come from here is

considered to be so remote as to be virtually impossible.

The other place bearing this name is on the coast at the

west end of Cape Barren I. in the Furncaux Group. Bass

Strait. At this locality, which I visited in February' 1969,

there is a thin sequence of Tertiary bryozoal calcarenite

sitting on granite within a kilometre of the sea. Again it

is most unlikely that the specimen could have come

from here, though Strzeiecki is known to have visited

the Furncaux Group. This species has never been

recorded from Tasmania, only from South Australia

and Victoria. Strzeiecki (1845) docs not mention this

fossil in his book at all, whereas virtually all the other

fossils described by Lonsdale. Morris and Sowerby are

mentioned either in Strzelecki’s discussion of the geo¬

logy of the country or under fossil fauna, in some cases

in both places. The two species of land snail. Bidimus

gimni and Helix lasmaiiiensis, illustrated on the same

plate as Cypraea eximia are mentioned hy Strzeiecki

twice, but neither C. eximia nor Terehralitia compla. a

brachiopod from Port Fairy, Victoria, also figured on

the plate, are mentioned at all. Strzeiecki does not even

record the genus Cypraea any where in the book, though

he mentions other genera of gastropods as occurring in

the fertiary of Tasmania. This suggests that there is an

error in the locality.

There is some matrix still adhering to the specimen

and also present inside the shell. The specimen is light

in colour with traces of yellow grey silt adhering to it.

Inside the shell were some polished shell fragments as

well as pieces of the grey silt. Of the Victorian occur¬

rences of the species, the area which contains sediments

which best matches the matrix is Geelong. I’he Fyans¬

ford Formation in the Geelong district, for example at

Red Hill, Shelford. is slightly weathered to a yellow

grey colour and has similar texture to that in and on the

type specimen. It is also pertinent to note that some

specimens from the Fyansford Formation around Gee¬

long seem to have the posterior canal twisted to the

right almost as in the holotype. a feature not so common

in specimens from other areas.

Strzeiecki was in Melbourne and could have received

the specimen there or even in Tasmania, because many

362

THOMAS A. DARRAGH

of the Port Phillip settlers came from Tasmania and

travelled back and forth. Sowerby described Terehnii-

ula compta from Port Fairy. Victoria, following his

description of Cypraca eximia which shows that

Strzelecki did have access to Victorian specimens, even

though he did not travel to Geelong or Port Fairy.

The weight of the evidence, though circumstantial,

seems to indicate that the true type locality was in

Victoria, probably some source within the Fyansford

Formation in the Port Phillip Embayment of the Otw ay

Basin.

Occurrence cmil material. Batesfordian. Fishing Point

Marl: Fishing {= Fischers) Point. Hordern Vale (1 frag¬

ment). Gellibrand Formation: PL3()43 Kennedy Creek

cutting (2 specimens). Wuk VVuk Marl: PL3054 Skin¬

ners (1 fragmentary specimen).

Balcombian. Muddy Creek Formation: PL3082

Clifton Bank (27 specimens): Muddy Creek. Hamilton

(24): Miocene. Hamilton (2); Grange Burn Gorge 3.

Fyansford Formation: PL3()6‘) Red Hill. Shelford (5);

PL3()7IW of Glen Leigh (I); PL3072 Fyansford (5);

PL3077 Allona Bay Coalhafl (4 and 3 fragments);

PL3()74 Oingleyhaft (2 and 3 fragments: PL3078 Fossil

Beach. Balcombc Bay (35); PL3079 lower beds. Grices

Creek (5): Old Quarries. Balesford (2); Overburden.

Batesford Limestone Quarry (5); Native Hut Creek

(upper) (I): Momington (6): south-eastern trunk sewer

spoil dump. Carrum (2): Braeside Tunnel spoil (1);

Shelford (3). Cadell Marl: River Murray clifls. 4 miles

south of Morgan (1 and 2 fragments). Morgan

Limestone; Broken Cliffs opposite Waikerie. River

Murray. South Australia (selenite replacement,

AM P29732).

Baimsdalian. Gellibrand Formation: PL3087 Lake

Bullcnmerri (3 and 3 fragments); Lake Gnotuk (I):

PL3088 east of Glenample Steps (2); Cowleys Creek

(1); Between Gibsons Beach and Point Ronald (10).

Rutledges Creek Member of Port Campbell Limestone:

PL3093 Rutledges Beach (1). Fyansford Formation:

PL3098 Native Hut Creek (3 fragmentary): PL3I()0

Murgheboluc 4A (7); PL3102 Warrambine Creek (2);

PL3I03 downstream Grices Creek (I); PL31()4

Manyung Rocks (15); Grices Creek (36): Leigh River at

Shelford (I): Inverleigh (I and I fragment): Mount

Eliza (1): Mouth of Grices Creek (1). Bookpumong

Beds: PL3298 Wookool (3 and 2 fragments).

Description. Shell globosely pyriform, with max¬

imum convexity about two-thirds length of shell

from anterior end: tapering rapidly anteriorly to

form long narrow, rounded anterior rostrum; ros¬

trum supported by 2 strong lateral elongate sub-

triangular extensions of base and bearing 2 promi¬

nent dorsal tubercles. Transverse section of last

whorl subcircular. Spire covered with thin glaze,

all whorls visible.

Posterior canal long to short, posterior end

slightly reflexed dorsally, twisted to left or right,

on most specimens to left; canal supported by

short flattened triangular extensions of base.

Anterior canal elongate, embraced by rostrum.

Aperture sinuous, outer lip with 23^0 teeth, on

most specimens present along entire length, on

some specimens fading at posterior third; col¬

umella bearing 23-37 prominent teeth, extending

along length of aperture, on .some specimens very

short, on most, extended into long ridges across

base away from aperture, sometimes bifurcating;

teeth encroaching on fossula on many specimens.

Fossula very narrow if at all present.

Dorsal surface on most specimens smooth,

shining, showing traces of mantle edges nmning

as shallow' sulcus from end of posterior canal

across left side of spire umbilicus across dorsum

between dorsal tubercles and onto anterior ros¬

trum; some specimens with very shallow depres¬

sions over shell.

Dimensions

Holotype BMNll

G9694

BMNll G40069

BMNll 70406

P5296

P12136

SAM T821

P12I35

103

P16I71

Figured specimen

P302779

PL3079

Figured specimen

P3027I2

PL3087

Figured specimen

P302716

crushed

Rose Hill

Figured specimen

P302682

PL3298

Time range. Batesfordian-Baimsdalian, Early

Miocene-Middle Miocene.

Remarks. This .species was the first Tertiary mol¬

lusc to be described from Australia. It is the most

common species of the larger cowries and one of

the most highly variable in morphology. The vari¬

ation is random through time and across the geo¬

graphic distribution of the species. This variabil¬

ity was not taken into account either by Tate, or

Schilder in particular, when erecting new taxa, so

that it probably has more synonyms than any

other Australian Tertiary mollusc. McCoy

remarked how variable the species was in the

degree of development of the anterior canal and

erected a variety brevis based on two small spec¬

imens with particularly short canals but without

illustrating them. This name was elevated to the

status of species by Schilder (1935), who did not

sec the specimens. There is every variation in size

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

363

of specimens and degree of development of the

eanals in the one population from Grices Creek

and from other localities, e.g.. Muddy Creek, con¬

sequently brevis is not recognised here as a valid

taxon.

Tate distinguished his speeies Cypraea

spluierodonui from C. eximia by its globose body

whorl, the abrupt sinistral curvature of the poste¬

rior half of the aperture, and the strong torsion of

the posterior canal. Specimens of similar shape

ean be found intergrading with other fomis in

populations from Fossil Beach (Fyansford For¬

mation) and Muddy Creek (Muddy Creek Forma¬

tion), so that this name may be regarded as a

synonym of Umbilici eximia.

The holotype of U. eximia. which matches most

other specimens in its general morphology, has

the distal end of the posterior canal twisted prom¬

inently to the left when viewed from the dorsal

side, whereas most other specimens have a pos¬

terior canal which is straight or has the distal end

twisted slightly to the right. 1 lowevcr, there are all

variations between the condition shown by the

holotype and that shown by the specimens figured

by McCoy (1876), in which the canal is bent

slightly to the right in dorsal view. This feature

was used by Sehilder (1932) to differentiate

between U. (U.) eximia and his new species,

U. (U.) eximia maccoyi (based on McCoy's fig¬

ured specimens), but given the variability in pop¬

ulations, the distinction cannot be sustained and

maccoyi is also placed in synonymy with U. (U.)

eximia.

Other features used by Sehilder in differentiat¬

ing his other new taxa, U. (V.) brevis fremk-

stonensis and V. (U.) brevis montismarthae, are

also variable. The columellar teeth, which are

produced into prominent ridges extending across

the base in the holotype and most other speci¬

mens, can be considerably reduced to ridges

extending across half the base or merely to the

edge of the columella. These ridges commonly

extend onto the fossula, such as it is; that is, they

commonly extend well into the aperture. The dor¬

sal tubercles are strongly developed in most spec¬

imens, but can be quite weak in others from the

same population. The holotype off/. (U.) brevis

monli.smarihae is water-worn, with extremities

that are broken and not ‘e.xtremely short' as main¬

tained by Sehilder. The sulcus connecting the

spire and dorsal side of the posterior canal, cited

as a diflerentiating feature of montismarthae, is

present to a greater or lesser extent on all speci¬

mens examined, including the lectotype of (7. ((/.)

brevis brevis. The dorsal tubercles present on the

holotype of U. (U.) brevis frankstonensis are no

weaker than those found on other specimens with

prominent ridgelike teeth from the same locality.

There are all degrees of variation in specimens

from Grices Creek. For these reasons, both

Schilder's names arc also placed in synonymy

with Umbilia eximia.

Only two undoubted specimens of U. (U.)

eximia arc known from the Gippsland Basin,

though internal moulds probably of this species

are common in the Bairnsdale Limestone. The

two specimens come from PL3054, Skinners, on

the Mitchell River (Batesfordian) and Rose Hill

(no other locality data), near Bairnsdale. The

Rose Hill specimen (Fig. 7F-G) does not have

matrix typical of the Tambo River Fonnation at

that locality and may have come from the top of

the underlying Bairnsdale Limestone. Specimens

of U. (U.) hesitate! occur in the Tambo River For¬

mation at Rose Hill, so if the Rose I fill specimen

of U. (U.) eximia does come from there, it

would make an ancestor/descendent relationship

between these two species unlikely. Both U. (U.)

hesitata and U. (U.) eximia also occurr in the

Bookpumong Beds at Loxton, though as yet not

recorded from the same localities within the

fomiation.

Umbilia (Umbilia) hesitata (Iredale, 1916)

Figures 7A; 9A-H; lOA-l; 1 lA-H; 12A, C,

F-H; I3D-E; I4A-C; 19C

Cypraea umbiiicata Sowerby. 1825, appendix; xxx,

pi. non Dilwyn, 1823.—Beddome, 1898: 564 (with dis¬

cussion on early literature).—Verco, 1912: 211 (with

synonymy).

T Cypraea amygcialina Talc. 1890: 209.—Tate, 1892,

pi. 6, fig. 8, non Grateloup, 1847.

7 Cypraea lalei Cossmann, 1903: 160, pi. 7 figs 4, 6,

nomen novum for Cypraea amygcialina Tate, 1890.

Cypraea hesitata Iredale, 1916; 93, nomen novum

for Cypraea umbiiicata Sowerby, 1825.

Umbilia hesitata beddomei Sehilder, 1930: 77.

Umbilia hesitata howetli Iredale, 1931: 220-224, figs

1 - 2 .

Umbilia cera Cotton, 1947: 667, pi. 21, figs 1-3.—

Ludbrook, 1958: 45.

Cvpraea tatei Cossmann.—Ludbrook, 1973, pi. 27

figs'87-88.

Cypraea (Umbilia) hesitata. —Wilson, 1993: 192, pi.

30 tigs 7,9-11.

Type and figured material. Lectotype BMNH

1950.8.28.22, Broderip Collection (Sowerby's figured

specimen chosen as lectotype here).

Holotype of U. cera Cotton, SAM P8339, donated

H.S. Pratt, 1925.

Syntypes of Cypraea amygdalina Tate (= tatei

364

THOMAS A. DARRAGH

Cossmann), SAM T8I2A. B. Ralph Tate Collection.

T8I2A Tate's figured specimen chosen as lectotype

here. Matrix associated with the two specimens is a car¬

bonate rich glauconitic sand. The type locality cited was

a well sinking in the Murray Desert. Tate had material

from wells at Tareena in southwestern New South

Wales and Mindarie in South Australia and it is not

known from which place the type specimens came,

since Tate (1899: 104) recorded his species from both.

Whatever the origin, the shells seem to have come from

the Bookpurnong Beds.

Holotype of Umhilia lie.tilaln heddoniei Schilder,

1930. Schilder collection no. 821, Zoologisches Institut,

Humboldt Museum, Berlin (locality unkown),

purchased Fulton.

Holotype of Umhilia hesilala howelli Iredale, 1931.

AM C57762.

Figured specimens. P3027I7- P302718. John Den-

nant Collection. Figured specimen P302720, G.B.

Pritchard Collection. P302780, collected and presented

J.M. Warren. P302770. T.A. Darrach. D.M. Shanks and

H.E. Wilkinson. 8 Feb 1969. P30278I. T.A. Darragh.

D.M. Shanks and H.E. Wilkinson, 11 Feb 1969. F4526.

J.H. Maepherson. 6-11 Jun 1948. F23I66, presented

T.A. Garrard. Figured specimens WAM 89.636a-b.

G.W. Kendrick. 27-30 Oct 1988. Geological Sursey of

South Au.stralia Ml335, M.J. Paul (fieured Ludbrook.

1973).

Occurrence and material. Baimsdalian. ? Dry Creek

Sands: Abattoirs Bore (fragmentary specimens).

Bookpurnong Beds: 1.5-3 m below bed of River

Murray. Loxlon, South Australia.

Mitchellian. Tambo River Formation: Rose Hill

Farm near Bairnsdale (2 specimens).

Cheltcnhamian. Black Rock Sandstone: Beaumaris

(3). Jemmys Point Formation: PL3115 Lake Bunga

Crossing (2 fragmentary specimens); PL3I23 Ferndale

Parade (2 fragmentary specimens).

Pliocene. Jemmys Point Formation: PL3275 Gosnells

Point (I specimen). Cameron Inlet Fomiation: PLI250

(2). PL1258(3). PL126I (1), PLI264(17), PLI265(2),

PLI287 (2), PL1296 (I). West end of North Memana

Drain. Memana, Flinders I. (3); Flinders 1. (1); South

side of Nelson Lagoon Drain, Flinders I. 018588 (1);

Land Settlement Division Drain. Flinders 1. (3). Roe

Calcarenite: PL3166 2.5 km N Hampton Tower (4),

PL3I65 16 km S of Madura (1), PI3I67 1.5 km N

Hampton Tower (I), PL3172 Hampton Tower (2).

Records of living species are from north-east of Cape

Morton, Queensland to Robe. South Australia, includ¬

ing northern and western Tasmania as far south as

Port Davey, 70—JOO m. Pliocene records from the

Roe Calcarenite lie outside of the recorded modern

distribution.

Description. Shell of small to average size for

genus, highly glazed, subpyrilbmi, tapering gen¬

tly anteriorly and produced into short rounded

rostrum bearing traces of 2 dorsal tubercles,

slightly rostrate posteriorly. Spire umbilicate,

covered with glaze, about 5 whorls visible.

Posterior canal very short, margins thickened,

twisted to left in ventral view. Anterior canal

short, embraced by rostrum, supported by very

weak lateral llanges. Aperture sinuous, outer lip

with 30^0 teeth; inner lip with 28-37 teeth

which become slightly weaker on posterior half of

columella. Fossula very narrow, very slightly

depressed. On some specimens slight trace of sul¬

cus running from left side of posterior canal

across dorsum onto anterior rostrum between

tubercles.

Colour white with irregular sized brown spots

over dorsum and large brown patches on anterior

rostrum and on most specimens on either side of

posterior canal; ventral surface white. Some

specimens almost entirely while.

Time range. Baimsdalian?, Mitchellian-Recent,

Middle Miocene?, Late Miocene-Recent.

Remarks. This species may be descended from

Umhilia (U.) eximia but, as noted above, there

could be an overlap in the time ranges of the two

species in the Middle to Late Miocene. U. (U.)

LT CT

33 31

31 27

32 31

28 28 off Botany Bay

34 35

29 25 heddomei form

38 31 holotype of U. howelli Iredale.

26 22 holotype of U. cera Cotton

22 29 lectotype of Cypraea lalei

Cossmann.

- 30

24 22

29 23

33 30

36 30

Dimensions

Lectotype BMNH 1950.8.28.22

Figured specimen P302781

Figured specimen P302780

Figured specimen P302770

Figured specimen F4526

103

Figured specimen F23116

AM C57762

111

SAM P8339

Figured specimen SAM T812A

Figured specimen P302717

Figured specimen P3027I8

Figured specimen P302720

Figured specimen WAM 89.636a

Figured specimen WAM 89.636b

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

365

liesUata dilTers from U. {U.) eximia in having less

prominent columellar teeth and a less prominent

anterior rostrum and posterior canal. The dorsal

tubercles on the anterior rostrum are also weakly

developed and the lateral llanges supporting the

rostrum are obsolete in U. hesitata. The differ¬

ences between U. hesitata and the other southern

Australian living species, U. armettiaca, are not

great. U. hesitata tends to be slightly less tumid

than U. armettiaca and has a slightly longer ante¬

rior rostrum. On our present knowledge of the

distributions of both ta.xa, there is a considerable

gap between the range extremities of both

species.

The type specimens of both Cypraea tatei

Cossmann (probably Bookpumong Beds at

Tareena or Mindarie) and Umbilia cera Cotton

(Dry Creek Sands) are both small and fragmen¬

tary. They seem to be similar if not identical to

small specimens of U. hesitata, but until a range

of material from the Bookpumong Beds and Dry

Creek Sands is available, it is not possible to be

certain of the identity of Cypraea tatei in particu¬

lar, since the apertures on the specimens look

immature. For this reason I have included the

name with a quer>' in the synonymy above, at the

same time preserving the stability of the name of

the well known common living species, Umbilia

hesitata. Since both tatei and cera seem to come

from Middle Miocene strata, there is also a possi¬

bility that they are small specimens of U. eximia.

This seems unlikely, but the anterior rostra on all

specimens are broken, so there is doubt about

their identity.

Late Miocene and Early Pliocene specimens of

U. hesitata are not common and most arc frag¬

mentary or poorly preserved; however, there is

sufficient well preseiA'cd material available to be

confident of the identification.

Umbilia hesitata beddomei was based on a

small fomi, with somewhat brighter coloured dor¬

sum, base with less callus and whiter. The holo-

type is from an unknown locality (Dr Matthias

Glaubrecht pers. comm. January 2000), but

Schilder cited Port Stephens, New South Wales as

a locality based on Beddome’s (1898) description

and figure of a similar specimen. Such small

specimens (Figs 12A, C) are found throughout the

geographic range of the species as well as in the

fossil record (Figs 90-11, lOA-C) and intergradc

with larger specimens from the same locality. For

these reasons Umbilia hesitata beddomei is

regarded as a synonym of hesitata and cannot be

accepted as a geographic subspecies.

Umbilia hesitata howelli was erected by Iredale

(1931) as a new subspecies ((type locality

164-274 m, olTCape Everard (Point Hicks), Bass

Strait, Victoria) for a large, pure white variety

previously called variety alba Cox, 1879 (non

Blainville, 1826, nec Sowerby, 1832). Apart IVom

its colour, it cannot be distinguished from other

specimens of hesitata and is regarded as a mere

colour form and therefore synonymised.

Utiihilia {Umbilia) prosiia sp. nov.

Figures 15A-C, E, H

Type material. Holotypc P3087I6, Paratype P308717,

F.A. Cudmore collection.

Type locality. Ledge, Bird Rock Cliffs, Torquay, Victo¬

ria. Jan Juc Formation, Janjukian, Late Oligocene,

Occurrence ami material. Jan Juc Formation. Ledge,

Bird Rock, Torquay (9 specimens); Spring Creek,

Torquay (8); Bird Rock cliffs, Torquay (2); below Bird

Rock cap between Fishemians Steps and Bird Rock,

Torquay (1); Geological Survey of Victoria locality

Ad24, Bird Rock (I) and Ad23, Bird Rock (1).

Description. Shell very small for genus, delicate,

highly glazed, umbilicate posteriorly, very glo¬

bose with short, very narrow anterior rostrum;

rostrum supported by very weak extensions of

base. Spire depressed, covered with thin callus, of

3 teleoconch whorls. Protoconch 3.5 smooth

whorls, axis tilled slightly from axis of shell.

Posterior canal very short, strongly bent to left,

bounded on columellar side by prominent blunt

ridge. Anterior canal short, deeply sunk into ros¬

trum. Aperture virtually crescent shaped, narrow¬

est medially; outer lip with 20-26 relatively thick,

elongate teeth, extending along entire lip; inner

lip with 16-22 well developed, short teeth,

extending along entire lip, almost nodulate on

anterior half, more elongate posteriorly. Fossula

wide, Hat to slightly concave, slightly notched

anteriorly on interior side and bounded anteriorly

by weak ridge.

Dimensions

Holotypc P3()8716

Paratype P3()87I7

Time range. Janjukian, Late Oligocene.

Remarks. Specimens of this species are not com¬

mon. This is the smallest species of the genus and

seems to be ancestral to Umbilia (U.) leptorhyn-

cha (McCoy). It differs from U. (U.) lepto-

rhyneba by its smaller size, being nearly

half the length, by having a relatively short

anterior rostrum, and by having relatively

thicker teeth. The anterior Hanges supporting the

rostrum present in leptorhyncha are scarcely

developed in this species. The species shows

little resemblence to the putative ancestor of

366

THOMAS A. DARRAGH

Umbilia, IPalaeocypraea eripnides sp. nov.,

or to the other species of Umbilia found at the

same horizon, U. (U.) platyrhyncha. It may have

developed from an undescribed species of

Cypraeorbia from the Late Eocene of Western

Australia by weakening of the fossula and

extension of the anterior canal. Cypraeorbis

sp. has a short anterior canal and a deep concave

fossula bounded on the anterior by a thick

ridge and with a notch in the interior margin

immediately behind the ridge.

Umbilia (Umbilia) leptorhyncha (McCoy, 1877)

Figures 16A-H

Cypraea (Liiponia) leptorhyncha McCoy, 1877: 35,

pi. 49 fig. 1, la-c.—Harris, 1897: 207.

Cypraea leptorhyncha. —Tate, 1890: 208.

Gisortia leptorhyncha. —Vredenburg, 1927: 42.

Umbilia (Rhyneboevpraea) leptorhyncha. —Schildcr,

1935:343, fig.'27.

Rhynchocypraea leptorhyncha. —Lorenz. 1989: 7,

fig-

Type material. Lectotype P12133, McCoy’s figured

specimen chosen herein. Paralectolypes P308726-

P308729. Origin unknown (possibly J. Kershaw, 1875).

Type locality. “Near foot of Mount Eliza and Mount

Martha, Momington". The paralectotypes arc labelled

near Mount Martha. The matrix on the lectotype indi¬

cates that it was collected at Fossil Beach, i.e. PL3078

Shore platform at F-ossil Beach, 3 km S of Momington.

Victoria. AMG Western Port 273653. Fyansford For¬

mation. Balcombian, Middle Miocene.

Occurrence and material. Batesfordian. Gellibrand

Formation: PL3043 Kennedys Creek cutting (2

specimens): Fyansford Fonnation: Curlewis (2).

Balcombian. Fyansford Formation: PL3078 Fossil

Beach (16 specimens); Balcombe Bay (29): near Mount

Martha (4); Momington (6); Schnapper Point (2);

PL3069 Red Hill (I): PL3072 Orphanage Hill (I);

PL3077 Altona Bay Coal Shaft (1); Shelford (I).

Muddy Creek Formation: PL3082 Clifton Bank (25):

Muddy Creek (6); Hamilton (3). Cadell Marl: Murray

River cliff. 4 miles downstream from Morgan, South

Australia.

Baimsdalian. Fyansford Fonnation: Grices Creek (4

specimens); Lower beds. Grices Creek (2); Mount Eliza

(1); PL3104 S of Manyung Rocks; PL3097 Murghe-

boluc 2B (2); PL3100 Murghcboluc 4A (1 and I frag¬

ment); Inverlcigh (1 fragment); Overburden, Batesford

Limestone Quarry (1).

De.tcription. Shell of medium size for genus, thin

shelled, delicate, highly glazed, umbilicatc poste¬

riorly. globosely pyrifonn, tapering abruptly to

narrow anterior rostrum; rostrum stipported by

narrow triangular extensions of base. On some

specimens, rostrum bearing deep oblique groove

on dorsal surface formed by anterior end of dorsal

line (mantle margins). Spire depressed, covered

with thin callus; 5 teleoconch whorls. Protoconch

of 2.5 smooth whorls with axis slightly tilted from

axis of teleoconch whorls.

Posterior canal, very short, .strongly bent to left,

bounded on columellar side by low sharp ridge.

Anterior canal short, deeply sunk into rostrum.

Aperture very sinuous, narrowest medially,

widest posteriorly; outer lip with 35 39 thin, well

developed, well spaced teeth, extending along

entire lip; inner lip with 32-34 well developed,

thin, widely spaced teeth, extending along entire

lip; teeth longer in middle section, with some

short teeth intercalated, becoming very short pos¬

teriorly and extending onto posterior canal wall;

teeth almost lamellar on some specimens. Fossula

moderately developed, rather elongate, narrow,

depres.sed, inner border slightly thickened and

weakly indented. Base and portion of sides cov¬

ered with very thin callus.

Dimensions

Lectotype PI2133

Paralectotype

P308726

Paralectotype

P308728

Time range. Batesfordian-Bairnsdalian, Middle

Miocene.

Remarks. The geographic distribution of this

species is probably much wider than that given

above and it is more common than the numbers

would suggest, because specimens arc generally

so fragile that they have not survived intact at

many localities. The specimens from Curlewis

and Kennedys Creek have the columellar side of

the posterior canal very well developed when

compared to specimens from younger horizons.

The apertural teeth arc aLso not so well developed

and look more like those of U. (U.) prosila sp.

nov.

Umbilia petilirostris sp. nov.

Figures 13A-C, I5D, F-G

Umbilia capricornica Lorenz. 1989: 2, pi. 2 (part,

deep water variants).

Cypraea capricornica. —Wilson, 1993: 192, pi. 30,

fig. 6 (part).

Type material. Hololype F86962, paratypes F86963,

F86967, presented Alan Limpus, 30 Jun 1997.

Type locality. Capricorn Channel, off Lady Musgrave

1., Queensland. 264-252 m.

Occurrence. Capricorn Channel, Queensland - type

loeality; off Fitzroy Reef 249 m; off One Tree I.; off

Swains Reef 188-201 m.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

367

Description. Shell of medium lo large size for

genus, thin shelled, highly glazed, po.steriorly

umbilicate, very globose, tapering abruptly to

short anterior rostrum; rostrum supported by very

narrow, thin extensions of the base and bearing 2

very weakly developed tubercles, separated on

some specimens by wide, shallow, oblique

groove. Spire depressed, covered with thin

callus; teleoconch whorls 3; protoconch of 2.5

smooth whorls, axis tilted slightly from axis of

teleoconch.

Posterior canal very short, strongly bent to left,

bounded on columellar side by thin high wall.

Anterior canal relatively wide, deeply immersed

in rostrum. Aperture sinuous; outer lip with

24-34 short, well developed teeth, extending

along entire lip; inner lip with 20-32 short, well

developed teeth, extending along entire lip and on

some specimens extending onto posterior canal

wall; teeth more elongated posteriorly and almost

nodulate anteriorly above fossula. Fossula

relatively long and wide, concave with weak

depression on inner border.

Dorsum grey brown with brown spots particul¬

arly near base. Base white with dark brown patch

on columellar side.

Dimensions. L W 11 LT CT

Holotypc F86962 76 49 40 34 32

Paratype F86963 67 47 38 28 28

off Fitzroy Reef

Paratype F86967 53 34 28 29 24 off

Swains Reef

Remarks. This species has been confused with

Umhilia (U.) capricornica Lorenz, but differs by

its more globose shape, more weakly developed

posterior rostrum, and by having the columellar

side of the posterior rostrum developed as a thin

wall. It seems to be the living descendent of U.

(U.) leptorhyncha from which it differs by its

much larger size, the presence of a seemingly

relatively larger fossula, and by having the pos¬

terior canal as a prominent thin wall on the

columellar side, whereas U. (U.) leptorhyncha

has a low ridge. The two species are very similar

in shape though perhaps U. (U.) petilirostris is

even more globose and with shorter labial teeth

than U. (U.) leptorhyncha.

Umhilia (Umhilia) siphoiiata Chapman, 1922

Figures I7A-F; 18A-F; 19A-B

Cypraea spluierodoma var. ? Tate, 1890: 210.

Cypraea siplwiutia Chapman. 1922; 12, pi. 3 fig. 16.

Gisortia hrevipUcata Schilder 1926: 361,373.

Umhilia (Palliocvpraea) hrevipUcata Schilder, 1935:

345, fig. 30.

Umhilia sp.— Burgess, 1989: 11, fig. C.—Lorenz,

1989: fig. 5.

Type ami figured material. Holotype PI3243, F.A.

Cudmore, presented 8 Oct 1920.

Holotype of Gisortia hrevipUcata P26904, John

Dennant Collection. There are two specimens in the

Dennant Collection from Muddy Creek labelled

Cypraea sphaerndoma. One is near complete

(P302803), but lacks the anterior canal and so could not

be that referred to by Tate. It is also too small. The other

specimen was found in fragments. This shell is very

fragile, the dorsum having had been subjected to boring

organisms (a common feature on many specimens of

larger cowries) and at some stage it had been broken.

This specimen was repaired to enable measurements

to be taken, and on repair it was found that Tate's

measurements fitted it.

Figured specimens. PI2569, purchased R.H. Annear,

23 Jan 1912. P302803, John Dennant Collection.

PI4835, Noel .1. Shaw. 13 Mar 1950.

Type locality. 'Below Overland Corner (left bank), and

second cliff showing strata, below Waikerie. Murray

River, South Australia. From upper part of the cliff

below the Kalimnan beds’.

Occurrence and material. Balcombian. Muddy Creek

Formation: PL3082 Clifton Bank (I specimen); Muddy

Creek (6 and 11 fragments). Fyansford Formation:

PL3069 Red Hill, Shclford (I and 2 fragments);

Orphanage Hill, Fyansford (I fragmentary specimen).

PL3074 Heatherton lest shaft. Morgan Limestone:

Broken Cliffs opposite Waikerie, River Murray cliffs.

South Australia (selenite replacement. 1 fragmentary

specimen. SAM P35237).

Baimsdalian. Bainisdale Limestone: I mile from

Baimsdale (1 natural internal mould)?; Nicholson River

(I specimen with remains of shell). Fyansford Forma¬

tion: PL3097 Murgheboluc 2B (2 specimens); PL3098

Native Hut Creek, S of Hamilton Highway (1 frag¬

ment); PL3102 Warrenbine Creek (1); PL3I()4

Manyung Rocks (I and 1 fragment); Grices Creek (2).

Gellibrand Formation: PL3087 Lake Bullen Merri (I

and several fragments). Bookpumong Beds: PL3I33

Wookool Bend (I); PL3298 Wookool 1 lomestead (I

fragment). PL6667 NW of Wookool Homestead (2

fragmentary specimens).

Description. Shell large for genus, subhemispher-

ical, tapering rapidly anteriorly to form tiarrow

canal; prominently rostrate posteriorly. Trans¬

verse section of last whorl oval. Spire umbilicate,

covered with glaze, whorls scarcely visible.

Protoconch of 3.5 smooth, dome-shaped

whorls, tilted at an angle to teleoconch whorls.

Teleoconch of about 3 whorls.

Posterior canal extremely elongate, rellexed

dorsally on most specimens, on others straight;

weak longitudinal sulcus present on left dorsal

side extending from end of canal into umbilicus;

base of canal supported by lateral extensions of

368

THOMAS A. DARRAGH

base; left extension triangular, prominent; right

extension elongate, narrow. Anterior canal mod¬

erately short (much shorter than posterior canal),

reflexed dorsally on some specimens (extremely

so on I specimen), almost straight on others,

sometimes bearing 2 very weak dorsal tubercles;

dorsal surface of canal bearing long sulcus, obsol¬

ete on some specimens; base of canal supported

by narrow elongate extensions of the base, right

extension longer than left.

Aperture sinuous; outer lip wide, slightly con¬

cave close to inner edge of lip, convex towards

periphery of base, bearing 30-34 relatively thin

but prominent teeth, well separated by interspaces

about twice width of teeth; teeth fading poster¬

iorly; inner lip with 26-28 teeth slightly wider

than labial teeth, very short anteriorly, longer and

slightly weaker posteriorly. Fossula broad, very

slightly depressed.

Base on columellar side slightly swollen in

middle and concave posteriorly and anteriorly

towards canals.

Dimensions

Holotype PI3243

135

P26904

133

Figured specimen

144

PI 2569

Figured specimen

111

P302803

Figured specimen

168

PI4835

Time range. Balcombian-Baimsdalian, Middle

Miocene.

Remarks. This is the second largest species of

Australian fossil cowry, exceed only by speci¬

mens of ZoUa gigas (McCoy). The species was

recorded as a questionable and unnamed variety

of Cypraea sphaerochma by Tate (1890). based

on a specimen from Muddy Creek in the collec¬

tion of John Dennant. In 1922 Chapman named

C. siphonata based on a natural mould in lime¬

stone from the Murray River near Waikerie and

compared it with C sphaerachma. Chapman con¬

fused anterior and posterior canals in his descrip¬

tion. Schilder (1926) recognised that Tate's

variety was a new species and fonnally named it.

based on Tate’s description and material. Later

Schilder (1935) stated that C. siphonata was

‘allied’ to C. hreviplicata, including both species

in the subgenus Palliocypraea on the basis of

their basal extremities being much expanded. In

fact, the extensions of the base on these two arc

no different from those of any other of the fossil

species included in Umhilia (Umhilia) and for this

reason, the two species are placed in this sub¬

genus.

The holotype of C. siphonata has sufficient

characters preserved to show that hreviplicata is a

synonym. The long posterior canal is the same as

that on a specimen of hreviplicata from Grices

Creek and the whorl profile is also identical in

shape. A rubber cast taken from the holotype

mould shows teeth identical to those on speci¬

mens of hreviplicata and the base has the same

slight swelling in the middle of the columella

side.

This species is rare in the Fyansford Fomiation

but not uncommon in the Muddy Creek Forma¬

tion. The occurrence of both this species and

Umhilia eximia in the Bookpurnong Beds of tlie

Murray Basin suggests that this formation is prob¬

ably Baimsdalian, that is Middle Miocene, rather

than Late Miocene in age.

Of the two specimens from the Baimsdale

Limestone, one is an internal mould. It is recorded

with a query but the si/e of the specimen and the

elongated anterior canal seems to indicate that it

is probably this species. The other specimen is

incomplete but has significant remnants of ori¬

ginal shell preserved, so that the identification

seems certain. It has the typical oval outline of the

dorstim and teeth typical of U. siphonata.

Umhilia (Umhilia) armeniaca Verco, 1912

Figure I2B. D-E

Cvpraea imihilicata var. armeniaca Verco, 1912:

211,213, pi. 10 figs 1-3.

Cypraea (Umhilia) armeniaca. —Wilson. 1993: 191,

pi. 30 figs 12-16.

Type ami figured material. Type stated to be in J.C.

Verco collection. Verco presented his collection to the

South Australian Museum, but the holotype cannot he

located there (T. Laperousa/, Collection Manager,

Marine Invertebrates, pers. comm., 12 Oct 1999).

Paratype AM C35583 (ex E3842), Verco’s ■slightly

older specimen'. Paratype AM 121153 (ex E3843).

Verco's ‘youngest example’. The holotype is not in the

Australian Museum collection.

Figured specimen. F27194, C.J. Gabriel collection,

presented 29 Aug 1963.

Type locality. ‘100 fathoms. Great Australian Bight. 60

miles from shore. 80 miles west ofEucIa’.

Description. Shell solid, globular, pyrifomi. of

large size for the genus, tapering abruptly anteri¬

orly and produced into a very short rostrum. Spire

deeply umbilicate, covered with glaze, 2 whorls

visible.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

369

Posterior canal very short, margins thickened,

twisted to left. Anterior canal very short,

embraced by rostrum, supported by weak lateral

flanges. Aperture sinuous, outer lip with 32-40

teeth, inner lip with 26-31 teeth, which become

slightly weaker posteriorly. Fossula on some

specimens wide, distinctly depressed.

Colour variable; apricot, orange, cream to

almost lilac blotches on white to grey ground,

darker patches on anterior and posterior rostra.

Ventral surface very pale, with many specimens

having dark patch in centre of base.

Dimensions L W H LT CT

Figured Specimen 89 57 51 39 31

F27194

Distribution. Albany, Western Australia to Port

Lincoln, South Australia, 30-200 metres.

Remarks. To date, this species has not been found

as a fossil. This may be accounted for by its

inhabiting deep water over most of its known

range and there being no outcrops of deep water

sediments of appropriate age, though specimens

of U. (U.) armeniaca have been collected by

divers in Thorny Passage near Port Lincoln and at

Esperance in 30 metres (C. Goudey collection).

Umhilia (Unihiiia) capricornica Lorenz, 1989

Figure 14D-H

Umhilia capricornica Lorenz, 1989: 2-8, pi. 2 (part),

ftgs 3-4.

Cypraea (Umhilia) capricornica.- —Wilson, 1993:

192; pi. 30 figs 1-5, 8.

Type and figured material. Haus der Natur-Cismar,

Germany. Holotype HNC 22453.

Figured specimens F85329, F85330, presented Alan

Limpus, 30 Jun 1997.

Type locality. Swains Reef, 120-125 fm, trawled Feb

r989.

Description. Shell somewhat small for genus,

pyriform, tapering relatively abruptly anteriorly

to form a moderately developed rostrum bearing

2 moderately developed dorsal tubercles; some¬

what rostrate posteriorly. Spire umbilicate,

covered with thick glaze

Posterior canal short, margins thickened,

twisted to left. Anterior canal embraced by ros¬

trum supported by weak lateral flanges. Aperture

sinuous, outer lip with 28-34 teeth; inner lip with

22-29 teeth. Fossula well developed, rather wide,

concave, projecting slightly into body of shell. On

some specimens slight trace of sulcus running

from left side of posterior canal across dorsum on

to anterior rostrum between tubercles.

Colour cream to white with irregular brown

spots and patches on anterior rostnim and on

either side of posterior canal; ventral surface

brown to white.

Dimensions H W H LT CT

Figured specimen 82 45 39 29 29

F85329

Figured specimen 60 36 29 30 22

F85330

Distribution. Swains Reefs, 165-420 m; Capri¬

corn Channel, off Lady Musgrave L, 220 m.

Remarks. Lorenz (1989) distinguished this

species from Umbilia (U.) lies data by the pres¬

ence of well developed tubercles on the anterior

rostrum, the coarser apcrtural dentition and by the

presence of a visible fossula margin. The general

colour and colour pattern of U. (U.) capricornica

are very similar to those of U. (U.) hesitata, the

posterior rostrum is very similar, the anterior ros¬

trum is more slightly produced and the dorsal

tubercles are more strongly developed; however,

the overall morphology of U. (U.) capricornica,

including the apertural dentition, is not very dif¬

ferent from that of northern specimens of U. (U.)

hesitata. Even the fossula in some specimens of

U. (U.) hesitata has a slight resemblance to that in

U. (U.) capricornica. Nevertheless, there do seem

to be sulTicient differences between them to

maintain specific separation of the two taxa.

Lorenz stated that the U. (U.) capricornica was

more closely allied with the fossils U. (U.) e.ximia

and U. (Palliocvpraea) gastroplax than with the

other living species; however, as noted above,

U. (U.) capricornica is not greatly different from

U. ((7.) hesitata. The resemblance to U. (U.)

eximia is not so great, but both U. (U.) hesitata

and U. (U.) capricornica are probably descended

from U. (U.) eximia.

Umhilia {PaUiocypraea) Cossmann, 1906

Rhvnchocvpraea (PaUiocypraea) Cossmann, 1906:

239 ■

Gisortia (PaUiocypraea). —Vredenburg, 1927: 26,

Umhilia (PaUiocypraea) Schildcr, 1935: 342.—

Sdiilder, 1939: 186.'

Type species. Cypraea gastroplax McCoy, 1867 by

original designation; Miocene, Victoria.

Diagnosis. Shell relatively thin, of medium size

for the family, body of shell pyriform with promi¬

nent thin Hat ftange extending around periphery

of body to give a circular outline. Rostra and

siphonal canals incor-porated into flange. Other

shell features similar to Umbilia.

370

THOMAS A. DARRAGH

Time range. Batesfordian-Balcombian, Early Miocene

to Middle Miocene.

Distribution. Victoria

Remarks. The overall shell morphology of the

only species assigned to the subgenus is very sim¬

ilar to that of Umhilia sensu stricto, with the

exception of the prominent lateral flange encir¬

cling the body of the shell. This flange can be

envisaged as an extensive development of the tri¬

angular basal extensions that support the rostra in

species of Umhilia sensu stricto. As mentioned

above, U. siphonata, included in this subgenus by

Schilder (1935), has basal flanges similar to those

of other species of Umhila {Umhilia), and is

placed in the latter subgenus.

Umhilia (Palliocypraea) gastroplax

(McCoy, 1867)

Figures 19D-E; 20A-D; 2IA-D

Cypraea gasiroplax McCoy, 1867a: 18.—McCoy.

1867b: 194.

Cypraea (Aricia) gasiroplax McCoy, 1875:20, pi. 16

figs 1-la, pis 17-18 figs 2-2a. Figures reversed in

lithography.

Rhynchocypraea (Palliocypraea} gasiroplax .—

Cossmann. 1906: 239. pi. 9 figs 10-11.

Pallion’praea gasiroplax. —Chapman, 1929:

202-205.’pis 19-20.—Lorenz. 1989: fig. 5.

Umhilia (PalHoc\praea) gasiroplax. —Schilder.

1935:342. 345.

Type and figured malerial. llolotype I’12140. The

original drawings for McCoy's plate are dated Jul 1861

which proves that the specimens were collected before

then and so almost certainly by Alfred Selwyn, Geo¬

logical Survey of Victoria, during his survey of the

Mornington Peninsula in 1854. The specimen was orig¬

inally registered under number 8060 (registered in

1861) under the old registration system, but the entry

has no further information beyond the locality 'Ml.

Eliza to Mount Martha'.

Figured specimens. PI3373. presented by Walter

Greed 3 Mar 1924. P24869, G.B. Pritchard Collection.

P302852.T.A. Darragh 23 Oct 1971. P302854, Nielson

Collection, presented Mrs W.A. Nielson, I May 1985.

Type locality. ‘Tertiary limestone of the tract between

Mount Eliza and Mount Martha'. The preservation of

the holotype in a block of septarian limestone indicates

that it came from PL3078 Fossil Beach.

Occurrence and malerial. Batesfordian. Gellibrand

Formation: PL3048 Boomong Rd cutting (2 frag¬

ments).

Balcombian. Muddy Creek Formation: PL3()82

Clifton Bank (2 specimens and 2 fragments). Fyansford

Formation: PL3077 Altona Bay Coal Shaft (I frag¬

ment); PL3()78 Fossil Beach (4 specimens and 2

fragments); Overburden, Batesford Limestone Quarry

(1 specimen and 2 fragmentary specimens).

Description. Shell of average size for genus; last

whorl pyriform, tapering gently anteriorly with

prominent wide flange extending around peri¬

phery of whorl; llange width about half width of

last whorl, flange thickness about 1 mm. Trans¬

verse section of last whorl subtriangular. Spire

umbilicate; spire whorls scarcely visible, covered

with glaze.

Posterior canal incorporated into shell flange,

slightly reflexed dorsally, subcylindrical. almost

tubelike anteriorly, almost closed on some speci¬

mens. closed on others; weak longitudinal sulcus

present on left dorsal side of posterior canal and

extending from middle of canal into umbilicus

and across dorsum to posterior canal on some

specimens. Anterior canal subcylindrical incor¬

porated into shell flange; dorsal surface of canal

bearing very weak sulcus. Aperture narrow, sinu¬

ous; with 30-39 labial teeth; teeth prominent,

thin, short, tooth interspaces about twice width of

teeth; inner lip with 24-30 short thin teeth about

half width of interspaces; teeth set on narrow

rounded ridge running from anterior canal to

posterior canal. Fossula elongate, narrow, slightly

depressed. Ventral surface on both sides of

aperture uniformly convex.

Dimensions

llolotype PI2140

Figured specimen

102

P24869

Figured specimen

364-

P13373

Figured specimen

P302852

Figured specimen

121

P302854

Time range. Batesfordian-Balcombian, Early

Miocene-Middle Miocene.

Remarks. Chapman (1929) suggested that the

flange was an adaptation to enable the animal to

creep over ‘an even-surfaced oozy sea-bed’. The

specimens from PL3048, Boornong Rd. cutting,

consist of fragments of the flange. It is assumed

that they belong to this species, though the

locality is of Batesfordian age, whereas all the

other specimens have been recorded only from

localities of Balcombian age.

Acknowledgments

1 am grateful to Alan Limpus for providing me

with specimens of Umhilia capricornica and

Umhilia pelilirosiris sp. nov., to Chris Goudey

and Thora Whitehead for the loan of material and

for provision of distributional information, and to

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

371

Ken Bell for collecting material at Batesford

Quarry. Mark Darragh kindly undertook all the

photographic work and Sally Rogcrs-Davidson

and Peter Bubulya scanned the plates. For the

loan of type and other specimens, 1 thank John

Cooper. Natural History Museum, London; Ben

McHenry and Wolfgang Zeidlcr, South Aus¬

tralian Museum; George Kendrick, Western Aus¬

tralian Museum; Lyn Broadbridge, Geological

Survey of South Australia; and Alan Beu, New

Zealand Institute of Geological and Nuclear Sci¬

ences. Phillip Maxwell generously waived his

interest in describing the Palaeacypraea speci¬

men. 1 thank Kathie Way, Natural History

Museum, London, for photographs of the syntype

of Cvpmea iimhilicatcr. Philippe Bouchet,

Museum National d’llistoirc Naturelle, for pro¬

viding dates of publication; Ronald Janssen,

Senckenburg Museum, for copies of literature

unavailable in Australia; Martin Gomon, Museum

of Victoria, for undertaking X-rays of specimens

for me; and David Holloway, Museum of Victo¬

ria, George Kendrick, Western Australian

Museum, and Barry Wilson for commenting on

the manuscript.

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Pacific Mollusca 1 (8); 457-484.

Figure 1. Sections through protoconchs. Bar scale 1 mm. A, Umbilia (U.) angustior (Pritchard), P304357, Picnic

bed, Hordern Vale. B, Umbilia (U.) eximia (Sowerby), P304355, PL3078, Fossil Beach.

THE AUSTRALIAN GASTROPOD GENUS UMBIUA

373

Figure 2. A-D, 'IValaeocvpraea eripnides sp. nov. A, C, NZGNS TM 8124, hololype, Pitt I., New Zealand, x 1.5.

B D, NZGNS TM 8125,’paratype, Pitt 1., New Zealand, x 1.2. E-F, H-K, Umhilia (U.) platyrhynclici {McCoy). E,

P12138, holotype. Ad 22, Bird Rock cliffs, X 1.0. F, P59169, Ad 22, Bird Rock cliffs, x 1.1. H-l, P302784, Bird

Rock cliffs. X 0.9, X 0.8. J-K, P12I38, holotype. Ad 22, Bird Rock cliffs, X 0.7. G, Umhilia (U.) angustior

(Pritchard), P2671, holotype. Table Cape, Tasmania, x 1.1.

374

THOMAS A. DARRAGH

Figure 3. A-F, Umhilia (U.) platyrhyrtcha (McCoy). A-B, P59169, Ad 22, Bird Rock cliffs, x 0.8. C-D, P302697,

cliffs west of Bird Rock, X 0.8. E, PI2I38, holotype. Ad 22, Bird Rock cliffs, x 0.75. F, P302784, Bird Rock cliffs,

X 0.8. G-l, Umhilia (U.) angtv.v/ior (Pritchard), P302688, PL3032, x 1.0.

Figure 4. A-1, Umhilia (U.) anguslior (Pritchard). A, E, G, P2671, holotype. Table Cape, Tasmania, x 0.9, x 1.0,

X 1.0. B-D, P302703, PL3037, Hordern Vale, x 1.1. F, H-I, P302785, Table Cape, Tasmania, x 1.0.

376

THOMAS A. DARRAGH

Figure 5. A-H, Umhilia (U.) e.ximia (Sowerby). A-B, P5296, syntype of Cypraea eximia brevis McCoy, Aw 9,

Gellibrand River, x 0.9. C, E, G, BMNH G9694, hololype, Franklin Village. Tasmania, X 0.8, x 0.8, X 0.9. D, F, H.

BMNH G40069. holotype of Umhilia (U.) brevis frankslonensis Schilder, Grices Creek, X 0.7, X 0.8, x 0.7.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

377

Figure 6. A-H, Umbilia (U.) eximia (Sowcrby). A, C, P16171, syntype of Umhilia (U.) eximia maccoyi Schilder,

between Mt Eliza and Mt Martha, X 0.7. B, D, G, PI2135, lectotype of Umbilia (U.) eximia maccoyi Schilder,

between Mt Eliza and Mt Martha, x 0.7. E-F, H, P12136, lectotype of Cypraea eximia brevis McCoy, between Mt

Eliza and Mt Martha, x 0.9.

378

THOMAS A. DARRAGH

Figure 7. A, Umhilia (U.) hesitata (Iredale), P30278I, PL 1264, Flinders 1., x 0.75. B-1 Umbilia (U.) eximia

(Sowerby). B, P12135, leclolype of Umbilia (U.) eximia maccoyi Schilder, between Mt Eliza and Mt Martha, x 0.9.

C, P5296, syntype of Cypraea eximia brevis McCoy, Aw 9, Gellibrand River, x l.l. D-F, SAM T82I, holotype of

Cypraea sphaerodoma Tate, near Morgan, South Australia, x 0.8. G-1, BMNH 70406, holotype of Umbilia (U.)

brevis monlismarlbae Schilder, Mount Martha. X 1.0.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

379

Figure 8. A-H, Umhilia (U.) eximia (Sowerby). A-B, P3027I2, Lake Bullenmerri, X 1.0. C, P302779, lower beds,

Griees Creek, X 0.8. D-E, H, P302782, PL3298, x 0.8, x 0.9, X 0.9. F-G, P302716, Rose Hill, x 1.0, x 0.9.

THOMAS A. DARRAGH

380

Figure 9. A-H, Umhilia (U.) hesitata (Iredale). A. C, E, WAM 89.636a, 2.5 km N of Hampton Tower, Roe Plains,

Western Australia, x 0.9, x 0.8, x 0.9. B, D, F, WAM 89.636b, 2.5 km N of Hampton Tower, Roe Plains, Western

Australia, x 0.8, x 0,9, x 0.9. G-H, P302717, Rose Hill, X 0.9, x 0.8.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

381

Figure 10. A-1, UnthUia (U.) hesilala (Iredale). A, C, SAM T8I2B, syntypc o\' Cypraea lalei Cossniann, Murray

Desert, x 1.1. B, D, SAM P8339, holotype of Umbilia cera Cotton, Abattoirs Bore, South Australia, x 1.1. E, 1,

P3027I8, Rose Hill, x 1.1. F, SAM T812A, lectotype of Cypraea lalei Cossmann, Murray Desert, x 1.1. G-H,

P302720, Beaumaris, X 1.1.

Figure 11. A-H, Umhilia (U.) hesitata (Iredale). A-C, P302770, PL 1250, Flinders Is, Tasmania, x 1, x 1.1, x 0.9,

D-E, P302780, North Memana Drain, Flinders 1., Tasmania, x 0.8. F-Fl, P302781, PL 1264, Flinders Is, Tasmania.

xO.8.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

383

Figure 12. A, C, Umhilia (U.) hesitata (Iredale), ‘beddomei’ form, F23116, 50 fm, off Botany Bay, New South

Wales, X 1.0, X 0.9. B, D-E, Umhilia (U.) armeniaca (Verco), F27194, Great Australian Bight, uncoated, X 0.7,

X 0.8, X 0.8. F-H, Umhilia (U.) hesitata (Iredale), F4526, off Lakes Entrance, Victoria, X 0.8, x 0.7, x 0.7.

384

THOMAS A. DARRAGH

Figure 13. A-C, Umbilia (U.) petilirosiris sp. nov., F86962, holotype, Capricorn Channel, off Lady Musgrave I.,

Queensland, 264-252 metres. A, C x 1.1, D x 1.0. D-E, Umbilia (U.) hesitata (Iredale), BMNH 1950.8.28.22,

lectotype, X 0.9. ,

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

385

Figure 14. A-C, Umhilia (U.) hesitata (\Kda\e). A, C, SAM T8I2A, lectotype of Cypmea latei Cossmann, Murray

Desert, x l.l. B, SAM P8339, holotype of Umhilia cera Cotton, Abattoirs Bore, x l.l. D-H, Umhilia (U.) capri-

cornica Lorenz. D-E, F85330, 108-110 fm, Swains Reef, Queensland, x 1.2. F-G, F85329, 103-105 fm, Swains

Reef, Queensland, x 1.0.

386

THOMAS A. DARRAGH

Figure 15. A-C, E, H, Umbilia (U.) prosila sp. nov. A. H, P308717, paratype. Bird Rock cliffs, Torquay, x 1.4.

B-C, E, P308716, hololype. Bird Rock cliffs, Torquay, x 1.4. D, F-G, Umbilia (U.) petilirosiris sp. nov. F86963.

paratype, 249 m. off Fiztroy Reef, D x 0.8, F-G x 0.9.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

387

Figure 16. A-H, Umhitia (U.) leptorhyncha (McCoy). A, D, P308728, paralectotype, near Mount Martha, x 1.2.

B-C, E-F, PI2133, lectotype, B x I.l, C X 1.0, E x 0.9, Fx 1.2, G-H, P308726, paralectotype, near Mount Martha,

X 1.2.

388

THOMAS A. DARRAGH

Figure 17. A-E, Umbilia (U.) siphonala (Chapman). A-B, P26904, holotype of Gisortia breviplicata Schilder,

Muddy Creek, x 0. 6. C, PI4835, Grices Creek, X 0.6. D-E, P302803, Muddy Creek, x 0.8.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

389

Figure 18. A-F, Umhilia (U.) siphonala (Chapman). A-C, PI4835, Grices Creek, x 0.6. D-F, PI2569, Muddy

Creek. X 0.6, X 0.7, X 0.6.

390

THOMAS A. DARRAGH

Figure 19. A-B, Umhilia (U.) siphoiuila (Chapman), PI3243, holotype, Murray River cliffs near Waikerie, South

Australia, X 0.7. C, Umhilia (U.) hesitata (Iredale), SAM T812B, syntype of Cypraea tatei Cossmann, Murray

Desert, x 1.1. D-E, Umhilia (Palliocypraea) gaslroplax (McCoy), P12140, holotype, between Mt Eliza and Mt

Martha, x 0.8.

THE AUSTRALIAN GASTROPOD GENUS UMBILIA

391

Figure 20. A-D, Umhilia (Palliocypraea) gastroplax (McCoy). A-B, P302854, Balcombe Bay, X 0.7. C-D,

P302852. Clifton Bank, x 0.8.

392

THOMAS A. DARRAGH

Figure 21. A-D, Umbilia (Palliocypraea) gasiroplax (McCoy). A-B, PI3373, Clifton Bank, x 0.8, C-D, P24869,

Balcombe Bay, x 0.8, x 0.7.

H-

Memoirs of the Museum of Victoria 59(2): 393-437 (2002)

THE AUSTRALIAN SPECIES OF CHIMARRA STEPHENS

(TRICHOPTERA: PHILOPOTAMIDAE)

David 1. Cartwright

13 Brolga Crescent, Wandana Heights, Victoria 3216, Australia

Abstract

Cartwright, D.I., 2002. The Australian species of Chimarra Stephens (Trichoptera:

Philopotamidae) Memoirs of Museum Victoria 59(2): 393^37.

Descriptions, keys and a checklist are provided for males of 26 species of the cosmopolitan

caddisfly genus Chimarra (Philopotamidae), including 23 new species from Australia.

Females of 22 species are also keyed and described.

Introduction

The widespread caddisfly genus Chimarra

Stephens, 1829 is one of the largest genera in the

order Trichoptera with about 400 species

described, but with numbers of species approach¬

ing 600 with works in progress (Blahnik, 1997;

Morse, 1999). The genus Chimarra is diverse,

common and widespread in faster flowing areas

of rivers and streams in Australia, although only

four species have been described previously,

C. atisiralis Navas, 1923, C. australica Ulmer,

1916, C. monticola Kimmins, 1953 and

C. iiranka Mosely, 1953. In reference to C. aus¬

tralis Navas, described only from a female,

Kimmins slated that he was “unable to recognize

this species from the description” (Kimmins in

Mosely and Kimmins, 1953: 404); and the

whereabouts of the holotype female is

unknown (Neboiss, 1988: 212). This species is

not considered further in this paper.

Cartwright (1990) described and figured

females of C. australica and C. monticola from

Victoria. Wells (1991) partly figured four species

Chimarra in a key to Northern Territory (Alli¬

gator Rivers Region) Trichoptera. Wells and

Cartwright (1993) listed four species of Chimarra

amongst Trichoptera collected from Cape York

Peninsula, north Queensland. Walker ct al.

(1995) listed nine species of Chimarra, including

six undescribed species, from the Queensland

Wet Tropics. Cartwright (1997) listed 24 Chi¬

marra ‘species' in a checklist, and noted that the

only record of Chimarra from Tasmania was a of

single male of C monticola.

In this taxonomic revision of the Australian

Chimarra nearly 9400 male and female

specimens were examined and referred to 26

species. Half of these specimens belong to

C. Iiranka, while another 20% were identified as

C. karakara sp. nov. Distributions of Australian

species are summarised in Table I. The break¬

down in distribution of Australian species in each

province and region is: two (endemic) species

recorded from the Eyrean province (Pilbara

region of North Western Australia, N-WA); a

total of 21 species recorded within the Torresian

province of northern Australia, including seven

species from the Kimberley region of N-WA,

nine species from the northern part of the North¬

ern Territory (N-NT), 11 species in northern

Queensland (N-Qld); and five species in the

Bassian province of southeastern Australia (SE-

Qld, New South Wales, Victoria and Tasmania)

(Table 1). Chimarra is recorded from all

Australian states and territories except South

Australia and the Australian Capital Territory and

is also not recorded from southwestern Australia.

Most of the Chimarra species described are from

the warmer northern part or Torresian province of

Australia. Neboiss and Wells (1998) found a sim¬

ilar preponderance of northern species in the

Australian leptocerid genus Triaenodes, and sug¬

gested that Triaenodes may be a relatively recent

arrival in Australia. Chimarra is well represented

in the tropical and subtropical areas of Asia and

Africa, and the genus as a whole can be described

as warm adapted (Blahnik, 1998). The distribu¬

tion of Chimarra differs from that of the other

Australian philopotamid genus Hydrobiosella

Tillyard, 1924, which is most common in south¬

eastern Australia, particularly Tasmania and also

occurs in southwestern Australia. The ‘southern’

distribution of Hydrobiosella is more indicative

of an older Gondwanan origin.

393

394

D. CARTWRIGHT

Table 1. Eyrean (western-central), Torresian (northern) and Bassian (south-eastern) species of Aus¬

tralian Chimarra.

Eyrean species (Pilbara, northwestern WA)

C. luminaris, C. yoolumba

Torresian species (Kimberley, north-western WA, NT, northern Qld)

C. adalutna, C. akruiia, C. bibaringa. C. bungoona, C. kaiya, C. karakara, C. larapinta. C. locolo,

C. mouldsi, C. nabilla, C. natalicia, C. neboissi. C. onimbera, C. pillara, C. pita. C. raniika,

C. stclainie, C. wooroonooran. C. yandala

Bassian species (southeastern Australia including Tasmania)

C. kewarra. C. monticola, C. tallawalla

Species common to both Bassian and Torresian provinces (eastern and northern Australia)

C. aiistralica (mainly Bassian), C. uranka (mainly Torresian)

Ross (1956) and Blahnik (1997, 1998) have

published studies on the phylogeny and bio-

geography of various groups and subgroups of

Chimarra. Ross (1956) studied species of Chi¬

marra from higher altitudes, although he noted

that Chimarra is not primarily a montane group.

He stated that from an Asiatic ancestor, a group

spread throughout .Asia and to Australia and

Africa, where offshoot lines occur. Blahnik

(1998 ) in a study of Neotropical Chimarra. sug¬

gested that New World Chimarra may represent a

monophylctic group with several lineages, and

that a phylogenetic analysis of Old World species

is required to resolve unanswered questions.

Blahnik (1998) postulated a common northern

South America-Africa origin before continent

separation to explain the Old and New World

distribution of Chimarra.

Common but often variable key characteristics

of the Australian fauna include; eastern Aus¬

tralian species with colour uniformly brown to

almost black and northern Australian species with

pale yellowish head, often with a brownish trian¬

gular area between the ocelli and darker wings

and body; wings usually unicolorous, length of

forewing in males 3.7-8.5 mm, females usually

slightly larger; forewing commonly with Rs

curved basal to discoidal cell, which often has an

enlarged node or thickened veins; hindwing with

or without fork I, including at least one species

where this character seems to be variable within

the species; male genitalia with the ventral pro¬

cess on segment IX usually short, keel-like;

tergum X usually mostly membranous mesally,

not divided, with mesal or lateral lobes or pro¬

cesses; inferior appendages variable, usually

short, in some species more elongate; phallus

often with obvious embedded or projecting phal¬

lic or endothecal spines; female genitalia usually

short with segment Vlll membranous dorsally;

sternum IX with paired membranous or partly

sclerotised plates; cerci usually short. Size, body

and wing colour can be useful characters, but are

variable. Colour should be considered with

caution since the colour often fades with time in

alcohol. The Austndian species can be placed in

groups based mainly on male gcnitalic characters.

Some species arc placed in groups which may be

superficially similar but not necessarily natural or

monophylctic. The groups can be characterised as

follows: species with a ‘pale window’ (an area of

semi-transparent membrane) in the wings -

C. luminaris and C. locolo-, species with hindwing

with fork 1 present, relatively elongate inferior

appendages and elongate lateral processes of ter¬

gum X usually with distinct ‘barb-like projec¬

tions’ without apparent sensilla - C. kaiva,

C. bungoona. C. larapinta, C. onimbera, C. pil¬

lara and C yandakr. species with a relatively

elongate ventral process projecting between bases

of the inferior appendages - C. natalicia and

C. neboissi-, species with a single large dorsal

phallic projection or apicodorsal extension of the

phallotheca — C. ranuka and C. uranka-, species

with a pair of elongate phallic spines -

C. tallawalla and C. wooroonoonan; species with

one or two pairs of pigmented mesal processes on

tergum X (and/or tergum IX) and dorsal sclcro-

tised ‘hood-like’ projection on the phallus -

C. adaluma, C. yoolumba, C. nabilla, C. akriina

and C. pita-, unifomily dark species without the

above characters but with relatively slender

upturned inferior appendages - C monticola.

C. aiistralica and C. kewarra-, and the rest (prob¬

ably not a natural group), again darker species

typically with shorter inferior appendages -

C. mouldsi, C. bibaringa, C. karakara and

C. stclairae.

Most of the material studied was made

available by Dr Arturs Neboiss. Depositories for

specimens are abbreviated as follows: Museum

Victoria, Melbourne (NMV), Australian National

Insect Collection, Canberra (ANIC), the

Natural History Museum, London (BMNH),

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

395

Naturhistoriska Riksmuseet, Stockholm (NRS),

Museum and Art Galleries of the Northern Terri¬

tory, Darwin (NTM) and the Queensland

Museum. Brisbane (QM). All specimens, includ¬

ing types, mentioned in the text are lodged in the

NMV unless stated otherwise.

Males and females of each species are most

readily distinguished by genitalic features, often

requiring clearing of the abdomen in potassium

hydroxide. Females were paired with respective

males on the basis of similarities in coloration,

particularly on the head, and on wing venation

and locality and some were associated by rearing

out from larv'ae or pupae.

Figured specimens are identified by the note¬

book numbers of Dr Arturs Neboiss (NMV), pre¬

fix PT-; or the author; prefix CT-. Terminology

used generally follows that of Nielsen (1957,

1981), and Blahnik (1998). Abbreviations for

genitalic parts are indicated on Figs 2-4 (male)

and 80-81 (female) and additionally where neces¬

sary. Typically, setae or spines are illustrated only

on the right side of the figure (as viewed) to

enable a better view of the underlying structures.

Names of prolific collectors have been abbrevi¬

ated in the text as follows: J.E.Bishop - JEB; J.

Blyth - JB; P. Dostine - PD, M.S. Moulds -

MSM; A. Neboiss - AN; P. Suter - PS; A. Wells

-AW.

Chimarra Stephens

Chimarra Stephens, 1829: 318.—Mosely and Kim-

mins. 1953: 398.

Type species. Phryganea imirginalaC. Linnaeus, 1767,

by monotypy.

Diagnosis. A revised diagnosis of the genus Chimarra

was provided recently by Blahnik (1998: 14).

Key to males and females of species of Chimarra from Australia

1. Males.2

— Females.27

2. Forewing with ‘pale window’ (Figs I, la).‘C. himinaris-groxip'... 3

— Forewing without ‘pale window’.4

3. Inferior appendages in lateral view, sub-rectangular, not tapered apically

(Fig. 2); hindwing without ‘pale window’ (Fig. I) N-WA (Pilbara).

.C Iiiminaris

— Inferior appendages in lateral view, not subrectangular, tapered apically

(Fig. 5); hindwing with ‘pale window’ (Fig. la)N-NT......C locolo

4. Hindwing usually with fork 1 present; lateral processes of tergum X elon¬

gate, usually with at least I ‘barb-like projection’, usually longer than infe¬

rior appendages; inferior appendages relatively long with small digitiform

projection apically (Figs 8, 11, 17).‘C. Aa/y«-group’ ... 5

— Hindwing usually with fork 1 absent; lateral processes of tergum X , if pre¬

sent, usually short without ‘barb-like projections’, usually shorter than

inferior appendages; inferior appendages usually short, robust (Figs 26, 35)

.10

5. Mesal process of tergum X single in dorsal view (Figs 10,13, 16, 19).6

— Mesal processes of tergum X paired in dorsal view (Figs 22,25).8

6. Inferior appendages, in lateral view, angled strongly near midventral margin;

lateral processes of tergum X with 2 small subapical ‘barb-like projections’

(Figs 8, 10) N-NT, N-Qld.C. kaiya

— Inferior appendages, in lateral view, not angled strongly near mid ventral

margin; lateral processes of tergum X without or with I subapical ‘barb-like

projection’ or only weakly developed (Figs 11, 13, 14, 16, 17, 19).7

7. Phallus with pair of projecting endothecal spines obvious dorsally (Fig. 13);

lateral processes of tergum X with relatively large apical hook and subapical

‘barb-like projection’ (Figs II, 13) N-NT.C. hungoona

— Phallus without pair of projecting endothecal spines apparent dorsally (Figs

16, 19); lateral processes of tergum X without relatively large apical hook

or ‘barb-like projections’, paired processes not apparent or only weakly

developed (Figs 14, 16, 17, 19).8

D. CARTWRIGHT

Lateral processes of tcrgum X with paired processes weakly developed and

without dorsal branch (Figs 14, 16); hindwing with fork I present, N-WA....

. C. lanipinici

Lateral processes of tergum X without paired processes but with distinctive

dorsal branch (Figs 17, 19); hindwing with fork 1 absent, N-WA.

. C. oriimbeni

Lateral processes of tergum X with relatively large upturned apical hook

(Fig. 20); paired mesal processes of tergum X relatively short (Fig. 22)

N-WA.C. pillara

Lateral processes of tergum X without upturned apical hook (Fig. 23);

paired mesal processes of tergum X relatively long (Fig. 25) N-Qld.

.C. ycmdctla

Ventral process of segment IX relatively elongate, acute, projecting between

bases of inferior appendages (Figs 26, 27,29.30)....‘C. natalicia-group'... 11

Ventral process of segment IX relatively short, obtuse, not projecting

between bases of inferior appendages (Figs 32, 33, 35, 36).12

Inferior appendages in ventral view, with digitifonn apical mesal projection,

narrowly separated from the subapical angle, forming a deep notch (Fig. 27)

N-Qld. C. natalicia

Inferior appendages in ventral view, without digitifonn apical mesal projec¬

tion, but with basal mesal projection widely separated from the apical angle,

forming a wide notch (Fig. 30) N-Qld.C. neboissi

Phallus with a single projecting, elongate, dorsal ‘phallic projection' or

‘apicodorsal extension of the phallotheca’ (Figs 32, 32a, 34, 35, 35a, 37).

.‘C. iiranka-gmup' ... 13

Phallus with paired projecting ‘phallic spines’ or spines not apparent (Figs

38,38a, 40,41,41a, 43).14

Mesal processes of tergum X relatively short, not reaching apex of dorsal

‘phallic projection’ (Figs 32, 34) Qld, N-NT, N-WA.C. iiranka

Mesal processes of tergum X relatively long, reaching apex of dorsal

‘phallic projection’ (Figs 35,37) N-NT.C. nvmka

Phallus with paired long and slender ‘phallic spines’ (Figs 38, 38a, 41, 41a)

.‘C. IcillawaUa-group' ... 15

Phallus without paired projecting long and slender ‘phallic spines’ apparent,

single to many endothecal spines may be present (Figs 44, 50).15

Phallus with paired very slender, very elongate ‘phallic spines’ dorsally,

attached near base of phallotheca. often extending past apex of phallus

(Fies 38-^0, 38a), although sometimes withdrawn and so not as apparent

(Fiis 38b. 40a) E-NSW, SE-Qld.C. killawalla

Phallus with paired slender, long ‘phallic spines’ laterally, attached near base

of endotheca, never extending past apex of phallus (Figs 41, 4Ia, 43) N-Qld

.....C. wooroonooran

Mesal processes of tergum X or IX in dorsal view', present as 1 or 2 pairs of

relatively elongate spine-like projections; phallus with dorsal sclerotised

‘hood-like’ projection (Figs 46, 49).‘C culaliima-gtoup' ... 17

Mesal processes of tergum X in dorsal view, not present as I or 2 pairs of

(usually dark) relatively elongate spine-like projections (Figs 61,64).21

Mesal processes of tergum X in dorsal view, present as one pair of spine-like

projections (Fig. 46) N-WA.C. adctluma

Mesal processes of tergum X in dorsal view, present as two pairs of spine¬

like projections (Figs 49. 52. 55).18

Inner pair of mesal processes of tergum X relatively widely separated,

situated adjacent to outer pair (Fig. 49) N-WA (Pilbara).C yoolumba

Inner pair of mesal processes of tergum X not relatively widely separated,

not usually situated adjacent to outer pair (Figs 52, 55, 58).19

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

397

19 .

20 .

21 .

22 .

23.

24.

25.

26.

27.

28.

29.

30.

31.

Inner pair of mesal processes of tergum X shorter than outer pair (Fig. 52)

N-WA, N-NT.C. nabilla

Inner pair of mesal processes of tergum X as long as or longer than outer pair

(Figs 55, 58).20

Outer pair of mesal processes of tergum X as long as inner pair (Fig.52)

N-NT. C. aknina

Outer pair of mesal processes of tergum X about two-thirds as long as inner

pair (Fig. 58) N-WA, N-NT.C. pita

Inferior appendages in lateral view, relatively long, tapered gradually api-

cally to acute apices (Figs 59, 62, 65), and in ventral view apices inflexed

(Figs 60, 63, 66).‘C. monlicola-group'...22

Inferior appendages in lateral view, relatively short, not usually tapered

gradually apically to acute apices (Figs 68, 71, 74), and in ventral view

apices not usually inllexed (Figs 69, 72, 75).‘C. mouldsi-group'.... 24

Lateral processes of tergum X not produced into obvious projections (Figs

59, 61) NSW, Vic., Tas.C monticola

Lateral processes of tergum X produced into obvious projections, slightly

hooked apically (Figs 62,64.65.67).23

Lateral processes of tergum X, with relatively short, upturned or out-turned

apices (Figs 62, 63, 64, 64b,c); phallus with a ventral process or

‘phallotremal sclerite' (Figs 62, 63) E-Aust. C. australica

Lateral processes of tergum X with relatively long, upturned or out-turned

apices; phallus without a ventral process (Figs 65,66) SE-Qld.C. kewarra

Inferior appendages in ventral view, subquadrate, truncate apically (Figs 69,

72).25

Inferior appendages in ventral view, not subquadrate, not truncate apically

(Figs 75, 78).26

Inferior appendages in lateral view, subquadrate, truncate apically (Fig. 68)

N-Qld.C. mouldsi

Inferior appendages in lateral view, not subquadrate, not truncate apically

(Fig. 71) N-Qld.C. siclairae

Lateral processes of tergum X apically with obvious elongate processes

ventral to phallus; apex of phallus without obvious dark spines (Figs 74, 75)

N-Qld.C. hiharingci

Lateral processes of tergum X withotit elongate processes ventral to phallus;

apex of phallus with (three) obvious small dark spines (Fig. 77, 78) N-Qld

.C. karakara

Forewing with 'pale window’ (Figs I, la).‘C. /«/)7/7?am-group’....28

Forewing without 'pale window'.29

Sternum IX ventrolatcrally with distinctive ‘knob-like processes’ (Figs 80,

81); hindwing without ‘pale window" (Fig. I) N-WA (Pilbara)...C. hmiinaris

Sternum IX ventrolaterally without distinctive ‘knob-like processes’ (Figs

82, 83); hindwing with ‘pale window’ (Fig. la) N-NT. C. locolo

Ilindwing usually with fork I present; cerci pigmented; genitalia usually

relatively long (Figs 85, 87, 89).‘C. A-i:7n’a-group'....30

Ilindwing usually without fork I present; cerci usually unpigmented;

genitalia relatively short (Figs 95, 97, 99).34

Posteroventral margin of sternum VIII with 2 areas of setae not separated by

small notch (Figs 85, 87). 31

Posteroventral margin of sternum VIII with 2 areas of setae separated by

small notch (Figs 89, 91, 93). 32

Cerci and genitalui relatively elongate (Figs 84, 85); hindwing with fork 1

absent; N-WA, N-NT.C orumhera

Cerci and genitalia not relatively elongate (Figs 86, 87); hindwing with fork

1 present; N-NT, N-Qld.C. kaiya

D. CARTWRIGHT

Posteroventral margin of sternum Vlll with small notch about as long as

wide (Fig. 89); N-NT.C. himgoona

Posteroventral margin of sternum VIII with small notch about twice as long

as wide (Figs 91,93).33

Posterior margin of sternum IX with rounded ‘comers’ (Fig. 91) N-Qld.

.C. yanciala

Posterior margin of sternum IX with angular ‘comers’ (Fig. 93) N-WA.

. C. pillara

Head pale with body and wings usually darker (N-Australia).35

Head dark with body and wings usually dark (E-Australia).39

Sternum IX vcntrally with obvious pigmented areas (Figs 95,97).36

Sternum IX venlrally without obvious pigmented areas (Figs 99, 101).

.37

Sternum IX ventrally with obvious pair of pigmented triangular shaped areas

(Fig. 95) Qld, N-NT, N-WA.C. tiranka

Sternum IX ventrally with pigmented areas not triangular (Fig. 97) N-NT,

N-WA...,...,.C. akruiui

Posteroventral margin of sternum VIII with 2 areas of setae relatively widely

separated (Fig. 99); N-WA, N-NT.C. pita

Posteroventral margin of sternum Vlll with 2 areas of setae relatively

narrowly separated (Figs 101, 103);.38

Posterior margin of .sternum IX rounded (Fig. 101) N-WA (Pilbara).

.C. yooltimba

Posterior margin of sternum IX angular (Fig. 103) N-WA, N-NT.

.C. nahilla

Po.steroventral margin of sternum VIII with 2 areas of setae separated

bynotch (Fig. I05)N-Old.C. natalicia

Posteroventral margin of sternum Vlll with 2 areas of setae not separated by

notch (Figs 107, 109).40

Sternum IX ventrolaterally with pair of‘pockets’ (Figs 106. 107) E-NSW,

SE-Qld.C. tallawalla

Sternum IX ventrolaterally without pair of‘pockets’ (Figs 109, 111, 113)...

Po-steroventral margin of sternum Vlll with pair of pigmented sclerites

joined to form ‘T-shaped’ pigmented area (Fig. 109) N-Qld.C. nehoissi

Posteroventral margin of sternum Vlll with pair of pigmented sclerites not

joined to form ‘T-shaped’ pigmented area (Figs 111,113).42

Sternum IX ventrally without obvious pair of pigmented areas (Fig. Ill)

N-Qld...C. wooraonoonan

Sternum IX ventrally with obvious pair of pigmented areas or depressions

(Figs 113, 115).43

Sternum IX ventrally with pair of small squarish depressions (Fig. 113)

N-Qld. C. siclairae

Sternum IX ventrally without pair of small squarish depressions (Figs 115,

117). 44

Sternum IX ventrally with obvious pair of pigmented areas relatively widely

separated (Figs 115,117).45

Sternum IX ventrally with obvious pair of pigmented areas relatively

narrowly separated (Figs 119, 121).‘C. monticola-group'... 46

Posteroventral margin of sternum Vlll with pair of pigmented sclerites

widely separated; sternum IX ventrally with pair of pigmented areas with

length less than width (Fig. 115) N-Qld.....C. mouldsi

Posteroventral margin of sternum VIII with pair of pigmented sclerites nar¬

rowly separated; sternum IX ventrally with pair of pigmented areas with

length greater than width (Fjg. 117) N-Qld.C. karakara

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

399

46. Sternum IX venlrally with pair of non-triangular pigmented areas or sclerites

(Fig. 119) Vic., E-NSW, E-QId.C. australica

— Sternum IX ventrally with pair of triangular pigmented areas or sclerites

(Figs 121, 123).47

47. Sternum IX ventrally with pair of triangular pigmented areas or sclerites

with length greater than width (Fig. 121) SE-Qld. C. kewarra

— Sternum IX ventrally with pair of triangular pigmented areas or sclerites

with length less than width (Fig. 123) E-NSW, Vic., Tas.C. monticola

Chimarra /Hw/iwom-group

The C. lurninciris-gToup of two species, C. lumi-

naris and C. locolo, is characterised primarily by

forewings with ‘pale window’ near discoidal cell.

Other common but not exclusive characters are

pale head and brownish body and wings, hind¬

wing with fork 1 absent, forewing with vein Rs

thickened and curv'ed slightly basal to discoidal

cell; male genitalia tcrgum X with a pair of

relatively widely separated mesal processes,

phallus with large, protruding ventral or apical

spine(s), ventral process on .segment IX small and

keel-like; female genitalia relatively short and

broad. Both members of this group are from

northern Australia, one each from N-WA and

N-NT.

Chimarra luniiiiaris sp. nov.

Figures 1-4, 80, 81

Chimarra sp. nov. CT-262.—Cartwright. 1997: 17.

Type material. Ilolotype male. Western Australia.

Fortescue R.. Millstream, S of Roeboume, 22 Feb 1977,

M.S. and B.J. Moulds (NMV, T-I7548). Paratypes. 24

males (specimen CT-262 figured). 15 females

(specimen CT-263 figured), collected with holotype

(NMV).

Other material examined Western Australia. 9 males, 2

females. Lockyer Gorge, Harding R., Pilbara, 19 Oct

1979, JB; 24 males. 12 females. Millstream. 21-22 Apr

1972, N.R. Mitchell ( ANIC); I male, 1 female, Fortes¬

cue R., Millstream National Park, 2i°34’S, I I7°03'E,

24-25 Apr 1992, P.S. Cranston; 16 males, 16 females.

Forteseue R.. Millstream. S of Roeboume, 22 Feb 1977,

M.S. and B.J. Moulds; 7 males, 6 females. Crossing

Pool, Millstream, Pilbara. 21 Oct 1979, JB; 2 male

pupae, Pilbara. Ashburton R., Wallbrook Pool,

22°28.54’S, 116°28.05'E, date?, P. Cranston (ANIC); 2

males, 1 female, 28 km NE Tom Price, 22°30'S,

117°58'E, 26 Sep 1995, D.C.F. Rent/ and J. Otto

(ANIC); 7 males, 3 females, Wittenoom Gorge, llamer-

sley Range National Park. 20 Feb 1977, M.S. and B.J.

Moulds; 6 males, I female, Fortescue Falls, flamersley

Range National Park. 27 Oct 1979. JB; I male, llamer-

sley National Park, Fortescue Falls, 23°38 S, I I8°33'E,

23 Apr 1992, Cranston and Gullan (ANIC).

Diagnosis. Chimarra liiminaris resembles C

locolo in possessing ‘pale semi-transparent

window’ in the forewing, but it is distinguished

by the absence of the ‘pale window’ in the

hindwing.

Description. Head pale, body and wings brownish

to dark brown, forewings with ‘pale semi-trans¬

parent window’ near discoidal cell; hindwings

without ‘pale window'; length of forewing: male

5.0-5.4 mm. female 5.3-7.1 mm; wing venation:

forewing with forks I, 2, 3 and 5 present; hind¬

wing with forks 2, 3 and 5 present; in forewing,

vein Rs thickened and curved slightly basal to dis¬

coidal cell (Fig. I).

Male. Ventral process on segment IX small,

keel-like; inferior appendages short, in lateral

view subrectangular, length about 1.5 times width

(Fig. 2), apices slightly intlexed; mesal processes

of tergum X short, usually with bifid apices,

sometimes asymmetrical; lateral processes of ter¬

gum X with 3 small apical lobes; phallus with

robust hooked endothecal spine projecting at apex

and with dorsal sclerotised ‘hood-like’ projection

(Figs 2-4).

Female. Female genitalia relatively short,

broad; sternum VII with small keel-like process.

Posterolateral margin of segment VIII with dark

sclerites and associated setae dorsally, near mid¬

dle and ventrally. Ventral pair of sclerites rela¬

tively widely separated, with connecting posterior

margin of segment Vlll fairly straight and without

notches. Segment Vlll dorsally with membranous

area almost dividing segment. Ninth sternum with

pair of faint, oblong pigmented areas and laterally

with distinctive pairs of knob-like processes. Ter¬

gum X fomiing 2 lobes with numerous setae, each

with short apical cercus (Figs 80, 81).

Etymology^ Luminaris - Latin for window

(wings).

Remarks. Chimarra luminaris has been col¬

lected from live sites in the Pilbara region of

northern Western Australia (latitudinal range

2I°00'-23°38'S).

400

D. CARTWRIGHT

Chimarra locolo sp. nov.

Figures la, 5-7, 82, 83

Chimarra sp. nov. E .—Cartwright, 1997: 17.

Type material. Holotype male. Northern Territory,

Litchfield National Park, UV light, 6 Jun 1991, Wells

and Webber (NMV, T-17588). Paratypes. 21 males

(specimen CT-279 figured), 22 females (specimen

CT-299 figured), collected with holotype (NMV,

NTM).

Other material examined. Northern Territory. I female.

Jim Jim Ck, 3 km below falls. Kakadu National Park, 1

Sep 1979. JB; 1 male. Berry Springs, 12°42'S.

130°58’E, 7 May 1992, A.W. (NTM); 3 males. 2

females, Florence Falls. Litchfield National Park. MV

It, 9 Apr 1991. Horak and Wells (NTM); 7 males, 13

females, Litchfield Park, MV light, 3 Apr 1991,3. Web¬

ber and R. De Jong (NTM); 1 male. Devil Devil Ck, 70

km SW Daly R. Mission, 23 Aug 1979, JB; 1 male,

Katherine R. Gorge National Park, 26 Jan 1977, M.S.

and B.J. Moulds.

Diagnosis. Chimarra locolo resembles C. lumt-

naris in possessing ‘pale semi-transparent

window' in the tbrewing, but it is distinguished

by the presence of the ‘pale window’ in the hind¬

wing.

Description. Mead pale, with brown triangular

area between ocelli, body and wings brow'nish,

fore- and hindw ings with ‘pale window' centred

near discoidal cell; length of forewing: male

4.7-5.2 mm, female 4.9 -6.5 mm; wing venation:

forewing with forks 1, 2, 3 and 5 present; hind¬

wing with forks 2, 3 and 5 present: in forcw'ing,

vein Rs thickened and curved slightly basal to

discoidal cell (Fig.la).

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

broadbased, narrowing strongly at about middle

to slender apices (Fig. 5); mesal pair of processes

of tergum X and IX margin slender, widely

spaced (Fig. 7); phallus robust with asymmetrical

pair of strong, serrate-edged cndothecal spines

projecting ventrally (Fig. 6).

Female. Female genitalia relatively short,

broad; sternum VII with small keel-like process.

Posterolateral margin of segment Vlll with dark

sclerites and associated setae dorsally, near mid¬

dle and ventrally. Ventral pair of sclerites rela¬

tively narrowly separated, with connecting poste¬

rior margin of segment Vlll fairly straight and

without notches. Segment Vlll dorsally with

membranous area almost dividing segment. Ninth

sternum with pair of faint, oblong pigmented

areas. Tergum X comprising 2 setose lobes with

numerous setae, each with short apical cercus

(Figs 82, 83).

Etymology. Locolo - Northern Territory Abori¬

ginal word for circle (wings).

Remarks. This species has been collected

from the northern half of the Northern Territory

(latitudinal range 12°42'-14°29'S).

Chimarra Ara/ra-group

The C. kaiya-gmup of six species, C. kaiya, C.

bungoona, C. larapinta, C. orumhera, C. pillara

and C. yandala, is characterised primarily by

male genitalia with inferior appendages elongate

and slender with an apical finger-like process,

tergum X with elongate lateral processes, often

with ‘barb-like projections'; female genitalia in

all but one species (C. kaiya) relatively elongate

with pigmented cerci. Other common but not

exclusive characters are pale head and brownish

body and wings, wings unicolorous, hindwing

with fork 1 present (except C. orumhera),

(brewing with vein Rs thickened and curved

slightly basal to discoidal cell, male genitalia with

tergum X either with a pair of mesal processes or

single mesal process, phallus with large, often

protruding spine(s), ventral process on segment

IX small and keel-like. All members are from

northern Australia.

Chimarra kaiya sp. nov.

Figures 8-10, 86, 87

Chimarra sp. C.—Wells, 1991: 20.

Chimarra sp. n. C.—Wells and Cartwright, 1993:

227.

Chimarra sp. nov. CT-270.—Cartwright, 1997; 17.

Type material. Holotype male. Northern Territory,

ARRS Radon Springs, It tr.. 13-14 Apr 1988, AW and

PS (NMV, T-17632). Paratypes. 21 males (specimen

CT-270 figured), 6 females (specimen CT-297 figured),

collected with holotype (NMV).

Other material examined. Northern Territory, Kakadu

National Park-Radon Springs. Baroalba Springs.

Koongarra, 15 km E of Mt Cahill, Stag Ck at BIIP

camp, Jim Jim Falls campsite, Litchfield National

Park-Walker Ck, Ada Ck (ANIC, NTM. NMV).

Queensland. Cape York Peninsula-Lockerbie Scrub.

Jardine R.. Heathlands area. Iron Range. Mcliwraith

Range. Mt Webb National Park (ANIC. NMV, QM).

Diagnosis. In general form of male genitalia,

C. kaiya resembles others in the group, but it is

distinguished by the strongly angled inferior

appendages and the lateral processes of tergum X

with 2 subapical ‘barb-like projections’ and no

apical hook.

Description. Head pale, body and wings brown¬

ish; length of forewing: male 4.7-6.1 mm, female

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

401

5.0-6.5 mm; wing venation: Ibrewing with forks

1,2, 3 and 5 present; hindwing with forks I, 2, 3

and 5 present; in forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view

long, length about 3.5 times width, angled

strongly near middle of ventral margin, with

small digitifomi process apically (Fig. 8); single

mesal process of tergum X short, simple in lateral

view (Fig. 10); lateral processes of tergum X

longer than inferior appendages, with 2 small dor¬

sal ‘barb-like projections' subapically and

towards middle, no apical hook; phallus robust

with 2 long slightly curved endothecal spines

projecting at apex (Figs 8, 9).

Female. Female genitalia relatively short,

broad; sternum Vll with small keel-like process.

Posterolateral margin of segment VIII with dark

sclerites and associated setae dorsally, near

middle and ventrally. Ventral pair of sclerites

relatively narrowly separated, with connecting

posterior margin of segment VIII fairly straight

and without notches. Segment VIII dorsally with

a membranous area almost dividing segment.

Ninth sternum without any pigmented areas. Ter¬

gum X fonning 2 lobes with numerous setae, each

with a short, pigmented apical cercus (Figs 86,

87).

Etymology. Kaiya — Queensland Aboriginal word

for spear with two barbs (lateral processes on ter¬

gum X).

Remarks. This is a common and widespread

species across Cape York Peninsula and northern

Northern Territory (latitudinal range 10°46'-

I5°14'S).

Chimarra hungoona sp. nov.

Figures 11-13, 88, 89

Chimarra sp. D.—Wells, 1991: 20,

C/»>«arra sp. nov. CT-272.—Cartwrighl. 1997: 17.

Type material. Ilolotype male. Northern Territory.

Radon Springs. It tr., 14 Apr 1989, AW and PS (NMV,

T-17660). Paratypes. 1 male (CT-272, drawn speci¬

men), Randon (Radon?) Ck, Kakadu National Park, 3

Sep 1979, JB (NMV); 2 females (specimen C'T-298 fig¬

ured), ARRS Radon Springs. It Ir., 18-19 May 1988,

AW and PS (NMV).

Other material examined. Northern Territory, 1 male,

ARRS Radon Springs, It tr., 13-14 Apr 1988, AW and

PS? (NTM); 2 males. Koongarra, 15 km E of Mt Cahill.

I2°52'S, 132°50'E. 12-13 Jun 1973, J.C. Cardale

(ANIC); 2 males, Litchfield National Park. Walker Ck,

UV It, 18-19 Apr 1992, AW (NTM); I male, Litchfield

National Park, Ada Ck at jump up. It tr, 24-25 Jun

1992, Wells and Webber (NTM); 2 males, Umbrawarra

Gorge, 14°00'S, I3I°38'E, MV It, 23 Aug 1982, J. and

I. Archibald (ANIC).

Diagnosis. This species closely resembles others

in the group in general form of male genitalia, but

it is distinguished by the combination of a single

mesal process on tergum X and lateral processes

of tergum X with subapical ‘barb-like projection’

and apical hook.

Description. Head pale with brown triangular

area between ocelli, body and wings brownish;

length of forewing: male 4.9-5.2 mm, female

4.9-6.3 mm; wing venation: forewing with forks

1, 2, 3 and 5 present; hindvving with forks I, 2, 3

and 5 present; in forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view

long, length about 3.5 times width, not angled

strongly near middle of ventral margin, with

small digitifomi process apically (Figs II, 12);

single mesal process of tergum X short, slender

(Fig. 13), simple in lateral view'; lateral processes

of tergum X longer than inferior appendages, w'ith

relatively large dorsal ‘barb-like projection’ sub¬

apically and apical hook; phallus robust with pair

of endothecal spines projecting dorsally and a

single spine projecting apically (Figs II, 13).

Female. Female genitalia relatively long, elon¬

gate; sternum VII with small keel-like process.

Posterolateral margin of segment VIII w'ith dark

sclerites and associated setae dorsally, near

middle and ventrally. Ventral pair of sclerites

relatively narrow'ly separated, with connecting

posterior margin of segment VIII fairly straight

and with small notch. Segment VIII dorsally with

a membranous area almost dividing segment.

Ninth sternum without any pigmented areas. Ter¬

gum X forming 2 lobes with numerous setae, each

with relatively long, pigmented apical cercus

(Figs 88, 89).

Etymology. Bungoona - Australian Aboriginal

word for sandy creek (typical habitat).

Remarks. Chimarra hungoona is known from

five sites in the northern Northern Territory

(latitudinal range I2°45'-14°()0'S).

Chimarra larapinta sp. nov.

Figures 14-16

Chimarra sp. nov. U.—Cartwright, 1997: 17.

Type material. Holotype male. Western Australia,

Marun' CALM site 8/4, Prince Frederick Harbour,

I5°00'S, 125°2rE, at light, 6-11 Jun 1988, I.D.

402

D. CARTWRIGHT

Nauniann (ANIC). Paralype. 1 male (specimen CT-308

figured), collected with holotype (ANIC).

Other material examined. Western Australia. 1 male, nr

Mitchell Plateau airfield. 14°48'S, 125°49'E, at light,

15 May 1983, J.C. Cardale (ANIC); 2 males, 1 female.

Carson Escarpment. 14“49'S. 126°49’E, Drysdale 1975

Survey site Bl, 9-15 Aug 1975, I.F.B. Common and

M.S. Upton (ANIC); I male (specimen CT-289. geni¬

talia missing). ‘Marun’ CALM site 8/4. Prince Freder¬

ick Harbour, 15'^00'S. l25'’2rFL at light. 6-11 Jun

1988. l.D. Naumann (ANIC); 7 males. CALM site 28/3.

4 km W of King Cascade, 15°35'S, I28°I5'E. 17-20

Jun 1988, T.A. Weir (ANIC).

Diagnosis. Chimarra larapinta closely resembles

others in the group in general form of male geni¬

talia, but it is distinguished by the combination of

a single mesal process on tergiim X and lateral

processes of tergum X with paired subapical

‘projections' only weakly developed.

Description. Head, body and wings pale; length

of forewing: male 4.6-4.9 mm; wing venation:

forewing with forks 1. 2. 3 and 5 present; hind¬

wing with forks 1,2. 3 and 5 present: in forewing,

vein Rs thickened and curved slightly basal to

discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

long, length about 3.5 times width, slightly

narrowed but not angled strongly near middle of

ventral margin, with small digitiform process

dorsoapically (Figs 14, 16); single mesal process

of tergum X short, triangular (Fig. 16), simple in

lateral view (Fig. 14); lateral processes of tergum

X longer than inferior appendages, with paired

subapical projections weakly developed laterally

(Fig. 16); phallus robust with short curved ventral

endothecal spine visible subapically (Figs 14, 15).

Female unknown.

Enmolog}-. Larapinta - Australian Aboriginal

word for flowing water (typical habitat).

Remarks. This species has been collected from

four sites in the Kimberley region of northern

Western Australia (latitudinal range 14°48'-

I5'’35'S).

Chimarra onimhera sp. nov.

Figures 17 19, 84, 85

Chimarra sp. nov. CT-266.—Cartwright, 1997: 17.

Type material Holotype male. Western Australia, Kim¬

berley, Prince Regent R., 15°47'S. 125°24’E, May

1985. E. Bloomfield (NMV, T-17664). Paratypes.

Western Australia. 5 males, collected with holotype

(NMV); 1 male (specimen CT-266 figured), Barnett R.

Gorge. Barnett Station, Kimberley, 1 Oct 1979, J.B.

(NMV); 2 males, same data (NMV); I male, 1 female

(specimen CT-316 figured). Manning R. nr Mt Barnett,

16°40-S. 125‘’.56 E, 8 Sep 1996, I. Edwards (NMV); 1

female, Mitchell Plateau. Lone Dingo Ck, trib. of

Mitchell R.. 17 Feb 1979, J.E.B. (NMV).

Other material e.xamined. Western Australia. 2 males, 1

female. CALM site 13/4, 12 km S of Kalumburu Mis¬

sion. I4°25'S, I26°38'E. 7-11 Jun 1988, T.A. Weir

(ANIC); 1 male. ■Martin’ CALM site 8/4, Prince Fred¬

erick Harbour. 15°0(I'S, 125°2I’E, at light. 6-11 Jun

1988, l.D. Naumann (ANIC); 11 males, C.ALM site

28/3. 4 km W of King Cascade. 15°35'S. 128°15 E,

17-20 Jun 1988. T.A. Weir (ANIC); I male. El Questro

Station, Chamberlain R. pool. 15‘’58'S. 127“56 E, 26

Aug 1996, I. Edwards; 2 males. Backsten Ck. S of

Prince Regent Rcserse, UTOO'S, I25°29 E, 6 Sep

1996.1. Edwards; I female. Drv.sdale R. headwaters, 30

km NW Ml Elizabeth HS. 30 Sep 1979. JB; 4 males. 1

female. Drysdale R., upper reaches. I6°09’S, 125°58'E,

7 Sep 1996. 1. Edwards; 1 male, I female. Adcock

Gorge. Gibb R.-Derby Rd, Kimberley. 2 Oct 1979, JB:

I male. Bell Gorge, Melalcuka Hole. n^Ol'S,

125° 14'E. 13 Sep 1996. I. Edwards; I female. King

Edward R., 14°54'S. I26°12 E. 3 Sep 1996.1. Edwards.

Diagnosis. Chimarra onimhera is distinguished

in the group by the absence of both fork I on the

hindwing and 'barb-like projections’ on the lat¬

eral processes of tergum X. The lateral processes

of tergum X have a distinctive dorsal branch.

De.scription. Head pale yellow with brown trian¬

gular area between ocelli, body brown, wings

dark brown; length of forewing: male 3.7-4.7

mm, female 4.2-5.0 mm; wing venation:

forewing with forks 1, 2, 3 and 5 present; hind¬

wing with forks 2, 3 and 5 present: in forewing,

vein Rs thickened and curved slightly basal to

discoidal cell.

Male. Ventral process on segment IX small,

kcel-like; inferior appendages in lateral view,

long, length about 3.0 times width, slightly

narrowed but not angled strongly near middle of

ventral margin, with small digitifomi process

apically (Fig. 17); mesal process of tergum X

broad (Fig. 19), simple in lateral view (Fig. 17);

lateral processes of tergum X with dorsal branch

joined basally, ventral branch about as long as

inferior appendages, slightly laterally (lattened.

without ‘barb-like projections’, dorsal branch

slightly shorter and more slender (Figs 17, 19);

phallus robust with long endothecal spine

embedded subapically (Figs 18, 19).

Female. Female genitalia relatively long, elon¬

gate; stenium Vll with kcel-like process. Postero¬

lateral margin of segment Vlll with dark scleritcs

and associated setae dorsally, near middle and

ventrally. Ventral pair of sclerites relatively

narrowly separated, with connecting posterior

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

403

margin of segment VIII fairly straight and without

small notch. Segment VIII dorsally with a mem¬

branous area almost dividing segment. Ninth ster¬

num without any pigmented areas. Tergum X

forming 2 lobes with numerous setae, each with

relatively long, pigmented apical cercus (Figs 84,

85).

Etymology. Orumhera - Australian Aboriginal

word for large spear (genitalia).

Remarks. Chimarra orumhera is known only

from the Kimberley region of northern Western

Australia (latitudinal range 14°25'-I7°0rS).

Chimarra pillara sp. nov.

Figures 20-22, 92, 93

Chimarra sp. nov. T.—Cartwright. 1997: 17.

Type material, llolotype male, We.stcm Australia, ‘The

Crusher’ CALM site 9/1. 4 km S by W minimi camp.

Mitchell Plateau. 14°52'S. I25“50-E, 2-6 Jun 1988,

I.D. Naumann(ANIC). Paratypes. Western Australia. 4

males 10 km W by N of mining camp. Mitchell

Plateau. I4“45’S. 125°47'E. at light. II May 1983. J.C.

Cardale (ANIC); 4 males. I female, nr Mitchell Plateau

airfield. 14°48 S, I25'’49’E, at light, 15 May 1983, J.C.

Cardale (ANIC); 1 male (specimen CT-288 figured), 1

female (specimen CT-327 figured) ‘Marun’ CALM site

8/4. Prince Frederick Harbour, IS^OO'S. I25'’2I'E, at

light, 6-11 Jun 1988, I.D. Naumann (ANIC); I male ,1

female. Mining camp, Mitchell Plateau, 14°49 S,

I25‘’5()'E, 9-19 May 1983, I.D. Naumann and J.C.

Cardale (ANIC); I male, 2 females, 4 km S by W min¬

ing camp, Mitchell Plateau, at light, I4°52'S, 125°50'E.

13 May 1983, J.C. Cardale (ANIC); 1 male, ‘The

Crusher’ CALM site 9/1. 4 km S by W mining camp,

Mitchell Plateau. I4°52’S. 125“50'E, 2-6 Jun 1988,

ID. Naumann (ANIC); 1 male, Augustus Island.

CALM site 26/1. I5°25'S, I24°38'E, 11-16 Jun 1988,

1 D Naumann (ANIC); 3 males, I female, CALM site

25/1, Synnot Ck, 16°3rS, I25°I6'E, 17-20 Jun 1988,

T.A. Weir (ANIC).

Other material examined Northern Territory. 1 male. 8

females (specimen CT-327 figured), Litchfield Park,

MV light, 3 Apr 1991, J. Webber and R. De Jong

(NTM); I male. 7 females, Litchfield National Park.

UV light, 6 Jun 1991, Wells and Webber (NTM).

Diagnosis. This species is distinguished from oth¬

ers in the group by the combination of relatively

short paired mesal processes on tergum X and the

lateral processes of tergum X with subapical

‘barb-like projections’ and well developed apical

hook.

Description. Head, body and wings pale; length

of forewing: male 4.6-5.2 mm, female 5.2-5.6

mm; wing venation; forewing with forks 1, 2, 3

and 5 present; hindwing with forks 1, 2, 3 and 5

present: in forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

long, length about 3.5 times width, not angled

strongly near middle of ventral margin, with

small digitifonn process apically (Figs 20, 21);

pair of mesal processes of tergum X short

(Fig. 22), simple in lateral view (Fig. 20); lateral

processes of tergum X longer than inferior

appendages, slightly laterally flattened, with

relatively large ‘barb-like projection’ subapically

and large apical hook (Figs 20, 22); phallus long

with pair of robust endothccal spines visible

dorsally (Figs 20, 22).

Female. Female genitalia relatively long, elon¬

gate; sternum VII with a keel-like process.

Posterolateral margin of segment VIII with dark

sclerites and associated setae dorsally, near

middle and ventrally. Ventral pair of sclerites

relatively narrowly separated, with connecting

posterior margin of segment VIII with small

notch. Segment VIII dorsally with membranous

area almost dividing segment. Ninth sternum

without any pigmented areas, posterior margin

with angular ‘corners’. Tergum X fomiing 2 lobes

with numerous setae, each with relatively long,

pigmented apical cercus (Figs 92, 93).

Etymology. Pillara - Aboriginal word for spear

with two barbs (lateral processes on tergum X).

Remarks. This species is known from the Kim¬

berley region of northern Western Australia and

Litchfield National Park in the Northern Territory

(latitudinal range l4°45'-16°3rS). It has been

collected from April to June.

Chimarra yaiulala sp. nov.

Figures 23-25, 90, 91

Chimarra sp. n. G.—Wells and Cartwright. 1993:

227.

Chimarra sp. nov. CT-271.—Cartwright, 1997: 17.

Type material. I lolotype male, Queensland. Gordon Ck,

Iron Range, 16 Apr 1975. Moulds (NMV, T-17676).

Paratypes. I male (specimen CT-271 figured) same

locality, 10 Jun 1975. MSM (NMV); 2 males, 3

females, .same locality, 12 May 1975, MSM (NMV); 1

male. I female (specimen CT-301 figured), same local¬

ity, 18 Apr 1975, Moulds (NMV); 2 females, Gordon

Ck, Iron Range. 2 Jun 1975, MSM (NMV).

Other material examined. Queensland. I male, I

female. Burster Ck, KISS'S, 142°40'E. at light, 17 Oct

1992. P. Zborowski and T. Weir (ANIC): 1 male. Canal

Ck u/sjn Eliot Ck, UV It, ir23’S, I42“25'E, 6 Feb

1992, Cartwright and Wells (QM); 2 females, Canal Ck

u/sjn Eliot Ck, UV It, 1 l“23'S, 142°25'E, 6 Feb 1992.

404

D. CARTWRIGHT

Cartwright and Wells (QM): 1 t'emale, Dulhunty R.,

Telegraph Crossing, I TSO'S, I42°30'E, 10 Feb 1992.

Cartwright and Wells (QM); 1 male, 14 km ENE Heath-

lands, by hand, rainforest, I l'’4rS, 142°42'E, 28 Feb

1993. P. Zborowski (ANIC); I male. Dividing Range,

25 km W of Capt Billy Landing. I l°40'S, I42°45’E.

4-9 Jul 1975, G.R. Monteith (ANIC); I male. 9 km

ENE Mt Tozer. MV It, 12°43'S, 143“I7 E. 5-10 Jul

1986, J.C. Cardale (ANIC); 1 male. 2 females. Middle

Claudio R., Iron Range, 2-9 Oct 1974. Moulds; I male,

Archer R. crossing. Cape York Peninsula, 9 Sep 1974.

MSM; 4 males. 3 km NE Mt Webb. 15‘’03 S. I45'’09 E,

1-3 Oct 1980, J.C. Cardale (ANIC); 1 male, 7 km N of

Hopevale Mission, 15“I4 S. 145°07 E, 4 Oct 1980. J.C.

Cardale (ANIC); 5 males. 1 km N of Rounded Hill nr

Hopevale Mission. 15=I7'S. I45°13 E, 5-6 Oct 1980,

J.C. Cardale (ANIC); I male. 5 km W by N of Rounded

Hill nr Hopevale Mission. I5°17’S, I45°10'E, 7 Oct

1980. J.C. Cardale (ANIC); 1 male, same locality,

15°17 S. 145°10 E, 7 Oct 1980. D.H. Collcss (ANIC);

3 males, Mt Cook National Park. Cooktown. I5“29'S.

145°I6 E, 11-12 Oct 1980. J.C. Cardale (ANIC).

Diagnosis. Chimarra yandala is distinguished

from others in the group by the combination of

paired mesal processes on tergum X and the elon¬

gate and slender lateral processes of tergum X

with one subapical ‘barb-like projection'.

Description. Head pale with brown triangular

area between ocelli, body and wings brownish;

length of forew'ing; male 5,0-5.4 mm, female

4.9-6.2 mm; wing venation: forewing with forks

1,2,3 and 5 present; hindwing with forks 1,2,3

and 5 present; in forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

long, length about 3.5 times width, not angled

strongly near middle of ventral margin, with

small digitiform process apically (Figs 23, 25);

pair of mesal processes of tergum X relatively

long (Fig. 25). simple in lateral view (Fig. 23);

lateral processes of tergum X longer than inferior

appendages, slender, with I relatively small

‘barb-like projection’ subapically (Figs 24. 25);

phallus with short, robust endothecal spine visible

on dorsal margin subapically (Figs 23, 25).

Female. Female genitalia relatively long, elon¬

gate; stemum VI1 with elongate keel- like pro¬

cess. Posterolateral margin of segment VIII with

sclerites and associated setae near middle and

ventrally. Ventral pair of sclerites relatively nar¬

rowly separated, with connecting posterior mar¬

gin of segment Vlll with small notch. Segment

Vlll dorsally with membranous area almost divid¬

ing segment. Ninth stemum without pigmented

areas, posterior margin with rounded ‘corners’.

Tergum X forming 2 lobes with numerous setae.

each with relatively long, pigmented apical cercus

(Figs 90,91).

Etymology. Yandala - Queensland Aboriginal

word for spear with long point (lateral processes

on tergum X).

Remarks. This appears to be a common species on

Cape York Peninsula, north-eastern Queensland

(latitudinal range 10°55'-15°29'S).

Chimarra natalicia-gmup

The C. natalicia-group of two species. C. nalali-

cia and C. nehoissi, both characterised primarily

by male genitalia with ventral process on segment

IX which is relatively elongate and acute. Other

common but not exclusive characters are dark

head, body and wings, wings unicolorous,

forewing with vein Rs thickened and curved

slightly basal to discoidal cell, male genitalia with

tergum X without any apparent mesal processes

but with short, broad lateral processes, phallus

with more than one endothecal spine, inferior

appendages short; female genitalia relatively

short and broad. Both members of this group are

from north-east Queensland.

Chimarra natalicia sp. nov.

Figures 26-28, 104, 105

Chimarra sp. nov. CT-221.—Walker et al.. 1995:

26.—Cartwright. 1997: 17.

Tvpe material. Holotvpe male. Queensland. Birthday

Ck, 3.5 km WNN Paluma, I8°59'S. ITh^lO'E, al light.

8 Oct 1989. R. St Clair (NMV. T-17688). Paratypes. 14

males. 16 females (specimen CT-309 figured), col¬

lected with holotype(NMV); 1 male (specimen CT-221

figured), same locality, 1 Apr 1990, R. St Clair (NMV):

2 males, same locality, 17 Mar 1990, R. St Clair

(NMV).

Other material examined. Queensland. 1 male, I

female, Windsor Tableland, 20 Feb 1982, MSM; 2

males, 5 females. Upper Freshwater Ck, Whitfield

Range nr Cairns, 15 Dee 1974, Moulds; 1 male. Upper

Freshwater Ck. Whitfield Range nr Cairns, MV-lighl.

24 Aug 1974, Moulds; 1 male, 1.5 km SE Kuranda.

16-17 May 1980, I.D. Naumann and J. Cardale

(ANIC); 4 males. Bellenden Ker Range, Cableway

Ba.se Sin. 100 m, 17 Nov 1981, Earthwatch/Qld

Museum: I male. I female, Bellenden Ker Range.

Cableway Base Stn, 100 m. 17-24 Oct 1981, Earth-

watch/Qld Museum; 1 male, Koolmoon Ck. Atherton

Tablelands, site KM I, 27 Nov 1990, S. Bunn and M.

Gray; 1 male. Tully Falls, S of Ravenshoe, 11 Jan 1977.

M.S. and B.J. Moulds; 2 males, 1 female. Birthday Ck.

3.5 km WNN Paluma. I8°59 S, I46''I0'E, al light, 17

Feb 1990, R. St Clair; 1 female. Paluma, 27 Jan 1982.

MSM; 1 female. Birthday Ck below falls, Mt Spec State

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

405

Forest, 18°57'S, I46°I0'E, 760 m. It. tr.. 17 Mar 1994.

A.L. Sheldon; 2 males, 4 Icmales, same site, 29 Mar

1994- 7 females. Echo Ck tributary, Mt Spec State For¬

est, I8°57'S, I46°I0'E, 735 m. It. tr., 7 Nov 1993, A.L.

Sheldon; I female. Echo Ck, Mt Spec State Forest,

I8°57'S, I46°I0'E, 735 m. It. tr., 5 Mar 1994, A.L.

Sheldon; 1 female, Echo Ck. Mt Spec State Forest,

I8°57'S, I46°I0'E, 700 m. It. tr., 17 May 1994, A.L.

Sheldon; 1 female. Running River, Mt Spec State For¬

est, 18°57 S, 146°10’E. 640 m. It. tr., 16 Mar 1994,

A.L. Sheldon.

Diagnosis. Males of this species resemble those

of C. nehoissi having the ventral process on seg¬

ment IX relatively elongate, acute, projecting

between bases of inferior appendages, but differ

in that the apices of the inferior appendages are

turned mesally and extended into a digitiform

process, which is narrowly separated from the

subapical angle, forming a deep notch.

Description. Head brown, body and wings light

brown; length of forewing: male 4.8-5.4 mm,

female 5.7-6.4 mm; wing venation; forewing with

forks I, 2, 3 and 5 present; hindwing with forks 2,

3 and 5 present. In forewing, vein Rs thickened

and curved slightly basal to discoidal cell.

Male. Ventral process on segment IX relatively

elongate, acute, projecting between bases of infe¬

rior appendages (Figs 26, 27); inferior

appendages in lateral view, very short, robust,

length about 1.2 times width (Fig. 26), in ventral

view stout, with apices turned mesally and

extended into a digitifomi process, which is nar¬

rowly separated from the subapical angle, form¬

ing a deep notch (Fig. 27); mesal processes of

tergum X not apparent, lateral processes of ter-

gum X robust, inflected apically; phallus robust

with many small endothecal spines embedded

subapically (Figs 26-28).

Female. Female genitalia relatively short,

broad; sternum VII with a keel-like process. Pos¬

terolateral margin of segment VIII with scleritcs

and associated setae dorsally, near middle and

ventrally. Ventral pair of sclerites relatively nar¬

rowly separated, with connecting posterior mar¬

gin of segment VllI with a large notch. Segment

VIII dorsally with membranous area almost divid¬

ing segment. Ninth sternum with 2 lightly pig¬

mented areas. Tergum X fomiing 2 lobes with

numerous setae, each with relatively short apical

cercus (Figs 104, 105).

Etymology. Natalicia - Latin word for birthday

(type locality Birthday Creek).

Remarks. Chimarra natalicia is a common north

eastern Queensland species (latitudinal range

I6°10'-I9°00'S).

Chimarra nehoissi sp. nov.

Figures 29-31, 108, 109

Chimarra sp. nov. CT-223.—Walker et al., 1995:

26.—Cartwright, 1997: 17.

Type material. Holotype male, Queensland, Little Mul-

grave R.. 9 km SW ofGordonvale, 17 Nov 1979, A.N.

(NMV, T-17722). Paratypes. 7 males (specimen CT-

223 figured), 5 females (specimen CT-324 figured),

collected with holotype (NMV).

Other material examined. Queensland. Mt Webb

National Park, llopevale Mission, Cooktown area,

Whitfield Range nr Cairns, Mareeba district, nr Gor-

donvale. NW of Tully, Kirrama State Forest, Mt Spec

State Forest (ANIC, NMV).

Diagnosis. Males of Chimarra nehoissi resemble

those of C. natalicia having the ventral process

on segment IX relatively elongate, acute, project¬

ing between bases of inferior appendages, but

differ in that the apices of the inferior append¬

ages are separated widely from the basal angle,

fomiing a wide notch. The wing venation in

some specimens appears variable in that fork 1 in

the hindwing is sometimes missing or not

apparent.

Description. Head, body and wings dark brown to

blackish; length of forewing: male 4.5-5.3 mm,

female 5.1-5.9 mm; wing venation; forewing

with forks 1, 2, 3 and 5 present; hindwing with

forks 1, 2, 3 and 5 present. In forewing, vein Rs

thickened and curved slightly basal to discoidal

cell.

Male. Ventral process on segment IX relatively

elongate, acute, projecting between bases of

inferior appendages (Figs 29, 30); inferior

appendages in lateral view, short, tapering

slightly apically, length about twice wddth (Fig.

29), in ventral view stout, with apices separated

w'idely from basal angle, fomiing a w'ide notch

(Fig. 30); mesal processes of tergum X not appar¬

ent, lateral processes of tergum X short, robust,

situated below phallus; phallus robust with two

long slender endothecal spines embedded sub¬

apically (Figs 29-31).

Female. Female genitalia relatively short,

broad; sternum VII with a small keel-like process.

Posterolateral margin of segment VIII with scle¬

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclerites joined

forming a central 'T-shaped' pigmented area.

Segment VI11 dorsally with membranous area

almost dividing segment. Ninth sternum without

any lightly pigmented areas. Tergum X forming 2

lobes with numerous setae, each with relatively

short apical cercus (Figs 108, 109).

406

D. CARTWRIGHT

Etymology. Nehoissi - named after Dr Arturs

Neboiss (collector).

Remarks. Chimarra neboissi is a common dark

species known from north-eastern Queensland

(latitudinal range 15°04'-18°57'S). Superficially

C. neboissi males can be confused with another

very common north-eastern Queensland dark

species, C. karakara, but the species can be dis¬

tinguished by the length of the ventral process on

segment IX.

Chimarra uranka-group

The C. uranka-group of two species, C. itranka

and C. raniika, is characterised primarily by male

genitalia with phallus with large dorsal phallic

projection or ‘apicodorsal extension of the phal-

lothcca'. Other common but not exclusive charac¬

ters are brownish body and wings, wings uni-

colorous, forew'ing with vein Rs thickened basal

to discoidal cell, in males tergum X with pair of

mesal processes, phallus with one apical spine,

inferior appendages in lateral view, broadbased

and tapered apically, ventral process on segment

IX small and keel-like. Both are from northern

Australia.

Chimarra iiranka Mosely

Figures 32-34, 94, 95

Chimarra uranka Mosely in Mosely and Kimmins,

1953: 399, fig. 273.—Neboiss, 1986: 106.

Type material. Holotype male, Queensland,

Kuranda.IlOO ft, 21 Jun-24 Jul 1913, R. E. Turner

(BMNH). Type not seen.

Material e.xaminecl. Queensland. 1 male (specimen CT-

290 figured), I female (specimen CT-293 figured ), Irib.

Bertie Ck. 250 m SW Heathlands H.S., 11 Feb 1992,

Cartwright and Wells (QM); Cape York Peninsula,

Cairns and Townsville areas, near Monto, Goomeri and

Benarkin.

Northern Territory, Cape Crawford, Amhemland,

Kakadu, Litchfield and Katherine Gorge National

Parks. Victoria River Downs.

Western Australia. Kimberley-Mitchell Plateau.

Lake Argyle area. Bungle Bungle National Park, near

Halls Creek. Giekie Gorge and Tunnel Ck.

Diagnosis. Chimarra uranka most closely resem¬

bles C, ratuika from which it is distinguished by

the relatively short pair of mesal processes of ter¬

gum X and the dorsoventrally Battened apex of

the dorsal phallic projection.

Description. Head orange-yellow, body and

wings dark brownish; length of forewing: male

4.2-6.0 mm, female 4.4-7.0 mm; wing venation:

forewing with forks 1, 2, 3 and 5 present; hind¬

wing with forks 2, 3 and 5 present. In fore¬

wing, vein Rs thickened slightly basal to discoidal

cell.

Male. (Revised after Mosely in Mosely and

Kimmins, 1953). Ventral process on segment IX

small, keel-like; inferior appendages in lateral

view, relatively robust, tapering slightly apically,

length about twice width, upturned (Fig. 32), in

ventral view, directed slightly dislally (Fig. 33);

pair of mesal processes of tergum X relatively

short (Fig. 34), pair of lateral processes of tergum

X short, robust (Figs 33, 34); phallus robust with

single, short, upturned cndothccal spine visible

apically, dorsally with single, slightly dorsoven¬

trally llatlened elongate dorsal phallic projection,

with simple apex (Figs 32. 32a, 34).

Female. Female genitalia relatively short,

broad; sternum VII with keel-like process.

Posterolateral margin of segment VIII with scle-

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclerites relatively

narrowly separated, with connecting posterior

margin of segment Vlll straight and without

notch. Segment Vlll dorsally with membranous

area almost dividing segment. Ninth sternum with

2 triangular lightly pigmented areas. Tergum X

forming 2 lobes with numerous setae, each with

relatively short apical cercus (Figs 94, 95),

Remarks. Chimarra uranka is an extremely com¬

mon and widespread species throughout northern

Australia (latitudinal range 10°48’-26°53'S).

Mosely’s, (in Mosely and Kimmins, 1953) figures

have been redrawn to allow direct comparisons

and to accompany the description that is revised

in light of new interpretations of Chimarra

genitalic structures.

Chimarra raniika sp. nov.

Figures 35-37

Type material. Holotype male. Northern Territory,

Litehfield National Park, UV light, 6 Jun 1991. Wells

and Webber (NMV, T-17735). Paraiypes. 2 males

(specimen CT-291 figured), collected with holotype

(NMV); 2 males, same locality, 3 Apr 1991, J. Webber

and R. de Jong (NTM).

Diagnosis. This species most closely resembles

C. uranka from which it is distinguished by the

relatively long pair of mesal processes of tergum

X and the concave apex of the dorsal phallic pro¬

jection.

De.scription. Head, body and wings brownish;

length of forewing: male 4.3 5.1 mm; wing vena¬

tion; forewing with forks I, 2, 3 and 5 present;

hindwing with forks 2, 3 and 5 present. In

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

407

forewing, vein Rs thickened and curved slightly

basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view, rel¬

atively slender, tapering slightly apically, length

about 3.0 times width, upturned (Fig. 35). in ven¬

tral view, directed slightly distally (Fig. 36); pair

of mesal processes of tergum X long, slender

(Fig. 37), pair of lateral processes of tergum X

robust, apices slightly turned outwards (Figs

35-37); phallus robust with single endothecal

spine embedded apically, dorsally with single,

slender, elongate phallic projection with concave

ape.x (Figs 35, 35a, 37).

Female unknown.

Etymolog)’. Rcintika - anagram of uranka.

Remarks. Chimarra ranuka appears to be

restricted in distribution and probably is rare as it

has been collected from the type locality only

(latitude 13°28'S).

Chimarra /fl//flH’«//ff-group

The Chimarra tallawal/a-group of two species,

C. lallawalla and C wooroonoonan. charac¬

terised primarily by male genitalia with phallus

with long phallic (or endothecal) spines. Other

non-exclusive characters are dark head, body and

wings, wings unicolorous, forevving with vein Rs

not thickened and straight basal to discoidal cell,

in males tergum X with at most a small pair of

mesal or lateral processes, inferior appendages in

lateral view, slender and uptumed, ventral process

on segment IX small and keel-like. Both arc from

eastern Australia.

Chimarra tallawalla sp. nov.

Figures 38-40, 106, 107

Chimarra sp. nov. CT-224.—Cartwright, 1997: 17.

Type nuUeriai. llolotypc male. New .South Wales,

Cangi Ck. NW of Grafton. 29°34'S. 152°26 E. 15 Jan

1985. O. Theischinger (NMV, T-17738). Paratypes. 15

males (specimen CT-224 llgurcd), 18 females (speci¬

men CT-3()4 llgured), collected with holotype (NMV).

Other malerial examined Queensland. I male pupa,

Nerang R.. nr Advancetown, 8 Feb 1973. collector?; 1

male, Mt Superbus, 22 Jan 1971. S.R. Monteith

(ANIC); 5 males, 2 females, Wallangarra, Qld-NSW

border, 31 Dec 1956, J.K. New South Wales. 1 female.

Korumbyn Ck, 28°24 S. 153''19 E. E slope of Mt

Warning, 29 Jul 1988, AN; 2 females. UnderclilTe

Falls, 25 km E of Stanthorpe, II Oct. 1973, AN; 17

males, 20 females, Clarence R. at Yates Crossing, 26

Oct 1981. Wells and Carter; 34 males, 20 females,

Cangi Ck, NW of Grafton, 29°34’S, 152°26'E, 15 Jan

1985. Theischinger; 17 males, 19 females, Boyd R., 5

km E of Dalmerton on old Grafton Rd, 27 Oct 1981,

Wells and Carter; I male, Fembrook nr Dorrigo, 17 Dec

1975, A.B. Prose?; 2 males, Bellinger R., 30°26'S,

I52°44'E.31 Oct 1981, Wells and Carter; 6 males, Styx

R.. 12 km S of Ebor. 17 Oct 1973, AN; 12 males, 18

females, Styx R. at Forest Camp in Styx River State

Forest, 28 Oct 1981, Wells and Carter; 3 males, Apsley

R. at falls, SE of Walcha. 29 Oct 1981, Wells and

Carter; 8 males, 41 females. Wilson R., NW of Wau-

chopc. 3I°I4’S, 152°34'E. 30 Oct 1981, Wells and

Carter; I male. 8 females. Duffers Ck, 9 km WSW Bar¬

rington. 16 Feb 1980, A. Calder; 33 males, 55 females.

Gloucester R., 9 km SW of Gloucester, 17 Feb 1980, A.

Calder; 2 females, U. Allyn R.. 12 km N of Eccleston,

21 Feb 1980, A. Calder; 2 males, 3 females, Tuglo' 48

km N of Singleton, 3 Feb 1979, Smithers; 2 males. 6

females. Tuglo' 48 km N of Singleton, alt. 780m. 5 Oct

1975, MSM; 34 males, 58 females. Allyn R.. NE of

Eccleston, 3 Nov 1981. Wells and Carter; 3 males, 32

females. Allyn R. nr Allynbrook, 21 Mar 1973, AN; 2

male pupae, Tuross R., 3 Feb 1977, J. Dean.

Diagnosis. Males ot' Chimarra tallawalla resem¬

ble those of C. wooroonooran, but dilTer in

having the phallus with paired, very slender, very

elongate phallic (or endothecal) spines dorsally,

attached near base of phallotheca, often extending

past apex of phallus.

Description. Head, body and wings brown; length

of forewing: male 5.1-6.3 mm, female 5.4-7.5

mm; wing venation: forewing with forks 1, 2, 3

and 5 present; hindwing with forks 2, 3 and 5 pre¬

sent. In forewing, vein Rs thickened and curved

slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

short, slender, length about 4.0 times width,

uptumed slightly (Fig. 38). in ventral vievy, short,

broadbased (Fig. 39); mesal processes of tergum

X not apparent (Fig. 40), pair of lateral processes

of tergum X short. robu.st, situated below phallus

(Figs 38, 39); phallus relatively slender with pair

of very slender, very elongate phallic (or endo¬

thecal) spines dorsally, attached near base ol

phallotheca (Figs 38-40, 38a), although some¬

times withdrawn and so not as apparent (Figs

38b. 40a).

Female. Female genitalia relatively short,

broad; sternum VII with small keel-like process.

Posterolateral margin of segment Vlll with scle-

rites near middle and groups of setae ventrally.

Ventrally posterior margin of segment Vlll

straight and without notch. Segment Vlll dorsally

with membranous area almost dividing segment.

Ninth sternum with 2 pockets laterally. Tergum X

forming 2 lobes with numerous setae, each with

relatively short apical cercus (Figs 106, 107).

408

D. CARTWRIGHT

Etymology'. Tallawalla - NSW Aboriginal word

for forest country (habitat).

Remarks. Chimaira tallawalla is a common but

distinctive species found in eastern New South

Wales and southeastern Queensland (latitudinal

range 28°02'-36°06'S).

Chimarra wooraonooran sp. nov.

Figures 41-43, 110. Ill

Chimarra sp. nov. CT-228.—Walker el al., 1995:

26.—Cartwright, 1997: 17.

Type material. Ilolotype male (specimen CT-228 fig¬

ured), Queensland, Bellenden Ker Range. 0.5 km S

Cable Tower No. 7, 500m. 25-31 Oct 1981. Earth-

watchj'Qld Museum (NMV, T-17904). Paralype. 1

female, collected with holotype (specimen CT-314

figured. NMV).

Diagnosis. This species differs from C. tallawalla

in having the phallus with paired slender, long

phallic (or endothccal) spines laterally, attached

near base of endotheca, never extending past apex

of phallus.

Description. Head, body and wings brown; length

of forewing; male 4.2 mm, female 5.0 mm; wing

venation: forewing with forks 1, 2, 3 and 5 pre¬

sent; hindwing with forks 2, 3 and 5 present. In

forew'ing, vein Rs not thickened but curved

slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

slender, length about 3.0 times width, upturned,

tapering slightly apically (Fig. 41), in ventral

view, broadbased, apices turned slightly mesally

(Fig. 42); pair of short mesal processes of tergum

X pale, not obvious, lateral processes of tergum X

not apparent; phallus robust, with pair of long and

slender phallic (or endothecal) spines embedded

laterally, apices turned slightly outwards, and

with about 5 small dark endothccal spines

embedded subapically (Figs 41-43, 41a).

Female. Genitalia relatively short, broad; ster¬

num VII with small keel-like process. Posterolat¬

eral margin of segment VIII with scleritcs and

associated setae dorsally and near middle. Ven-

troposterior margin of segment VIII with a num¬

ber of long setae, relatively straight and without

notch. Segment VI11 dorsally with membranous

area almost dividing segment. Ninth sternum

without any apparent pigmented areas. Tergum X

forming 2 lobes with numerous setae, each with

relatively short apical ccrcus (Figs 110, 111).

Etymology. Wooraonooran ~ Queensland Aborigi¬

nal word for Bellenden Ker Range (type locality).

Remarks. Chimarra wooroonoonan is known

from only one male and one female specimen

from the type locality only in northeastern

Queensland (latitude 17°16'S).

Chimarra adaliima-group

The C. acJahima-gtoup of live species.

C. adaluma. C. yoohimba. C. nabilla. C. akruna

and C. pita, is characterised primarily by males

with tergum X and/or IX with one or two pairs of

slender and elongate mesal processes, and phallus

with a dorsal sclerotised ‘hood-like' projection.

Other common but not exclusive characters are

body and wings brownish, wings unicolorous.

forewing, with vein Rs thickened and curved

slightly basal to discoidal cell, inferior

appendages in lateral view, broadbased and

upturned apically, phallus with strong ventral

endothecal spine or ventral process, ventral pro¬

cess on segment IX small and keel-like. All mem¬

bers of this group are from northwestern Australia

(N-WA and N-NT).

Chimarra adaluma sp. nov.

Figures 44-46

Chimarra sp. nov. CT-268.—Cartwright, 1997; 17.

Type material. Holotype male. Western Australia, Kim¬

berley. Prince Regent R.. King Cascade, MV It, 27 Jul

1990, D.K. Yeates (NMV, T-17773). Paratypes. 14

males (specimen CT-268 figured), collected with holo¬

type (NMV).

Other material examined. Western Australia. 22 males.

CALM site 28/3, 4 km W of King Cascade. I5°35'S,

I28“I5'E. l7 -20Jun 1988, T.A. Weir(ANIC).

Diagnosis. Chimarra adaluma is distinguished

from others in the group by the I pair of curv ed

spine-like mesal processes of tergum X and the

lateral pair of processes of tergum X with apices

almost meeting ventral to the phallus.

Description. Head pale, body and wings light

brown; length of forewing: male 4.8-5.5 mm;

wing venation; forewing with forks 1, 2, 3 and 5

present; hindwing with forks 2, 3 and 5 present. In

forewing, vein Rs thickened and curved slightly

basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

robust, subquadrate, length about 1.1 times width

(Fig. 44), in ventral view, length about 2.5 times

width, with apices directed slightly mesally (Fig.

45); pair of mesal processes of tergum X or IX

slender, curved (Fig. 46), pair of lateral processes

of tergum X project below phallus; phallus

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

409

relatively robust with single, slender, ventral

endothecal spine and dorsal sclerotised slightly

flanged ‘hood-like’ projection (Figs 44-46).

Female unknown.

Etvmolog}’. Atlaliima - Australian Aboriginal

word for river (habitat).

Remarks. Chimarra adaluma has been found

at only two sites in the Kimberley region of

north Western Australia (latitudinal range

15°35'-15°38'S).

Chimarra yoohunha sp. nov.

Figures 47^9, 100, 101

Chimarra sp. nov. CT-264.—Cartwright, 1997; 17.

Type material. Ilolotype male. Western Australia.

Fortescue Falls, llamersley Range National Park. 27

Oct 1979, J. B. (NMV, T-17788). Paratypes. 5 males

(specimen CT-264 figured), 5 females (specimen CT-

265 figured), collected with holotype (NMV).

Other material e.uiminecl. Western Australia. 3 males,

Hamersley National Park. Fortescue Falls, 23°38'S,

I I8°33'F.. 23 Apr 1992, Cranston and Gullan (ANIC).

Diagnosis. This species most closely resembles

C. nahilla in general fonn of male genitalic struc¬

tures, but is distinguished by having the inner pair

of mesal processes oftergum X relatively widely

separated, situated adjacent to outer pair.

Description. Head pale brown, body and wings

brownish; length of forewing: male 5.3-5.7 mm,

female 6.6-7.1 mm; wing venation: forewing

with forks I, 2, 3 and 5 present; hindwing with

forks 2, 3 and 5 present. In forewing, vein Rs

thickened and curved slightly basal to discoidal

cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

robust, length about twice width, upturned

slightly (Fig. 47), with apices turned slightly

mesally and e.xtcnded into a digitiform process

(Fig. 49), in ventral view, length about twice

width (Fig. 48): 2 pairs of dark mesal processes of

tergum X, slender, straight, of similar length,

individual process of each pair widely separated,

but inner and outer processes adjacent (Fig. 49),

lateral processes of tergum X not apparent; phal¬

lus relatively robust, with ring of about 6 dark,

stout endothecal spines subapically and dorsal

sclerotised ‘hood-like’ projection (Figs 47, 48).

Female. Female genitalia relatively short,

broad; sternum VII with keel-like process. Pos¬

terolateral margin of segment VIII with sclerites

and associated setae dorsally, near middle and

ventrally. Ventral pair of sclerites relatively nar¬

rowly separated, with connecting posterior mar¬

gin of segment VIII straight and without notch.

Segment VIII dorsally with membranous area

almost dividing segment. Ninth sternum without

any lightly pigmented areas. Tergum X fomiing 2

lobes with numerous setae, each with relatively

short apical cercus (Figs 100, 101).

Elymolog)\ Yoolumha — Western Australian

Aboriginal word for the Fortescue River (type

locality).

Remarks. This species is known from the type

locality only in the Pilbara region, north Western

Australia (latitude 23°38'S).

Chimarra nabilia sp. nov.

Figures 50-52, 102, 103

Chinuaro sp. nov. CT-267.—Cartwright, 1997: 17.

Type material. Holotype male. Western Australia, Kim¬

berley, Prince Regent River. King Cascade, MV light,

27 Jul 1990. D.K. Yeates (NMV, T-17799). Paratypes.

Western Australia. 3 males (specimen CT-267 figured),

collected with holotype (NMV); 1 male, Backsten Ck.

S of Prince Regent Reserve, 16°00'S, I25°29'E, 6 Sep

1996,1. Edwards (NMV); 2 males, 2 females (specimen

CT-325 figured), Mitchell Plateau, Lone Dingo Ck.

trib. of Mitchell R., 17 Feb 1979. J.E.B. (NMV); 3

males, 'The crusher’, CALM site 9/1,4 km S by W

mining camp, Mitchell Plateau, I4°52’S, 125°50’E. at

light. 2-6 Jun 1988, 1. D. Naumann (ANIC).

Other material e.xamined. Northern Territory. 29 males,

42 females. Berry Springs, MV light, 26 Jun 1991,

Wells and Webber (NTM); 4 males. Berry Springs,

I2°42 S. 130°58’E, 7 May 1992, Wells (NTM). West¬

ern Australia. 3 males, nr Mitchell Plateau airfield,

I4°48-S, 125°49 E, at light. 15 May 1983. J.C. Cardale

(ANIC).

Diagnosis. Chimarra nahilla most closely resem¬

bles C yoolumha in general fonn of male geni¬

talic structures, but is distinguished by having two

pairs of dark, relatively stout mesal processes of

tergum X, inner pair of processes adjacent and

slightly shorter than outer pair which are situated

slightly ventrally with respect to inner pair.

Description. Head pale with brownish triangular

area anterior to ocelli, body and w'ings brownish;

length of forewing: male 4.0-5.2 mm. female

4.6-6.0 mm; wing venation: forewing with forks

I, 2, 3 and 5; hindwing with forks 2, 3 and 5. In

forewing. discoidal cell with distal veins thick¬

ened and Rs thickened and slightly sinusoidal dis¬

tal to discoidal cell.

Male. Ventral process on segment IX small.

410

D. CARTWRIGHT

keel-like; inferior appendages in lateral view,

robust, length about 1.5 times width, upturned

slightly (Fig. 50), with apices turned slightly

mesally and e.xtcnded into digitifomi process

(Fig. 52), in ventral view, length about 2.5 times

width (Fig. 51); 2 pairs ofdark mesal processes of

tergum X, relatively stout, slightly curved, inner

pair of processes adjacent and slightly shorter

than outer pair (Fig. 52). which are situated

slightly ventrally with respect to inner pair

(Fig. 50), lateral processes of tergum X not appar¬

ent; phallus relatively robust with ring of about

5 dark, stout endothecal spines subapically and

1 apically and with pale dorsal sclerotised

‘hood-likc' projection (Figs 50-52).

Female. Female genitalia relatively short,

broad; sternum Vll with keel-like process. Pos¬

terolateral margin of segment Vlll w ith sclerites

and associated setae dorsally, near middle and

ventrally. Ventral pair of sclerites relatively nar¬

rowly separated, with connecting posterior

margin of segment Vlll relatively straight and

without notch. Segment Vlll dorsally with mem¬

branous area almost dividing segment. Ninth ster¬

num without any lightly pigmented areas. Tergum

X forming 2 lobes with numerous setae, each with

relatively short apical cercus (Figs 102, 103).

Etymology. Nahilla - Australian Aboriginal word

for water.

Remarks. Chimarra nahilla is a slightly variable

species with male specimens from the Northern

Territory differing from the type material, mainly

in the positioning and angle of the spines on the

phallus. Known from the Kimberley region of

northern Western Australia and Berry Springs,

northern Northern Territory (latitudinal range

I2°42'-16°00'S).

Chimarra akrana sp. nov.

Figures 53-55. 96, 97

Chimarra sp. B.— Wells, 1991: 20.

Chimarra sp. nov. CT-269.—Cartwright. 1997: 17.

Type material, tiolotype male. Northern Territory,

Radon Springs, It ir.. 14 Apr 1989, AW and I’S (NMV.

T-17808). Paratypes. 13 males (specimen CT-294 fig¬

ured). 6 females (specimen CT-295 figured), collected

with hololype (NMV).

Other material examined Northern Territory. 1 male,

E. Alligator R. at Cahills Crossing, It tr., 27 May 1988,

AW and PS; 5 males. 5 females (including 3 copulating

pairs). ARRS Radon Springs. It tr.. 13-14 Apr 1988.

AW and PS; 4 males. 1 female, same locality. It tr., 14

Apr 1989, AW and PS; 4 males, 1 female, ARRS Radon

Springs, It tr., I2°45'S, 132“55'E, 13-14 Apr 1989,

AW and PS; 2 males, ARRS Radon Springs, It tr.,

12°45'S, 132°55’E, 13-14 Apr 1989, AW and PS; 7

males, 4 females, ARRS Radon Springs, It Ir.. 13-14

Apr 1988, AW and PS?; 2 males, ARRS Radon

Springs, It tr., 14 Apr 1989. AW and PS; 24 males. 27

females. ARRS Radon Springs, It tr.. 18-19 May 1988,

AW and PS; 3 males. Randon (Radon?) Ck, Kakadu

National Park 3 Sep 1979. JB; 2 males, 1 female. ARRS

Bowerbird Billabong outlet. I2°47 S. I32°02'E. 1 Oct

1988, PD; I male, ARRS Baroalba Springs. 22 May

1988, AW and PS; many males, females. Baroalba

Springs, Kakadu National Park, UV, 12°48'S,

132’49’E, 16 .Ian 1992, Wells (NTM); many males,

females, same locality. 4 Mar 1992, Wells (NTM); 1

male, I female, Jim Jim Ck. 3 km below falls. Kakadu

National Park, 1 Sep 1979, JB; 1 male. 2 females.

Graveside Gorge, It trap. 18 Jul 1988; P.D.; 12 males, 8

females. ARRS Graveside Gorge. It trap. 18 Jul 1988,

PD; 3 male.s, 7 females. Graveside Ck, It trap, 18 Jul

1988. PD; 4 males, 1 km below Twin Falls. Kakadu

National Park. It tr., 30 Aug 1979. JB; 12 males. 5

females, ARRS ck 5 km W of OSS, 13°33'S. 132°34E

19 Apr 1989. AW and PS; 2 males, 1 female. ARRS

St.ig Ck at BHP camp. It Ir., 25 May 1988. AW and PS;

I male, ARRS S Alligator R.above Fisher Ck jn. It tr.,

19-20 Apr 1989. AW and PS; 3 male.s. Devil Devil Ck,

70 km SW Daly R. Mission, 23 Aug 1979, JB.

Diagnosis. Chimarra akrtina closely resembles

C pita in general form of male genitalic struc¬

tures, but is distinguished by having 2 pairs of

dark, long and slender mesal processes of tergum

X, all of similar length, individual processes of

inner pair adjacent, outer processes slightly more

widely separated.

Description. Head pale with brownish triangular

area anterior to ocelli, body and wings brownish;

length of forewing: male 4.5-5.5 mm, female

4.9-5.7 mm; wing venation: forewing with forks

1.2, 3 and 5 present; hindwing with forks 2, 3 and

5 present. In forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

robust, length about twice width, upturned and

tapering slightly apically (Fig. 53), in ventral

view, length about 2.5 times width (Fig. 54); 2

pairs of dark mesal processes of tergum X, slen¬

der. slightly curved, of similar length (Fig. 55),

individual processes of inner pair adjacent, outer

processes slightly more widely separated (Fig.

55), robust pair of lateral processes of tergum X

situated below phallus (Figs 53 55); phallus rela¬

tively robust with broadbased process ventrally

(Fig. 54) and dorsal sclerotised ‘hood-like'

projection (Figs 53. 54).

Female. Female genitalia relatively short,

broad; sternum Vll with keel-like process.

AUSTRALIAN SPECIES OF CH/MARRA (TRICHOPTERA)

411

Posterolateral margin of segment VIII with scle-

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclerites relatively

narrowly separated, with connecting posterior

margin of segment VIII relatively straight and

witliout notch. Segment VIII dorsally with mem¬

branous area almost dividing segment. Ninth ster¬

num with pair of poorly deHned lightly pigmented

subtriangular sclerites, their inner margins touch¬

ing at base. Tergum X forming 2 lobes with

numerous .setae, each with relatively short apical

cercus (Figs 96, 97).

Eivntolog}’. Akriina - anagram of uranka.

Remarks. Chimarra aknma is a common species

throughout the Alligator Rivers region, north¬

ern Northern Territory (latitudinal range

12°26'-13°34'S).

Chimarra pita sp. nov.

Figures 56-58, 98, 99

Type material, l lolotypc male. Mitchell Plateau. Camp

Ck. It ir.. 13 Jul 1978. PS and M. Tyler (NMV, T-

I78'’8). Paratypes. I male, Mitchell Plateau, Camp Ck

at crusher, UV It. 15 Feb 1979. JF.B (NMV); 1 male,

Mitchell Plateau. Camp Ck at crusher. UV It, 15 Feb

1979. JEB (NMV); I male. Milehell Plateau, Kimber¬

ley, It tr., 30 .Ian 1978. JFB (NMV); I male. Mitchell

Plateau. Camp Ck. UV It tr., 20 Jul 1978. PS and M.

Tyler (NMV); I male (speeimen CT-269 figured).

Prince Regent R., Kimberley. I5°47'S, 125°24'E, May

1985, E. Bloomfield (NMV); I female (speeimen CT-

326 figured). Kimberley, Prince Regent R., 15°47 S,

125°24'E. May 1985, E. Bloomfield (NMV).

Other material e.xamincd Western Australia. I male, nr

Mitchell Plateau airfield, 14=48'S, I25°49'E, at light,

15 May 1983, J.C. Cardale (ANIC); I male. ‘Martin’

CALM site 8/4, Prince Frederick Harbour. I5°00 S.

l25°2rE, at light. 6-11 Jun 1988, l.D. Naumann

(ANIC); 5 males. C.ALM site 28/3. 4 km W of King

Cascade. I5'’35 S. I28“15'E. 17-20 Jun 1988. T.A.

Weir (ANIC). Northern Territory. 117 males. I female.

Litchfield National Park, UV light. 6 Jun 1991. Wells

and Webber (NTM); 4 males. 2 females, Litchfield

Park, MV light. 3 Apr 1991. J. Webber and R. De Jong

(NTM); I male. I female, Litchfield National Park,

lagoon nr Tolnier Falls. UV It. 24-25 Jun 1992, Wells

and Webber (NTM); 4 males. 5 females, Litchfield

National Park, billabong, UV, 18-19 Apr 1992. AW

(NTM); I male, Litchfield National Park. Walker Ck,

18 Apr 1992, AW (NTM).

Diagnosis. Chimarra pita most closely resembles

C. aknma in general fonn of male genitalic struc¬

tures. but is distitiguished by having two pairs of

dark slender, mesal processes of tergum X, outer

pair about half to two-thirds length of inner pair.

individual processes of inner pair adjacent, outer

processes narrowly separated from inner

processes.

Description. Head pale with brownish triangular

area anterior to ocelli, body and wings brownish;

length of forewing: male 4.0-4.8 mm, female

4.9-5.5 mm; wing venation: forewing with forks

1,2,3 and 5 present; hindwing with forks 2, 3 and

5 present. In forewing, vein Rs thickened and

curved slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

robust, length about twice width, upturned and

tapering slightly apically (Fig. 56). in ventral

view, length about 2.5 times width (Fig. 57); two

pairs of dark mesal processes of tergum X, slen¬

der, slightly curved, outer pair about halt to two-

thirds length of inner pair, individual processes ot

inner pair adjacent, outer processes narrowly

separated from inner processes (Fig. 58). pair of

lateral processes of tergum X situated below phal¬

lus, relatively slender with slightly out turned

apices (Figs 56-58); phallus relatively robust with

broadbased process ventrally (Fig. 57) and dorsal

sclerotised ‘hood-like’ projection (Fip 56, 58).

Female. Female genitalia relatively short,

broad; sternum VII with short keel-like process.

Posterolateral margin of segment VIII with scle¬

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclerites relatively

narrowly separated, with connecting posterior

margin of segment Vlll slightly convex and with¬

out notch. Segment Vlll dorsally with membra¬

nous area almost dividing segment. Ninth ster¬

num with poorly defined lightly pigmented areas.

Tergum X forming 2 lobes with numerous setae,

each with a relatively short apical cercus (Figs

98, 99).

Etymology. Pita — Australian Aboriginal word lor

four-pronged spear (four mesal processes on

tergum X).

Remarks. Chimarra pita is found in the Kimber¬

ley region, north Western Australia, Litchfield

Park, northern Northern Territory (latitudinal

range I2‘’58 -15°47'S).

Chimarra monticola-gmup

The C. monticola-group of three species, C. mon-

ticola, C. australiea and C. kewarra, is charac¬

terised by not possessing any of the primary dis¬

tinguishing characters of the previous groups, but

with male genitalia with relatively slender

upturned inferior appendages. Other common but

not exclusive characters are dark head, body and

412

D. CARTWRIGHT

wings, wings unicolorous, phallus with several

endothecal spines, ventral process on segment IX

small and keel-like; females with pair of dark

sub-triangular sclerites on ninth sternum. All

members of this group are from eastern Australia.

Chimarra monticola Kimmins

Figures 59-61, 122, 123

Chimarra monticola Kimmins in Mosely and Kim¬

mins, 1953: 402. fig. 275.—Neboiss, 1986: 105.—

Cartwright, 1990: 19, fig. 16 (female).

Type material. Holotype male. New South Wales,

Rules, Point, 4450 ft, 30 Dec 1934, R. J. Tillyard

(BMNH). Type not seen.

Material examined. Tasmania. 1 male (PT-802). Great

Lake, stream S of Breona, 27 Jan I960, F. McDonald.

Victoria. I male, (specimen CT-278 figured) OShan-

nassy R.. 12 Dec 1975, J. Dean and D. Cartwright; 1

female (specimen CT-322 figured). O'Shannassy

R.-Yarra R. jn, I Dec 1977. J. Dean and D. Cartwright;

Otway Ranges, Yarra River system, Mitta Milla River

system, north-east. East Gippsland. New South Wales.

South-east, north-cast. List of localities available from

author.

Diagnosis. Resembling C. attsiralica and

C. kewarra in general form of male genitalia, but

the fonn of the lateral processes on tergum X.

with a pair of short, broadbased ventrolateral pro¬

cesses, is quite distinct from the hooked processes

in those species.

Description. Head, body and wings black; length

of forewing: male 6.0-7.3 mm, female 7.3-8.5

mm; wing venation: forewing with forks 1, 2, 3

and 5 present; hindwing with forks 2. 3 and 5 pre¬

sent. In forewing, vein Rs not thickened or curv ed

basal to discoidal cell.

Male. (Revised after Kimmins in Mosely and

Kimmins, 1953 ). Ventral process on segment IX

small, keel-like; inferior appendages in lateral

view, length about 3.0 times width, upturned,

tapering slightly apically (Fig. 59), in ventral

view, broadbased, apices turned slightly mesally

(Fig. 60); no mesal processes of tergum X appar¬

ent, pair of lateral processes of tergum X short,

broadbased; phallus relatively long, slender, with

about 4 endothecal spines embedded subapically

(Figs 59-61).

Female. (Revised after Cartwright, 1990).

Female genitalia relatively short, broad; sternum

VII with weak keel-like process. Posterolateral

margin of segment VIII with sclerites and associ¬

ated setae dorsally. near middle and ventrally.

Ventral pair of sclerites relatively narrowly sepa¬

rated, with connecting posterior margin of

segment VIII relatively straight and without

notch. Segment Vlll dorsally with a membranous

area almost dividing segment. Ninth sternum with

pair of darkly pigmented triangular sclerites,

length less than width, their inner margins touch¬

ing at base. Tergum X forming 2 lobes with

numerous setae, each with relatively short apical

cercus (Figs 122, 123).

Remarks. Chimarra monticola is a common and

widespread species throughout eastern NSW and

Victoria, including one male specimen from cen¬

tral Tasmania (latitudinal range 21°02'-4i°47'S).

According to Neboiss (pers. comm.) the Tas¬

manian male (PT-802) is a close but distinct

species. Kimmins' (in Mosely and Kimmins,

1953) and Cartwright's, (1990) figures have been

redrawn to allow direct comparisons and to

accompany the description that is revised in light

of new interpretations of Chimarra genitalic

structures.

Chimarra australica Ulmer

Figures 62-64, 118, 119

Chimarra australica Ulmer, 1916: 3-5, figs 3-6.—

Mosely and Kimmins, 1953: 399, fig. 274.—Neboiss,

1986: 105.—Cartwright, 1990: 20, fig. 18 (female).

Type material. Holotype male. Queensland, Malanda,

date and collecter unknown (NRS). Type not seen.

Material examined. Queensland. I female. Zarda Ck nr

Mt Misery. W of Mossman, 1200m, 23 Dec 1974,

MSM; 1 male (CT-275). 25 km along Mt Lewis Rd.

SW of Mossman, 16 Jan 1977, M.S. and B.J. Moulds: I

male. Rocky Ck, 11 km N of Atherton. 3 May 1967,

D.ll. Colless (ANIC); I male (CT-276). Bellenden Ker

Range, Cableway Base Station, lOOm, 17-24 Get 1981.

Earthwatch/Qld Museum; I male (specimen CT-3()9

partly figured). Ml Spec Slate Forest. Birthday Ck

above weir, 18°57'S. 146°I0'E. It tr.. 13 Nov 1993,

A.L. Sheldon; 2 males. 1 female, same site, 6 Dee 1993,

A.L. Sheldon; 1 male, 1 female?, same site. 15 Oct

1993, A.L. Sheldon; 1 male, same loeality. 6 Nov 1993;

1 male (specimen CT-292 partly figured). Finch Hatton

Gorge. 2r()7'S, 148“38'E. 14 Nov 1982. Theischinger;

1 female, (specimen CT-323 figured); I male, MtNebo

area, slow stream at roadside. I May 1975, coll.?

(ANIC); New South Wales. North-east, central east and

south-east; 1 male (specimen CT-277 partly figured),

Brogo River. 22 Jan 1977, J. Dean?. Victoria. East

Gippsland, norlh-ea.st. Yarra River system, Lai Lai

Falls, near Lome. List of other localities available from

author.

Diagnosis. This species most closely resetnbles

C kewarra in general form of male genitalic

structures, but is distinguished by having lateral

processes of tergum X, with relatively short.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

413

upturned or out turned apices. The phallus has a

distinctive ventral process.

Description- Head, body and wings black to

brownish-black; length of forcwing: male 4.7-5.9

mm, female 4.9-7.0 mm; wing venation:

forcwing with forks I, 2, 3 and 5 present; hind¬

wing with forks 2, 3 and 5 present. In forewing,

vein Rs not thickened or curved basal to discoidal

cell.

Male. (Revised after Ulmer, 1916 and Mosely

and Kimmins, 1953). Ventral process on segment

IX small, keel-like; inferior appendages in lateral

view, length about twice width, upturned, taper¬

ing slightly apically (Fig. 62), in ventral view,

broadbased. apices turned slightly mesally (Fig.

63); mcsal processes of tergum X not apparent,

pair of lateral processes of tergum X broadbased,

relatively slender apically, apices hooked

upwards or sometimes outwards (Figs 62, 62a-c,

64, 64a-c); phallus relatively long, with ventral

subapical process and I or 2 endothecal spines

embedded subapically (Figs 62-64).

Female. (Revised after Cartwright, 1990).

Female genitalia relatively short, broad; sternum

VII with small keel-like process. Posterolateral

margin of segment VIII with sclerites and associ¬

ated setae dorsally, near middle and ventrally.

Ventral pair of sclerites relatively narrowly sepa¬

rated, with connecting posterior margin of seg¬

ment VIII relatively straight and without notch.

Segment VIII dorsally with a membranous area

almost dividing segment. Ninth sternum with pair

of darkly pigmented subtrapezoidal sclerites, their

inner margins almost touching at base. Tergum X

forming 2 lobes with numerous setae, each with

relatively short apical cercus (Figs 118, 119).

Remarks. Chimarra austraiica is a very common,

widespread and slightly variable species through¬

out eastern Australia (latitudinal range 16°34'-

38 ° 35 'S).UImer’s, (1916) and Cartwright’s,

(1990) figures have been redrawn to allow direct

comparisons and to accompany the description

that is revised in light of new interpretations of

Chimarra genitalic structures.

Chimarra kewarra sp. nov.

Figures 65-67, 120, 121

Type material. Holotype male (specimen CT-287 fig¬

ured). Queensland. Searys Ck, Rainbow Beach.

25°58'S, I53°04'E, 7 Jan 1986, G. Theischinger

(NMV, T-17906). I’aratypes. 2 females (specimen CT-

310 figured), collected with holotype (NMV).

Diagnosis. Chimarra kewarra most closely

resembles C. austraiica in general form of male

genitalic structures, but is distinguished by having

lateral processes of tergum X, with relatively

long, upturned or out turned apices and phallus

without a ventral process.

Description. Head, body and wings dark greyish-

brown to black; length of forewing; male 4.7 mm,

female 5.5-5.6 mm; wing venation: forewing

with forks I, 2, 3 and 5 present; hindwing with

forks 2, 3 and 5 present. In forewing, vein Rs

thickened and curved slightly basal to discoidal

cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

length about 2.5 times width, upturned, tapering

slightly apically (Fig. 65), in ventral view, broad¬

based, apices turned slightly mesally (Fig. 66);

mesal processes of tergum X not apparent, pair of

lateral processes of tergum X broadbased, later¬

ally flattened (Figs 65, 66), in ventral view rela¬

tively slender apically (Fig. 66), apices hooked

upwards; phallus robust, with pair of endothecal

spines embedded subapically (Figs 65-67).

Female. Female genitalia relatively short,

broad; sternum VII with small keel-like process.

Posterolateral margin of segment VIll with scle¬

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclerites relatively

nan'owly separated, with connecting posterior

margin of segment VIII relatively straight and

without notch. Segment VIII dorsally with

membranous area almost dividing segment. Ninth

sternum with pair of lightly pigmented triangular

sclerites, length greater than width, their inner

margins almost touching at base. Tergum X

fomiing 2 lobes with numerous setae, each with

relatively short apical cercus (Figs 120, 121).

Etymology. Kewarra - Southern Queensland

Aboriginal word for rainbow (type locality —

Rainbow Beach).

Remarks. At present Chimarra kewarra is known

only from the type locality in southeastern

Queensland (latitude 25°58'S).

Chimarra moiildsi-gmiip

The Chimarra moulcisi-group, complex of four

species, C. moukJsi. C. stclairae. C. hiharinga

and C. karakara, is almost certainly not a natural

group, but is characterised by not possessing any

of the primary distinguishing characters of the

previous groups, but all males have relatively

short inferior appendages. Other common but not

exclusive characters are dark head, body and

414

D. CARTWRIGHT

wings, wings unicolorous, tergum X without

apparent mesal processes and with short lateral

processes or lobes, ventral process on segment

IX small and keel-like. All members are from

north-east Queensland.

Chimarra nwtildsi sp. nov.

Figures 68-70, 114, 115

Chimarra sp. nov. CT-225.—Walker et al., 1995;

26.—Cartwright. 1997: 17.

Type material. Holotype male. Queensland, Upper

Freshwater Ck, Whitfield Range nr Cairns, 15 Dec

1974. Moulds (NMV. T-17835). Paratypes. 4 males

(specimen CT-225 figured). I female, collected with

holotype (NMV): 1 male, same locality. 24 Aug 1974,

MSM (NMV); 2 females (specimen CT-329 figured),

Bellendcn Ker Range, Cableway Base Sin, 100 m,

25-31 Oct 1981. Earthwalch/Qld Museum (NMV).

Other material examiiiecl. Queensland. I male. Black

Mt Rd, E ofMt Molloy, 5 Dec 1974. MSM; I male. 1.5

km SE Kuranda. 16-17 May 1980. I.D. Naumann and

J. Cardale (ANIC); I female, Cairns, Lake Morris Rd.

MV It. I6°55'S, 145“46'E. 16 Nov 1988, K. Walker; 1

female, Currunda Ck. Freshwater Ck Irib., on road to

Crystal Cascades, nr Cairns, 30 Apr 1979. AW; 2

males. Qld, Lock-Davies Ck Rd. Lamb Range, Mareeba

district, 10 Nov 1974. MSM; 1 female, 25 km Gordon-

vale. Gillies 11-way, 4 Sep 1974, MSM; 1 female, Bel¬

lendcn Ker Range. Cableway Base Stn, 100 m, 1-7 Nov

1981. Earthwatch/Qld Museum; 4 females. Carron Ck.

Kirrama Stale Forest. I7°50'S. 145°35'E, Apr 1993. G.

Theischinger; 2 females, Goodard Ck. Kirrama State

Forest, 18°06 S. I45°4I E. Apr 1993. G. Theischinger;

1 male (PT-1642). Yuccabine Ck. Kirrama State Forest.

I8°12 S, 145°45 E, 2 Jan 1985. R. Pearson.

Diagnosis. Grouped with C stclairae. C.

bibaringa and C. karakara but distinguished by

having inferior appendages which in lateral view

are subquadrate, truncate apically.

Description. Brown head, body and wings; length

of forewing: male 4.6-5.2 mm. female 4.6-5.2

mm; wing venation: forewing with forks 1. 2, 3

and 5 pre.sent; hindwing with forks 2, 3 and 5 pre¬

sent. In forewing, vein Rs thickened and curved

slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

short, robust, subquadrate, truncated, length about

1.5 limes width (Fig. 68), in ventral view, length

about same as width (Fig. 69); mesal processes of

tergum X not apparent, pair of lateral processes of

tergum X situated beside phallus, robust, tapering

slightly apically (Figs 68, 70); phallus relatively

robust with pair of dark, slender endothecal

spines embedded subapically (Figs 68-70).

Female. Female genitalia relatively short,

broad; sternum VII with short keel-like process.

Posterolateral margin of segment Vill with scle-

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of sclcrites relatively

narrowly separated, with connecting posterior

margin of segment VIII relatively straight and

without notch. Segment Vlll dorsally with mem¬

branous area almost dividing segment. Ninth ster¬

num with pair of irregular-shaped pigmented

areas. Tergum IX with pigmented scleroli.sed area

extending down sides and with truncate apices.

Tergum X forming 2 lobes with numerous setae,

each with relatively short apical cercus (Figs 114

115).

Etymology. Mouldsi - named after Max Moulds

(collector).

Remarks. Chimarra mouklsii is an uncommon

and distinctive north-eastern Queensland species,

(latitudinal range 16°4r-18°l2’S).

Chimarra stclairae sp. nov.

Figures 71-73, 112, 113

Chimarra sp. nov. CT-226.—Walker ct al., 1995:

26.—Cartwright. 1997: 17.

Type material. Holotype male, Queensland, Birthday

Ck, 3.5 km WNN Palunia, I8°59'S. 146°10'E. at light

17 Feb 1990. R. St Clair (NMV. T-17875). Paratypes. 2

males, saine locality and collector as holotype, 8 Oct

1989 (NMV); 7 males, 2 females, same locality and col¬

lector as holotype. 19 Jan I990(NMV);1 female (spec¬

imen CT-3()5 ligured). same locality and collector as

holotype. 17 Feb 1990 (NMV); I male (specimen CT-

226 figured), same locality and collector as holotype, 1

Apr 1990 (NMV); 2 males. 1 female. Birthday Ck, 3.5

km WNN Paluma, 18°59 S. UO^IO'E, al light. 23 Dec

1989. R. St Clair (NMV); 7 males, Birlhday'ck, 3.5 km

WNN Paluma. I8°59 S. ITO^IO'E, at light. 17 Mar

1990, R. St Clair (NMV); I male. 3 females. Birthday

Ck below falls. Ml Spec Slate Forest, I8°57'S.

I46®I0'E. 760 m. It. tr., 29 Mar 1994, A.L. Sheldon

(NMV); I male. Birthday Ck. iron cabin, Mt Spec State

Forest. I8°57'S, I46°I() E, 790 m. It. tr., 23 Apr 1994,

A.L. Sheldon (NMV).

Other material examined. Queensland. I male, 7 km N

of Hopevale Mission. 15°14'S. I45°07'E, 4 Oct 1980.

J.C. Cardale (ANIC); I male. 1 female, 8-13 km Mt

Lewis Rd, off Mossman-Molloy Rd, 22 Apr 1967. D.H.

Collcss (ANIC); 1 male, I female, Kirrama Slate For-

c.st. Western Fall. 30 May 1971, E.F, Rick (ANIC); I

male. Birthday Ck. iron cabin. Ml Spec State Forest.

I8'57 S. I46“I0-E, 790 m. It. tr.. 23 Apr 1994. A.L.

Sheldon; I male. 3 females. Birthday Ck below falls.

Mt Spec State Forest, I8'=57'S. 146- lO'E, 760 m. It. tr..

29 Mar 1994. A.L. Sheldon.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

415

Diagnosis. Chimarra stciairae is grouped with

C. niouldsi, C. biharinga and C. karakara but can

be distinguished by having inferior appendages

which in lateral view are short, upturned, tapering

slightly apically, in ventral view with small

process on inner margin.

Description. Head, body and wings brown; length

of forewing: male 4.6-5.5 mm, female 5.3-5.5

mm; wing venation: forewing with forks 1, 2, 3

and 5 present; hindwing with forks 2, 3 and 5 pre¬

sent. In forewing, vein Rs thickened and curved

slightly basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

short, length about twice width, upturned, taper¬

ing slightly apically (Fig. 71), in ventral view,

short, length about twice width, with small pro¬

cess on inner margin (Fig. 72); mesal and lateral

processes of tergum X not apparent; phallus

relatively short, with pair of long embedded

endothecal spines (Figs 71-73).

Female. Female genitalia relatively short,

broad; sternum VII with short keel-like process.

Posterolateral margin of segment VIII with scle-

rites and associated setae near middle and ven-

trally. Ventroposterior margin of segment VIII

relatively straight and without notch. Segment

Vlll dorsally with membranous area almost divid¬

ing segment. Ninth sternum with pair of squarish

depressions. Tergum IX with pigmented sclero-

tised area extending down sides and with rela¬

tively pointed apices. Tergum X forming 2 lobes

with numerous setae, each with relatively short

apical cercus (Figs 112, 113).

Etymology. Stciairae - named aller Ros St Clair

(collector).

Remarks. This species is known from only

northeastern Queensland (latitudinal range

I5°I4'-18°59'S).

Chimarra biharinga sp. nov.

Figures 74-76

Chimarra sp. nov. CT-227.—Walker et al., 1995:

26.—Cartwright, 1997: 17.

lype material. Ilolotype male, Queensland, Bellenden

Ker Range, Cableway Base Stn, 100 m, 17-24 Oct

1981, Earthwateh/Qld Museum (NMV, T-17844).

Paratypes. I male (specimen CT-227 figured), collected

with holotype (NMV); I male, same locality and col¬

lector. 1-7 Nov 1981 (NMV).

Ollier material e.xamined Queensland. I male. Moses

Ck. 4 km N by E Mt Einnigan, 14-16 Oct 1980. J.C.

Cardale (ANIC); I male, Mossman Gorge, 16 Nov

1979, AN; 2 males. Upper Freshwater Ck, Whitfield

Range nr Cairns, 15 Dec 1974, Moulds; I male.

Upper Freshwater Ck, Whitfield Range nr Cairns, 24

Aug 1974, MSM; I male pupa. Little Mulgrave R.,

28 Jun 1971, E.F. Riek (ANIC); I male. Base cable-

way, Mt Bellenden Ker, 80 m. I7°16'S. I45°54'E, 19

Oct 1981, E. D. Edwards (ANIC); 1 male. Williams Ck

tributary, Mt Spec State Forest, I8‘’57'S. I46°I0'E,

745 m. It. Ir., 13 Nov 1993, A.L. Sheldon; 4 malc.s.

Little Crystal Ck, Mt Spec. 29 May 1971, E.F. Riek

(ANIC).

Diagnosis. This species is grouped with

C. inouldsi. C. stciairae and C. karakara but can

be distinguished by having lateral processes of

tergum X with obvious elongate processes ventral

to phallus.

Description. Head, body and wings dark brown to

blackish; length of forew ing: male 4.5-5.2 mm;

wing venation: forewing with forks 1, 2, 3 and 5

present; hindwing with forks 2, 3 and 5 present. In

forewing, vein Rs thickened and curved slightly

basal to discoidal cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

short, subovate, length about 2.5 times width (Fig.

74) , in ventral view, length about twice width,

broadbased, tapering and curxed apically (Fig.

75) ; mesal processes of tergum X not apparent,

lateral processes of tergum X with elongate ven¬

tral processes, extending near apices of inferior

appendages; phallus relatively robust with no

embedded endothecal spines apparent (Figs

74-76).

Female unknown.

Etymologyi. Biharinga — Queensland Aboriginal

word for mountain.

Remarks. Chimarra biharinga is a distinctive and

uncommon north-eastern species (latitudinal

range I5°48'-I9°00’S).

Chimarra karakara sp. nov.

Figures 77-79, 116, 117

Chimarra sp. n. F.—Wells and Cartwright, 1993:

227.

Chimarra sp. nov. CT-222.—Cartwright, 1997: 17.

Type material. Holotype male. Queensland, Gunshot

Ck. Telegraph Crossing. UV It. I1°44'S. I42°29 E,

14-15 Feb 1992. Cartwright and Wells (NMV, T-

17847). Paratypes. I male, collected with holotype

(NMV); I male (specimen CT-273 figured) same local¬

ity and collectors, 18 Feb 1992; 2 males, 2 females

(specimen CT-3()() figured) same locality and collec¬

tors, 18 Feb 1992; 10 males, 11 females, same locality

and collectors, 17 Feb 1992 (NMV).

416

D. CARTWRIGHT

Other material examined. Queensland. Iron Range,

Mcllwraith Range, near Cooktown, near Cairns, Bel-

lenden Ker Range, Kirrama Stale Forest, Mt Spec State

Forest, Eungella National Park (ANIC, NMV, QM).

List of localities available from author.

Diagnosis. Chiniarra karakara is grouped with

C mouldsi, C. stclairae and C. bibaringa but

can be distinguished by having relatively long

phallus with bulbous head with 3 obvious and

characteristic small dark endothecal spines.

Description. Head, body and wings dark brown to

blackish: length of forewing; male 4.4-5.8 mm,

female 5.1-6,7 mm; wing venation; forewing

with forks 1, 2, 3 and 5 present; hindwing with

forks 2, 3 and 5 present. In forewing, vein Rs

thickened and curved slightly basal to discoidal

cell.

Male. Ventral process on segment IX small,

keel-like; inferior appendages in lateral view,

short, subovate, length about 1.5 times width (Fig.

77), in ventral view, length about 2.5 times width,

curved slightly apically (Fig. 78); no mesal pro¬

cesses of tergum X apparent, pair of lateral

processes of tergum X short; phallus relatively

long, in lateral view with bulbous head with 3

obvious and characteristic embedded small dark

endothecal spines (Figs 77-79).

Female. Female genitalia relatively short,

broad; sternum VII with short keel-like process.

Posterolateral margin of segment Vlll with scle-

rites and associated setae dorsally, near middle

and ventrally. Ventral pair of selerites relatively

narrowly separated, with connecting posterior

margin of segment Vlll slightly convex and with¬

out notch. Segment Vlll dorsally with membra¬

nous area almost dividing segment. Ninth ster¬

num with pair of oblong pigmented areas.

Tergum IX with pigmented sclcrotised area

extending down sides and with pointed apices.

Tergum X fonning 2 lobes with numerous setae,

each with relatively short apical cerciis (Figs 116,

117).

Etymology. Karakara - Queensland Aboriginal

word for black (body and wings colour).

Remarks. Chimarra karakara is a very common

and distinctive dark species found throughout

northeastern Queensland (latitudinal range

11°44'-2I°28'S).

Acknowledgements

1 thank Environment Australia, in particular Aus¬

tralian Biological Resources Study (ABRS) for

providing a grant to undertake this work, Drs

Arturs Neboiss and Alice Wells for providing

access to the specimens, and together with two

referees, for constructive comments on a draft of

this manuscript.

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Zealand. British Museum (Natural History):

London. 550 pp.

Neboiss. A., 1986. Atlas of Trichoptera of the SIV

Paeijic-Australian Region. Dr W. Junk: Dordrecht.

286 pp.

Neboiss. A.. 1988. Trichoptera. Pp 177-280 in Walton.

D.W. (ed.). Zoological catalogue of Australia. Vol¬

ume 6. Ephemeroptera. Megaloptera. Odonata,

Plecoptera, Trichoptera. Australian Government

Publishing Service: Canberra.

Neboiss, A. and Wells, A.,1998. Review of Australian

species of Triaenodes McLachlan (Trichoptera:

Lcptoceridae). Memoirs of the Mimnim of Victoria

57:89-132.

Nielsen, A., 1957. A comparative study of the genital

segments and their appendages in male

Trichoptera. Biologiske Skrifter 8: 1-159.

Nielsen, A., 1981. A comparative study of the genital

segments and the genital chamber in female

Trichoptera. Biologi.ske Skrifter 22: 1-200.

Ross, II., 1956. Evolution and classification of the

mountain vaddisflies. LIniversily of Illinois Press:

Urbana. 213 pp.

Stephens. J.F.. 1829. A systematic catalogue of British

insects. Pt 1. Baldwin: London. 416 pp.

Ulmer, G., 1916. Results of Dr E. Mjoberg'S, Swedish

Scientific Expedition to Australia. 1910-1913.

Trichoplcra. Arkiv fur Zoologi 10: 3-5.

Walker, K., Neboiss, A., Dean, J. and Cartwright. D.,

1995. A preliminary investigation of the caddis-

llies (Insecta: Trichoptera) of the Queensland wet

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

417

tropics. Australian Entomologist 22: 19-31.

Wells, A., 1991. A guide to the caddisjlies (Tri-

choptera) of the Alligator Rivers Region, Northern

Territory. Open File Record. Supervising Scientist

for the Alligator Rivers Region. 105 pp.

Wells, A., and Cartwright, D.I., 1993. Trichoptera.

Ephemeroptera, PIccoptera and Odonata of the

Jardine River area. Cape York, northern Queens¬

land. Cape York Peninsula Scientific Expedition

Wet Season 1992. Report 2: 221-230. Royal

Geographical Society of Queensland. Inc.

Index and checklist of species of Chimarra from Australia

Description

page no.

male figure

nos

female figure

nos

Chimarra adahtma sp. nov.

408

44^6

C. akruna sp. nov.

410

53-55

96-97

C australica Ulmer, 1916

412

62-64

118-119

C. hiharinga sp. nov.

415

74-76

C. btingoona sp. nov.

401

11-13

88-89

C. kaiva sp. nov.

400

8-10

86-87

C. karakara sp. nov.

415

77-79

116-117

C. kewarra sp. nov.

413

65-67

120-121

C. larapinta sp. nov.

401

14-16

C. locolo sp. nov.

400

la, 5-7

82-83

C. htminaris sp. nov.

399

1-4

80-81

C. moiiticola Kimmins, 1953

412

56-61

122-123

C. mottlJsi sp. nov.

413

68-70

114-115

C. nahilla sp. nov.

409

50-52

102-103

C. natalicia sp. nov.

404

26-28

104-105

C. nehoissi sp. nov.

405

29-31

108-109

C. orttmbera sp. nov.

402

17-19

84-85

C. pillara sp. nov.

403

20-22

92-93

C. pita sp. nov.

411

56-58

98-99

C. ramtka sp. nov.

406

35-37

C. sictairae sp. nov.

413

71-73

112-113

C. tallawalla sp. nov.

407

38^0

106-107

C. iiranka Mosely, 1953

406

32-34

94-95

C. wooroonooran sp. nov.

408

41^3

110-111

C. yatidala sp. nov.

403

23-25

90-91

C. yoolumba sp. nov.

409

47^9

100-101

418

D. CARTWRIGHT

Figures 1-7. Chimarra spp. 1-la: Wings. 1: Cliimcirni luminaris sp. nov. forewing and hindwing.la: Chimaira

locolo sp. nov. section of forcwing and hindwing. 2-7: Male genitalia in lateral, ventral and dorsal views. 2-4: Chi¬

marra luminaris sp. nov. 5-7: Chimarra locolo sp. nov.

Abbreviations: ia, inferior appendages; Ip X, lateral process of abdominal tergum X; mp X, mesal process of

abdominal tergum X; pha, phallus; vp, ventral process of abdominal segment IX. Scale lines: l-la, 0.5 mm; 2-7,

0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

419

Figures 8-13. Cbimarra spp. Male genitalia in lateral, ventral and dorsal views. 8-10: Chimarra kaiya sp. nov.

11-13: Chimarra himgoona sp. nov. Abbreviation: b, ‘barb-like projection'. Scale lines 0.1 mni.

420

D. CARTWRIGHT

Figures 14-19. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 14-16: Chimarra larapinla sp.

nov. 17-19: Chimarra ommbera sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

421

Figures 20-25. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 20-22: Chimarrapiltara sp. nov.

23-25: Chimarrayandala sp. nov. Scale lines 0.1 mm.

422

D. CARTWRIGHT

Figures 26-31. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 26-28: Chimarra nalalicia sp.

nov. 29-31: Chimarra neboissi sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA) 423

Figures 22-37. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 32-34: Chinuvni tiranka Mosely.

32a: lateral view of phallic complex. 35-37: Chimarra rarnika sp. nov. 35a: lateral view of phallic complex.

Abbreviations: mp X, mesal process of abdominal tergum X; pp, dorsal phallic projection. Scale lines 0.1 mm.

424

D. CARTWRIGHT

Figures 38^3. Chimaira spp. Male genitalia in lateral, ventral and dorsal views. 38^0: Chimarra tallawalla sp.

nov. 38a: lateral view of phallic complex. 38b, 40a: Chimarra tallawalla sp. nov. (CT-318-Chandlers Ck.,

NE-NSW), phallus and phallic spines lateral and dorsal views. 41-43: Chimarra wooroonooran. 41a: lateral view

of phallic complex. Abbreviation: ps, phallic spine. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

425

Figures 44-52. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 44^6: Chimarra adaluma sp. nov.

47^9: Chimarrayoolumba sp. nov. 50-52: Chimarra nahilla sp. nov. Scale lines 0.1 mm.

426

D. CARTWRIGHT

Figures 53-58. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 53-55; Chimarra akruna sp. nov.

56-58: Chimarra pita sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

427

"-r:-' / .,y J

Figures 59-64. Chimarni spp. Male genitalia in lateral, ventral and dorsal views. 59-61: Chimatra monticola Kim-

mins.62-64; Chimarro ausiralica Ulmer. 62a,b,c-64a,b,c; Chimarra ausiralica Ulmer (CT-277, Brogo R,. SE-

NSW; CT-292, Finch Hatton Gorge, NE-Qld; CT-309, Birthday Ck, NE-Qld), lateral process of tergum X, lateral

and dorsal views. Abbreviation: vpp, ventral process of phallus. Scale lines 0.1 mm.

428

D. CARTWRIGHT

Figures 65-67. Chimarm spp. Male genitalia in lateral, ventral and dorsal views. 65-67: Chimarra kewarra sp. nov.

Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

429

Figures 6S-72. Chimarra spp. Male genitalia in lateral, ventral and dorsal views. 68-70: Chimarra mouldsi sp. nov.

71-73: Chimarra stclairae sp. nov. Scale lines 0.1 mm.

430

D. CARTWRIGHT

Figures 74-79. Chimmra spp. Male genitalia in lateral, ventral and dorsal views. 74-76: Chimarra hiharinga sp.

nov. 77-79: Chimarra karakara sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

431

Figures 80-85. Chimarra spp. Female genitalia in lateral and ventral views. 80-81: Chimarra lum'maris sp. nov.

82-83: Chimarra locola sp. nov. 84-85: Chimarra orumbera sp. nov. Abbreviations: c, cercus; st VIII, sternum

abdominal segment VIII; st IX, sternum abdominal segment IX. Scale lines 0.! mm.

432

D. CARTWRIGHT

Figures 86-91. Chimarra spp. Female genitalia in lateral and ventral views. 86-87: Chimarra kaiya sp. nov. 88-89

Chimarra yandala sp. nov. 90-91: Chimarra biingoona sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

433

Figures 92-97. Chimarra spp. Female genitalia in lateral and ventral views. 92-93: Chimarra pillara sp. nov,

94-95: Chimarra uranka Mosely. 96-97: Chimarra akrtma sp. nov. Scale lines 0.1 mm.

434

D. CARTWRIGHT

Figures 98-103. Chimarra spp. Female genitalia in lateral and ventral views. 98-99; Chimarra yoolumba sp. nov.

100-101: Chimarra nabilla sp. nov. 102-103: Chimarra pita sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

435

Figures 104-109. Chimarra spp. Female genitalia in lateral and ventral views. 104-105: Chimaira natalicia sp.

nov. 106-107: Chimarra lallawalla sp. nov. 108-109: Chimarra nehoissi sp. nov. Scale lines 0.1 mm.

436

D. CARTWRIGHT

Figures 110-117. Chimarra spp. Female genitalia in lateral and ventral views. 110-111: Chimarra wooroonoonan

sp. nov. 112-113: Chimarra stclairae sp. nov. 114-115: Chimarra mouldsi sp. nov. 116-117: Chimarra karakara

sp. nov. Scale lines 0.1 mm.

AUSTRALIAN SPECIES OF CHIMARRA (TRICHOPTERA)

437

Figures 118-123. Chimarra spp. Female genitalia in lateral and ventral views. 118-119: Chimarra australica

Ulmer. 120-121: Chimarra kewarra sp. nov. 122-123: Chimarra monticola Kimmins. Scale lines 0.1 mm.

Memoirs of the Museum of Victoria 59(2): 439-455 (2002)

FIVE ARTOTROGIDS (CRUSTACEA; COPEPODA: SIPHONOSTOMATOIDA)

FROM EASTERN ANTARCTICA

R. JOHNSSON AND C. E. F. ROCHA

Universidade de Sao Paulo, Instituto Biociencias, Departamento Zoologia,

Caixa Postal 11461, CEP; 05422-970 Sao Paulo, SP, Brazil

(e-mail: johnsson@ib.usp.br)

Abstract

Johnsson, R. and Rocha, C.E.F., 2002. Five artotrogids (Crustacea; Copcpoda: Siphonostom-

atoida) from Eastern Antarctica. Memoirs of Museum Ficioria 59(2): 439-455

Five .species ol Artotrogidae are reported from Enderbyland, Eastern Antarctica. Four are

new species belonging to the genera Sesiropnniius. Bradxponlius, Arcloponlius and

Neohracfyponliiis; Pscudotrogus imcinalus (Brady, 1910) is redescribed. The new species of

Sestropomius differs from its only congener in the armature of PI, P2 and P4. In Brady-

pontius the new species differs from other species in the armature of PI and the setation of

the maxillule. The number of segments in the antennule and modifications on the distal

element of the antenna distinguish the new species of Neohradyponlius from its congeners. In

Arcloponlius the number of segments in the antennule is the key difference between species.

Introduction

The family Artotrogidae was revised by Eiselt

(1961) and since then new species have been

added from Korea (Kim, 1996), the Arctic Ocean

(Eiselt. 1986), Madeira I. (Johnsson. 2001) and

southern Australia (McKinnon, 1988). McKin¬

non (1988) treated Artotrogidae in the sense of

Sars, 1915, i.e. taxa possessing only three pairs

of swimming legs, while the species studied in

the present paper deals with Artotrogidae in the

broader sense, those with four pairs of swimming

legs. A second species of Sesiropontiiis Gies-

brecht is described, more than a century after the

first. A new species is added to BraJyponlius

Geisbrecht, increasing the number of known

species to 2 \. Neohracfyponliiis, erected by Eiselt

(1961) with species belonging to other genera

has its first new species described. The genus

Arcloponlius Sars has now three species and

Pseuciolrogus uncinalus (Brady, 1910) is

redescribed.

Due to the rarity of Artotrogidae most of the

species were partially dissected and these

appendages were clarillcd with lactic acid,

stained with Chlorazol Black and mounted on

slides with CMC-9 mounting medium. The

drawings were made with an Olympus CM30

microscope equipped with a camera lucida.

Armature formulae for swimming legs

are shown with outer margin first and Roman

numerals indicating spines and Arabic numerals

setae, according to lluys and Boxshall (1991).

Armature for caudal setae; I - anterolateral acces¬

sory seta, II — anterolateral seta. III - postero¬

lateral seta, IV - outer terminal seta, V - inner

terminal seta, VI - tenninal accessory seta and

VII - dorsal seta, according to Huys and Boxshall

(1991). Given the fact that the material is taken

from sled samples some elements from the

armature of the antcnnules may have been lost.

All material is deposited in Museum Victoria,

Melbourne, Australia (NMV).

Artotrogidae Brady

Sestropomius Giesbrecht

Sestropomius nickinnoni Johnsson, sp. nov.

Figures 1-2

Material examined. Holotypc. Southern Ocean, off

Enderbyland, Antarctica (65“56.40S, 50”52.10E), 365

m. silt and bryozoan shell, M. Norman, 15 Nov 1985,

WHOI epibenthic sled (stn HRD 10). NMV J47289-a

(female).

Description. Female: Body length (excluding

caudal setae) 2.41 mm. greatest body width 1.17

mm, and 2.1 times as long as wide (fig. la). Body

shape cyclopiform, cephalosome and pedigerous

somite 2 with pointed epimera. Pedigerous

somite 3 with lateral margins bearing sensilla and

439

440

R. JOHNSSON AND C.E.F. ROCHA

Figure 1. Sestropontius mckinnoni sp. nov.; female holotype, a: habitus dorsal showing siphon (dotted line),

b: urosome (not showing first somite), c; antennule, d: antenna, e: distal part of mandible. Scale bars: a: 200 urn;

b-d: 100 pm; e: 50 pm.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

441

Figure 2. Sestropontius mckiimoni sp. nov.; female holotype, a: maxillule, b: maxilla, c: maxilliped, d: PI, e: P2,

f; P3, g: P4, h: P5. Seale bars: 100 pm.

442

R. JOHNSSON AND C.E.F. ROCHA

reaching genital double-somite. Pedigerous

somite 4 also with sensilla on projected margins.

Length:width ratio of prosome 1.3. Ratio of

length prosome:urosome 1.8.

Urosome (fig. lb) with 5 somites, all with sen-

sillae. Genital double-somite 296 x 429 pm,

length:width ratio 0.7, rounded anterolaterally,

armed with plumose seta near laterally projected

genital opening. Posterior margin serrated. Three

postgenital somites 117 x 229 pm, 117 x 208 pm

and 167 x 200 pm, length:width ratios 0.5, 0.6

and 0.8 respectively. First and second postgenital

somites with posterior margins serrated. Caudal

rami, 154 x 92 pm, 1.7 times as long as wide, with

row ot hairs on inner margin and armed with 6

setae. Setae 1 absent. Lengths of setae 11-Vll, 117.

217, 308, 521. 354 and 162 pm respectively.

Setae Ill-Vl plumose, setae II and VII smooth.

Setae II and III located medially.

Antennule (fig. Ic) 906 pm long. 14-seg-

mented. Lengths of segments measured along

their posterior margins 189 pm (117 pm along

anterior margin) 43,46, 39. 29. 26, 31,34,49, 34,

63, 57, 57 and 117 respectively. Segmental

homologies and setation as follows; 1-1; II-2; III-

2; IV-2; V-2; VI-2; Vll-2; Vlll-2; IX-XIII-

5+spine; XlV-I+spine; XV-XVI-2; XVIl-XVIII-

2; XIX-XX-2; XXI-XXVIIl-8-(-ae. All setae

smooth. Aesthetasc on segment XXI 278 pm

long.

Antenna (fig. Id) 647 pm long (including distal

seta), with basis 178 pm long. Endopod 2-seg-

mented; first segment 100 pm long, unarmed;

second segment 167 pm long and armed with

1 smooth seta proximally and 2 setae distally

close to a claw-like element, 203 pm long.

Exopod I-segmented, 25 pm long, bearing 3

setae.

Oral cone (fig. la) produced into long siphon¬

like distal portion, 1.2 mm long. 0.5 times body

length. Mandible (fig. le) comprising stylet bear¬

ing distally many small teeth, palp absent. Maxil-

lule (fig. 2a) bilobed, inner lobe 189 pm long,

armed with a smooth stout seta, a short plumose

seta and a distally plumose seta, inner lateral

margin covered with setules. Outer lobe 139 pm

long, armed with a pinnate seta and a plumose and

pinnate seta.

Maxilla (fig. 2b) syncoxa 521 pm long; claw

588 pm long, cuiAcd distally, bearing seta sub-

distally, 2 small teeth on outer margin and claw

tip serrated. Maxillipcd (fig. 2c) 5-segmentcd;

syncoxa 144 pm long and bearing short seta on

inner margin and long setules on outer margin;

basis 417 pm long, with small seta subdistally on

inner margin. Endopod 3-segmented, segments

1 to 3 measuring 72, 19 and 111 pm long respec¬

tively. First endopod segment with 2 setae; sec¬

ond and third segments with short seta distally;

third segment with claw 225 pm long.

Swimming legs 1-4 (PI P4, figs 2d-g) bira-

mous, with 3-segmented rami. Armature fomiula

of P1-P4 shown in Table I.

Fifth leg (fig. 2h) with smooth seta near

insertion of free segment which bears 2 distal, 1

subdistal seta.

Male: Unknown.

Etymology. The species is dedicated to Dr David

McKinnon (Australian Institute of Marine Sci¬

ences) who studied artotrogids from southern

Australia.

Remarks. The genus Sestropontins has, so far, a

single species. Sestropontius hullifer Giesbrecht,

1899, described from the Mediterranean Sea from

a single male. More recently. Stock (1965)

described the female. It is possible to find many

dilTcrcnces between this and the new species.

Sestropontius mckinnoni shows the third endo¬

pod segment of PI and P2 with the armature I,

I +1, 3 and the third exopod segment of P4 with

eight segments. In Sestropontius hullifer the third

endopod segment of PI and P2 shows I. 2. 3 as

armature, and the third exopod segment of P4

has nine segments (Giesbrecht, 1899). Sestro¬

pontius mckinnoni has three setae on the third

exopod of the antenna instead of two as in

S. hullifer. In .S. huUiJer the antennule is

8-segmented without the ae.sthetasc while the

new species has 13 segments with an aesthetasc

on the last one.

Bradypontius Giesbrecht

Bradypontius poorei lo\\n‘i%on, sp. nov.

Figures 3-4

Material examined. Holotypc. Southern Ocean, olT

Fnderbyland, Antarctica (65’56.4()'S, 50’52.10'H), 365

m. silt and bryozoan shell, M. Norman, 15 Nov 1985,

WHOl cpibenthic sled (stn lIRD 10), NMV J48687

(female).

Description. Female’. Body length (excluding

caudal setae) 1.65 mm, greatest body width 0.80

mm, and twice as long as wide (fig. 3a). Body

shape cyelopifomi, prosome covered with sen¬

silla. cephalosome with rounded epimera. Pedi¬

gerous somites 2-4 with lateral margins pro¬

jected. Lcngth;width ratio of prosome 1.4. Ratio

of lengths of prosome:urosome 2.1.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

443

Table 1. Armature fomiulae of P1-P4 of five species of Artotrogidae.

Sestropontius mckinnoni sp. nov.

coxa

0-1

0-0

basis

1-1

1-0

exopod

1-1; 1-1; 111.1,4

I-l; I-l; 111,1,5

1-1; I-l; 111,1,5

1-1; 1-1; 111,1,4

endopod

0-1; 0-2; 1,1+1,3

0-1; 0-2; 1,2,3

0-1; 0-2; 1,1+1,2

Bradypontius poorei sp. nov.

0-1

1-1

I-l; I-l; 111,1,3

0-1; 0-2; i,2,J

0-1

1-0

1-1; I-l; 111,1,5

0-1; 0-2; 1,1+1.3

0-1

1-0

1-1; I-l; 111,1,5

0-1; 0-2; 1,1+1,3

0-0

1-0

1-1; 1-1; 111,1,5

0-1; 0-2; 1

Neohradyponlius akanthakontus sp. nov

PI 0-1

1-1

I-l; I-l; 111,1,4

0-1; 0-2; 1,2,3

0-1

1-0

1-1; I-l; 111,1,5

0-1; 0-2; 1,2,3

0-1

1-0

I-l; I-l; 111,1,5

0-1; 0-2; 1,2,3

0-1

1-0

I-l; 1-1; 111,1,5

0-1; 0-2; 1,2,2

Arctopontius novenaritis sp. nov.

0-1

1-1

I-l; I-l; 111,2,3

0-1; 0-2; 1,2,3

0-1

1-0

I-l; 1-1; 111,1,5

0-1; 0-2; 1,2,3

0-1

1-0

I-l; 1-1; 111,1,5

0-1; 0-2; 1,2,3

0-1

1-0

I-l; I-l; 111,1,4

0-1; 0,2,3

Pseudotrogus uncinalits (Brady, 1910)

0-1

1-0

I-l; I-l; 111.2,3

0-1; 0-2; 1,2,3

0-1

1-0

I-l; I-l; 111,1,2

0-1; 0-2; 1,2,3

0-1

1-0

I-l; I-l; 111,1,5

0-1; 0-2; 1,2,3

Urosome (fig. 3b) with 5 somites. Genital

double-somite 174 x 254 pm, length:width

ratio 0.7, rounded anterolaterally, smooth seta

near genital opening. Three postgenital

somites 58 X 143 pm, 71 X 125 pm and

116 X 161 pm, Icngthiwidth ratios 0.4; 0.6 and 0.7

respectively. Anal somite, caudal rami bearing

sensilla. Caudal rami elongate, 134 x 71 pm,

almost twice as long as wide, armed with

6 setae. Setae 1 absent. Lengths of setae ll-Vll,

89, 98, 281, 134 and 62 pm respectively.

Setae V broken on both rami. Setae Ill-Vl

plumose, setae II and VII smooth. Setae III

located subdistally.

Antcnnule (fig. 3c) 419 pm long not including

setae, 8-segmented. Lengths of segments mea¬

sured along posterior margins 94 pm (69 pm

along anterior margin) 119, 31, 19, 29, 25, 32

and 70 respectively. Segmental homologies and

setation as follows: 1-1; II-VIIl-8; IX-XIII-

1+spine; XIV-Hspine; XV-XVl-1; XVII-XVIII-

2; XIX-XX-2; XXI-XXVIlI-12+ae. All setae

smooth. Aesthetasc on segment XXI 162 pm

long.

Antenna (fig. 3d) 240 pm long (including distal

seta), with basis 52 pm long. Endopod 2-

segmented; first segment 44 pm long, unarmed;

second segment 56 pm long and armed with

1 plumose seta proximally, 1 seta subdistally and

2 distal plumose setae. None modified as a claw¬

like element. Exopod l-segmented, 7 pm long,

bearing 2 setae.

Oral cone (fig. 3a) produced into siphon-like

distal portion, 615 pm long, 0.4 times body

length. Mandible (fig. 3e) comprising stylet bear¬

ing distally many teeth, palp absent. Maxillule

(fig. 3f) bilobed, both lobes thin and nearly equal

in size. Inner lobe 116 pm long, amied with 2

long smooth setae and a short smooth seta, inner

margin covered with setules. Outer lobe 100 pm

long, armed with 2 long pinnate setae and a short

smooth seta.

Maxilla (fig. 3g) with syncoxa 275 pm

long; claw 400 pm long, curved distally, armed

444

R. JOHNSSON AND C.E.F. ROCHA

Figure 3. Bradypontiuspoorei sp. nov.; female holotype, a: habitus dorsal showing siphon (dotted line), b: urosome,

c: antennule, d: antenna, e: distal part of mandible, f: maxillule, g: maxilla, h: maxilliped. Scale bars: a-b: 200 urn;

c-h: 100 pm.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

445

Figure 4. Bradypontius poorei sp. nov.; female holotype, a: PI, b: P2, c: P3, d: P4. Scale bars: 50 pm.

446

R. JOHNSSON AND C.E.F. ROCHA

with seta subdistally. Maxilliped (fig. 3h) 5-

segmented; syncoxa 130 jim long, bearing short

seta on inner margin; basis 275 pm long, with

small seta medially on inner margin. Endopod

3-segmented, segments 1-3 measuring 30,42 and

105 pm long respectively. Each endopod segment

armed with single seta. Third segment with claw

190 pm long and curved distally.

Swimming legs 1^ (P1-P4, figs 4a-d) bira-

mous, with 3-segmented rami. Annature fonnula

of P1-P4 shown in Table I. Third endopodal seg¬

ment of P4 with setation extremely reduced, bear¬

ing single spine.

Fifth leg (fig. 3b) with free segment armed with

2 setae distally.

Mate: Unknown.

Etymology'. This species is named after Dr Gary

Poore, carcinologist of Museum Victoria who has

been contributing significantly to the knowledge

of the Australian crustacean fauna.

Remarks. The 21 species of Bradypontius can be

divided according to the number of segments in

the antennule which can vary from 8, 9 or more

than 10. Bradypmuius poorei has tin 8-

segmented antennule as seen in B. pichoni Stock,

1966; B. papillaltis (Scott, 1888) (Sars, 1915);

B. magniceps (Brady, 1880) (Sars, 1915) and

B. crassi.setus Kim, 1996.

Bradypontius pichoni shows the third endopod

segment of P4 armed with two short setae (Stock,

1966) while in B. poorei there is only a spine. The

remaining segments of the endopod of 5. pichoni

are unarmed while in the new species they have

the nomial annature of 0-1; 0-2. Bradypontius

pichoni has the third endopod segment of P2 and

P3 with two distal spines instead of one spine and

one seta as observed in B. poorei. The new

species also has three setae on each maxillule lobe

while B. pichoni has two setae on each. Brady¬

pontius inermis Nicholls, 1944 also has the third

endopodal segment of P4 with a single clement,

but the remaining segments of the leg arc

unarmed (Nicholls, 1944) as in B. pichoni and not

as in fl. poorei.

The mo.st distinguishing feature of Brady¬

pontius poorei is the third endopod segment of

PI with 7 elements (111-4) instead of 8 (111-5) as

seen in all other species of the genus. This feature

is one of the characteristics of the genus Arcto-

pontius. together with the 2-segmented endopod

of P4, according to Eiselt (1961). How'ever in

A. e.xpaimis Sars, 1915 there are two spines and

five setae (11-5) (Sars, 1915) and A. hanseni

Eiselt, 1986 has eight elements (111-5) (Eiselt

1986).

Neohradypontius Eiselt

Neohradypontius ukanthakontus

Johnsson, sp. nov.

Figures 5-6

Material examined. Holotype. Southern Ocean, off

Enderbyland. Antarctica (65'56.40'S, 50*52.lO'E), 365

m, M. Nortnan, 15 Nov 1985, silt and bryozoan shell,

WHOl epibcnthic sled (stn IIRD 10), NMV JI2791

(female).

De.scription. Female: Body length (excluding

caudal .setae) 1.80 mm, greatest body width 1.39

mm. and 1.3 times as long as wide (fig. 5a). Body

dorsoventrally flattened, prosome covered with

■sensilla, cephalosome and pedigerous somites

2-4 with lateral margins projected. Pedigerous

somite 4 projected beyond the posterior margin of

the genital double somite. Length:width ratio of

prosome I. Ratio of length of prosome;urosome

Urosome (fig. 5b) w'ith 5 somites. Genital dou¬

ble-somite 108 X 196 pm, lengthiwidth ratio 0.6,

slightly rounded laterally and bearing smooth seta

near genital opening, posterior margins postero-

laterally projected, reaching second postgenital

somite. Three postgenital somites 13 x 88 pm. 21

X 87 pm and 83 x 106 pm, length:width ratios

0.1,0.2 and 0.8 respectively. Anal somite bearing

sensilla. Caudal rami 52 x 42 pm, 1.2 as long as

wide and armed with 6 setae. Setae 1 absent

Lengths of setae ll-Vll, 42, 54, -, 204, 83 and 38

pm respectively. Setae IV broken in both rami.

Setae 111 VI plumose, setae II and Vll smooth.

Antennule (fig. 5c) 692 pm long, not includ¬

ing setae, 9-segmented. Lengths ol segments

measured along their posterior margins 139 pm

(111 pm along anterior margin) 44, 125, 56, 33,

56, 58, 61 and 119 respectively. Segmental

homologies and setation as follows: l-l; IM; ||l-

Vlll-12; IX-Xlll-6+spine; XlV-l+spinc XV-

XVl-2; XVIl-XVIll-2; XlX-XX-2; XXl-XXVlll-

13+ae. Third segment showing a subdivision. All

setae smooth. Aeslheta,sc on segment XXI 444

pm long.

Antenna (fig. 5d) 333 pm long (including distal

seta), with basis 89 pm long. Endopod 2-

segmented; first segment 59 pm long, unarmed;

second segment 109 pm long and armed with

I plumose seta proximally. I smooth seta sub¬

distally and 1 distal plumose setae near insertion

of short straight claw 76 pm. Exopod I-

segmented, 28 pm long, bearing 2 setae distally.

Oral cone (fig. 5a) produced into siphon-like

distal portion. 478 pm long, 0.3 times body

length. Mandible (fig. 5e) comprising stylet

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

447

Figure 5. Neohradypontius akanllmkonlus sp. nov.; t'emalc holotype. a: habitus dorsal showing siphon (dotted line),

b: urosome, c: antennule, d: antenna, e: distal part of mandible, f: maxillule. Scale bars: a: 200 pm; b-f: 50 pm.

448

R. JOHNSSON AND C.E.F. ROCHA

Figure 6. Neohradypontiiis akanthakonliis sp. nov.; female holotype, a; maxilla, b: maxilliped. c: PI, d: P2, e: P3,

f: P4. Scale bars: 50 pm.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

449

bearing distally many teeth, palp absent. Maxil-

lule (fig. 50 bilobed. Inner lobe 196 pm long,

proximal part enlarged, armed with 2 smooth

setae, inner margin covered with setules. Outer

lobe 117 pm long, amied with 2 long plumose

setae.

Maxilla (llg. 6a) with strong syncoxa 248 pm

long; claw 254 pm long, curved distally, bearing

denticles subdistally. Maxilliped (llg. 6b) 5-seg-

mcnted; syncoxa 75 pm long, bearing short seta

on inner margin; basis 176 pm long, with small

spiniform seta medially on inner margin and

setules on outer margin. Endopod 3-segmented,

segments 1-3 measuring 50, 37 and 78 pm long

respectively. First endopod segment unamied.

Second and third endopod segments armed with

single seta. Third segment with claw 120 pm

long, curved distally.

Swimming legs 1^ (P1-P4, figs 6c-f) bira-

mous, with 3-scgmented rami. Amiature formula

of PI-P4 shown in Table I.

Fifth leg (fig. 5b) with small free segment

armed with 2 setae distally.

Male'. Unknown.

Etymology. From the greck akantha, spine or

thorn and kontus, reduced, referring to the distal

short spine from the second endopod segment of

the antenna (noun in apposition).

Remarks. Neohradypontius was erected by Eiselt

(1961) to accomodate a group of species which

were previously placed in other genera of

Artotrogidae but have pleura of the third pediger-

ous somite extending backwards, at least to the

front edge of the first postgenital somite. Neo-

hrach’ponlius akanthakoutus is the only species of

the genus with a 9-segmented antennulc. All other

species have at least a 10-segmented antennule,

except N. scaber (Brady, 1910) which has eight

segments. No other species of the genus has the

second endopod segment of the antenna with the

distal element modified into a spine of similar

length to the distal and the subdistal setae.

Arctopontius Sars

Arctopontius novenuriiis Johnsson, sp. nov.

Figures 7-8

Material examined Ilolotype. Southern Ocean, off

Enderbyland, Antareliea (65'56.40'S, 50‘52.10'E), 365

m, silt and bryozoan shell, M. Norman, 15 Nov 1985,

WIIOI epibenthic sled (stn lIRD 10), NMV J47290

(female).

De.scription. Female: Body length (excluding

caudal setae) 2.32 mm, greatest body width 1.25

mm, and 1.9 times as long as wide (fig. 7a). Body

with prosome covered with sensilla, cephalosome

and pedigerous somites 2-4 with lateral margins

rounded and slightly projected. Length:width

ratio of prosome 1.3. Ratio of lengths of

prosomeiurosome 2.1.

Urosome (fig. 7b) with 5 somites. Genital dou¬

ble-somite 231 X 323 pm, lengthiwidth ratio 0.7,

slightly rounded anterolaterally, with smooth

seta, posterior margin laterally serrated. Three

postgenital somites 100 x 173 pm, 73 x 158 pm

and 142 X 192 pm, lengthiwidth ratios 0.6; 0.5

and 0.7 respectively. First postgenital somite with

posterior margin of somite serrated, second post-

genital and anal somites with posterior margin

serrated. Anal somite bearing sensilla. Caudal

rami, 138 x 85 pm, 1.6 as long as wide and armed

with 6 setae. Setae I absent. Lengths of setae

II-VII: 96, 162,481, 731, 235 and 81 pm respec¬

tively. Setae III-VI plumose, setae II and VII

smooth. Caudal rami with posterior margin

serrated and bearing sensilla.

Antennule (fig. 7c) 631 pm long, not including

setae, 9-segmented. Lengths of segments

measured along their posterior margins 125 pm

(71 pm along anterior margin) 135,27,52,31,50,

46, 50 and 115 respectively. Segmental homolo¬

gies and setation as follows: I-l; ll-VII-11; VIII-

1; IX-XIII-7+spine; XlV-l+spine; XV-XVI-2;

XVII-XVIII-2; XIX-XX-2; XXI-XXVIII-11+ae.

Aesthetasc on segment XXI 260 pm long.

Antenna (fig. 7d) 377 pm long (including distal

seta), with basis 82 pm long. Endopod 2-

segmented; first segment 55 pm long, unarmed;

second segment 93 pm long and armed with

I naked seta proximally, 2 short smooth .setae

subdistally and I distal plumose seta near inser¬

tion of long straight claw 148 pm. Exopod 1-

segmented, 18 pm long, bearing 2 setae distally

and 1 subdistally.

Oral cone (fig. 7a) produced into siphon-like

distal portion, 950 pm long, 0.4 limes body

length. Mandible (fig. 7e) comprising stylet bear¬

ing 2 groups of teeth distally, palp absent. Maxil-

lule (fig. 7f) bilobed. Inner lobe 198 pm long,

amied with long, distally plumo.se seta and short

naked seta, inner margin covered with setules.

Outer lobe 112 pm long, armed w'ith 2 stout setae,

one pinnate.

Maxilla (fig. 7g) with strong syncoxa 535 pm

long; claw 538 pm long, slightly curved distally.

Maxilliped (fig. 8a) 5-segmented; syncoxa 150

pm long, bearing short seta on inner margin; basis

362 pm long, with small seta medially on inner

margin. Endopod 3-segmentcd, segments 1-3

measuring 42, 96 and 123 pm long respectively.

450

R. JOHNSSON AND C.E.F. ROCHA

Figure 7. Arclopontiiis novenariiis sp. nov.; female holotypc, a: habitus dorsal showing siphon (dotted line),

b: urosome. c: antennule, d; antenna, e: distal part of mandible, f: maxillule, g: maxilla. Scale bars: a: 200 pm; b, g:

100 pm; c-f: 50 pm.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

451

Figure 8. Arctopontiiis novenarius sp. nov.; female holotype, a: maxilliped, b: PI, e: P2, d: P3, e: P4. Seale bars:

a; 100 pm; b-e: 50 pm.

452

R. JOHNSSON AND C.E.F. ROCHA

First endopod segment bearing 2 setae. Second

and third endopod segments armed with single

seta. Third segment with claw 200 pm long,

curved distally.

Swimming legs 1-4 (P1-P4, Figs 8b-e) bira-

mous, P1-P3 with 3-segmented rami. P4 with 3-

segmented e.xopod and 2-segmented endopod.

Armature fomiula ol'PI P4 shown in Table 1.

Fifth leg (fig. 7a) with small free segment

armed with 2 setae distally, and one subdistally

on outer margin.

Male'. Unknown.

Etymology. The specific name novenahus means

"consisting of nine", an allusion to the 9-seg-

mented antennule (noun in apposition).

Remarks. The most distinguishing features of

Arctopontius are the 2-segmented endopod of P4,

the third e.xopod segment of PI bearing three

setae on the inner margin and only two spines on

the outer margin (Sars, 1915). These characteris¬

tics were confinned by Eiselt (1961), but Arcto-

pontiiis hanseiii Eiselt, 1986, the second species

described for the genus, only shows a 2-seg-

mented endopod and the third endopod of P1 has

armature of 111,5. Arctopontius novenarius shows

the same pattern as .4. Iiaimmi in PI but the sec¬

ond endopod segment of P4 has setation of 0,2,3,

instead of a single seta as in the case of the other

two species. Arctopontius novenarius differs from

its congeners because it has a 9-scgmented anten¬

nule instead of 8-segmentcd. This difference orig¬

inates from the ancestral segment Vlll, which is

not fused with the previous segment in the new

species.

Pseudotrogus Eiselt

Pseudotrogus uncinatus (Brady)

Figures 9-10

Dystrogu.’i uncinatus Brady, 1910: 583, pi. LX figs

1- 8 (parlim).

Pseudotrogus uncinatus. —Eiselt: 1961: 324, fig. 4.

Material e.xamined. Southern Ocean, oO' Endcrbyland.

Antarctica (65"56.40'S, 50*52. lO'E), 365 m, silt and

bryozoan shell. M. Norman, 15 Nov 1985, WIIOI

epibenthic sled (stn HRD 10), NMV J47288 (2

females).

Description. Female'. Body length (excluding

caudal setae) 2.07 mm, greatest body width 1.83

mm, and 1.1 times as long as wide (fig. 9a). Body

dorsoventrally flattened, prosome covered with

sensillac, ccphalosomc and pedigerous somites

2- A with lateral margins rounded and projected.

Pedigerous somite 3 with lateral margin reaching

caudal rami. Length:width ratio of prosome 1.

Ratio of lengths of prosomeiurosome 6.1.

Urosome (fig. 9b) with 5 somites. Genital

double-somite 208 x 308 pm, length:width ratio

0.7, slightly rounded anteriorly and bearing small

smooth seta, posterolateral projections unequal,

however both projected. Right posterolateral pro¬

jection reaching distal margin of second postgen¬

ital somite, left posterolateral projection reaching

caudal rami. Left side bearing an empty ovigerous

sac. Three postgcnital somites 80 x 176 pm, 52 x

184 pm and 184 X 256 pm, lengthiwidth ratio 0.5,

0.3 and 0.7 respectively. All somites of urosome

bearing sensillac. Caudal rami slightly as long as

wide. 164 X 156 pm, and armed with 6 setae.

Setae I absent. Setae Ill-VI broken. Lengths of

setae II and VII: 44 and 76 pm respectively, both

smooth.

Antennule (fig. 9c) 668 pm long, not including

setae, 9-segmented. Lengths of segments mea¬

sured along their posterior margins 150 pm (80

pm along anterior margin) 48, 145, 57, 29,36, 36,

48 and 118 respectively. Segmental homologies

and setation as follows: 1-1; 11-1; lll-Vlll-6- IX-

XIll-5; XlV-l+spine; XV-XVl-2; XVll-XVIll-2-

XlX-XX-2; XXl-XXVlIl-IO+ae. Aesthetasc on

segment XXI 170 pm long.

Antenna (fig. 9d) 367 pm long (including distal

seta), with basis 105 pm long. Endopod 2-

segmented; first segment 68 pm long, unanned;

second segment 97 pm long and armed with

1 plumose seta proximally, 1 smooth seta sub¬

distally and 1 distal seta near insertion of straight

claw 97 pm long. Exopod I-segmented, 27 pm

long, bearing 2 setae distally.

Oral cone (fig. 9a) produced into siphon-like

distal portion. 556 pm long. 0.3 times the body

length. Mandible (fig. 9e) comprising stylet bear¬

ing distally a group of teeth, palp absent. Maxil-

lule (fig. lOa) bilobed. Inner lobe 292 pm long,

anned with 2 long setae, one distally plumose and

short naked seta, inner margin covered with

setules. Outer lobe 169 pm long, anned with 2

plumose setae.

Maxilla (fig. I Ob) with strong syncoxa 575 pm

long; claw 621 pm long, slightly curved distally

and bearing short seta subdistally. Maxilliped

(fig. lOc) 5-segtnented; syncoxa 233 pm long,

bearing short seta on inner margin; basis 479 pm

long, unarmed. Endopod 3-segmented. segments

1-3 measuring 87, 96 and 150 pm long respec¬

tively. All endopod segmenUs bearing a seta.

Third segment with claw 233 pm long and curv ed

distally.

Swimming legs 1-3 (PI-P3, figs lOd-lj bira-

rnous. P4 reduced to single process bearing 2

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

453

Figure 9. Pseiidotrogtis uncinatus (Brady, 1910); female, a: habitus dorsal showing siphon (doited line), b: P4 and

urosome, c: antennule, d: antenna, e: distal part of mandible. Scale bars: a: 200 pm; b: 100 pm; c-e: 50 pm.

454

R. JOHNSSON AND C.E.F. ROCHA

Figure 10. Pseuclotrogus uncinatu.i (Brady, 1910); female, a: maxillule, b: maxilla, e: maxilliped, d: PI, e: P2,

f: P3. Scale bars; 100 pm.

FIVE ARTOTROGID COPEODA FROM EASTERN ANTARTICA

455

setae distally (fig. 9b). Armature formula of

P1-P3 shown in Table I.

Fifth leg (fig. 9b) with small free segment

armed with 2 setae distally.

Male: Unknown.

Remarks. Pseuciotrogiis uncinaius was described

by Brady (1910) based on a specimen recorded

from the Gauss-Station during the Deutschen

Siidpolar-Expedition in 1902. Later, Eiselt (1961)

redescribed the species but many characteristics

such as P2, the endopod of PI and the antennule

setation remained unknown. This single species

shows the same body shape as P. uncinaius, and

is similar to P. sphaericus (Brady, 1910). In

P. sphaericus the margins of the third pedigerous

somite arc parallel to the urosome, and cover the

genital somite projections, unlike P. uncinaius.

Only one minor difference has been observed

between the present material and the original

description of P. uncinaius. The distal seta of

the antenna is not as long as described by

Eiselt (1961) and Brady (1910). However,

this difference is not considered specifically

significant.

Acknowledgements

I wish to thank Dr G.C.B. Poore (Museum Vic¬

toria) for providing the specimens and the

opportunity to study this material; Dr G. A.

Boxshall (The Natural History Museum) who

kindly offered his opinion about Pseudolrogus

uncinaius', and Dr D. McKinnon (Australian

Institute of Marine Sciences) who provided

valuable comments. This study was supported

by a grant from the Fundafao de Amparo a

Pesquisa do Estado de Sao Paulo (FAPESP -

98/15333-3).

References

Brady, G.S., 1910. Die marinen Copepoden der

Deutschen Siidpolar-Expedition 1901-1903.

Deutschen Sudpoiar-ExpeJiHon 11: 499-593.

Eiselt, J., 1961. Neubeschreibungen und Revision

siphonostomer Cyclopoiden (Copepoda, Crust.)

von der Siidlichen Hemisphare nebst Bemerkun-

gen Uber die Familie Artotrogidae Brady 1880.

Silztingsberichle Oeslerreichusche .-ikademie der

iVissenchaften Mathematisch-Naturwissenchaf-

tliche klasse A hie Hung I Biologische Wissen-

chaflen undErdwissenschaften 170: 315-366.

Eiselt, J., 1986. Siphonostomatoide copepoden aus der

Aktis. Cruslaceana 50(3): 295-311.

Giesbrecht, W., 1899. Die Asterocheriden des Golfes

von Neapel und der angrenzenden Meeres-

Abschnitte. Fauna und Flora des Golfes von

Neapel, Monographic 25: 1-217.

Htiys, R. and Boxshall, G.A., 1991. Copepodevolution.

Ray Society: London. 468 pp.

Johnsson, R., 2001.Two new artotrogids (Copepoda:

Siphonostomatoida) from Madeira Island,

Portugal. Hydrobiologia 453/454: 431 -440.

Kim, I.H., 1996. Copepoda of Artotrogidae (Siphonos¬

tomatoida) from the Sea of Japan. Korean Journal

of Systematic Zoolog)’ 12(4): 397-466.

McKinnon. A.D., 1988. Five artotrogid copepods

(Siphonostomatoida) from soutliern Australia.

Invertebrate Taxonomy 2: 973-993.

Nicholls, A.G., 1944. Littoral Copepoda from South

Australia, 2. Calanoida, Cyclopoida, Notodelphy-

oida, Monstrilloida and Caligoida. Records of the

South Australia Museum 8: 1-62.

Sars, G.O., 1915. /In account of the Crustacea of Nor¬

way. 6. Copepoda Cyclopoida - parts IX-X.

Bergen Museum: Bergen. Pp. 105-140, pis 65-80.

Stock, J.H.. 1965. Copepodes associes aux invertebres

des cotes du Roussillon. V. Cyclopoides Siphonos-

tomes spongicoles rares et nouveaux. Vie et Millieu

16 (IB): 295-324.

Stock, J. II., 1966. Cyclopoida Siphonostoma from

Mauritius (Crustacea, Copepoda). Beaufortia

159(13): 145-194.

Memoirs of the Museum of Victoria 59(2): 457-529 (2002)

NEW PHREATOICIDEA (CRUSTACEA: ISOPODA) FROM

GRAMPIANS NATIONAL PARK, WITH REVISIONS OF

SYNAMPHISOPUS AND PHREATOICOPSIS

George D.F. Wilson and Stephen J. Keable

Australian Museum. Sydney, NSW 2010, Australia

(buzw@austmus.gov.au; stephenk@austmus.gov.au)

Abstract

Wilson, G.D.F. and Keable S.J. 2002. New Phreatoicidea (Crustacea: Isopoda) from

Grampians National Park, with revisions of Synamphisopus and Phreatoicopsis. Memoirs of

Museum Victoria 59(2): 457-529.

The Grampians National Park, Victoria, has substantial environmental significance owing to

the diversity of endemic species restricted to this reserve. We reinforce this observation by

reporting six new species and two new genera of isopod crustaceans endemic to the

Grampians, and redescribe two previously known Victorian species representing formerly

monotypic genera. These isopods are members of the ancient suborder Phreatoicidea, and

show diverse morphologies. To demonstrate the basis for the classification of these species, we

present a phylogenetic analysis of exemplar species of most extant genera of Phreatoicidea.

Our analysis supports the sister group relationship of Phreatoicopsis and Synamphisopus. We

observe a mdimentary accessory flagellum on Ihe antennulae of both genera, but this isopod

plesiomorphy optimises on the cladograms as a reversal. Two new genera. Naiopegia gen. nov.

and Gariwerdeus gen. nov., are members of the Phreatoicidae, but are distinct from any

described laxa in this firmily. Various metazoan and protist cpibionts are commonly encoun¬

tered on these isopods. These species are described using detailed scanning electron

microscopy and inked drawings: firmily Amphisopodidae, Phreatoicopsis raffae sp. nov.,

Phreatoicopsis terricola Spencer and Hall, 1897, Synamphisopus doegi sp. nov., Synam¬

phisopus amhiguus (Sheard, 1936); family Phreatoicidae, Naiopegia xiphagroslis gen, nov.,

sp. nov., Gariwerdeus turretensis gen. nov., sp. nov., Gariwerdeus beehivensis sp. nov. and

Gariwerdeus ingletonensis sp. nov.

Table of Contents

Introduction.

Phylogenetic analysis.

Epibionts.

Systematics.

AiTiphi‘'^p<^di(lR‘' Nirhnlls

.458

.460

.461

.462

Plty^/ifnir'nnviv Snpnrpr nnH Hall

.462

Phr/^/ifnu'nnvi<: rnffne^ cn nnv

.463

Phrpntoirnp.Kis tprnrnin Spenrer and Hall .

.470

Synamphisopus Nicholls.

^yf^(iniphjKopU'^ sp nnv

.478

Synamphisopus ambiguus (Sheard).

phreafoiriclae; r’hilinn .

.487

.496

f\j(}inpp^in , , .

.497

Nciiopf^^id yipho^Kostis '^p ,.

.497

n<^v .

.502

rttrrpfpnviK nnv . .

.506

hpplii\/pnviv <i:n nnv

.512

{^nri\.vt>rrh>itK inafp/nriptiviv <sn nnv .

.519

Acknowledgments.

References.

.529

457

458

G.D.F. WILSON AND S.J. KEABLE

Introduction

We report six new endemic phreatoicidean isopod

crustacean species from The Grampians National

Park, in western Victoria, Australia. These

isopods are possibly threatened by human use of

their ecosystems because The Grampians, despite

containing unique landforms, flora and fauna, has

a long histoiy' of degradation. Anthropogenic

impacts include water impoundments, in-stream

structures, timber har\'esting, grazing and

tourism. Although the Park would not ordinarily

achieve a high conservation rating owing to this

history of human disturbance, it has much natural

significance (LCC. 1991; Doeg, 1997). A recent

study of invertebrates inhabiting riffle zones of

streams in the Grampians (Doeg. 1997) identified

at least 20 sites with conservation significance,

based on their content of rare or endemic species.

Standard limnological methods used in that study

collected only relatively large phreatoicidean

specimens of the genus Synamphisopus Nicholls,

1943 (Doeg, pers. comm.), and are unlikely to

collect the smaller groundwater-dependent taxa

reported in this paper. Taking aceount ot this

groundwater eomponent, the endemic aquatic

fauna of the Grampians National Park is more

diverse than previously documented.

A phylogenetic analysis combining the

Grampians phreatoicidean fauna with representa¬

tives of all other extant genera delimits the sys¬

tematic relationships of these taxa. This analysis

justifies the creation of two new genera for the

family Phrcatoicidae, Naiopegia gen. nov.

(monotypic) and Gariwerdeus gen. nov. (three

new species), as well as new species lor the

Amphisopodidae genera Synamphisopus and

Phrcaloicopsis Spencer and Hall. 1897. These

taxa contribute to an understanding of the phylo¬

genetic diversity of the Grampians, owing to their

ancient derivation (Wilson and Keable, 2001) and

apparently long isolation in this region.

Methods. Isopods described in this paper were

collected during a 3-day visit to the Grampians

National Park during September 1999. In all

cases, samples were taken from substrates on the

edges of streams or in springs and seeps using

small plastic hand sieves (~1 mm mesh), and field

sorted into small plastic containers. Specimens

were preserved in either sodium bicarbonate

neutralised 10% fomialin solution or 95%

ethanol. Preparation of this material for scanning

electron microscopy (SEM) included dissection

and isolation of individual limbs, ultrasonic

cleaning and C02 critical point drying. Images

were obtained using a Leo 435VP SEM equipped

with a Robinson backscatter detector, and then

saved as digital TIE files. The images were placed

into digital image plates with the background

removed. The pleopods, which often become

distorted during drying, and holotypes were illus¬

trated using light microscopy. Descriptions were

generated using the taxonomic database system

DELTA (Dallwitz, 1980; Dallwitz et al., 2()00a.

b; Wilson and Keable, 1999, 2001, 2002). The

diagnoses and descriptions below contain only

those characters that are diagnostic or that differ

among species of the same genus. Descriptions

arc based on the male holotype or paralypes;

female characters are those features that differ

from the male and differ among species in the

same genus. In the case of the monotypic new

genus Naiopegia, we have compared this species

with members of Gariwerdeus gen. nov. Methods

for creating figures and collecting morphological

data are described in Wilson and Keable (2002).

Phylogenetic analyses used PAUP* (ver.4b8.

SWofford, 2001), with the following commands to

search for shortest length trees: “hs addseq =

random nchuck = 3 chuckscore = 1 nreps = 200

randomize = trees; hs start = current nchuck = 0

chuckscore = ();”. Successive weighting (com¬

mands repeated three times: “reweight; hs”) was

used to assess the topology based on the least

homoplastic characters. Table 1 provides the tax¬

onomic data for the species in this analysis; the

data and results are reported in supplementary

information (see http://www.museum.vic.gov.au/

memoirs/index.html). Because internal structures

of the major clades are uncertain, we do not dis¬

cuss the characters and results other than as a

background for the classification of the new taxa

described below.

Table 1. Taxa used for phylogenetic analysis.

Taxa Outgroups

Source of Data

Spelaeogriphus lepidops (Spelaeogriphacea)

Gordon, 1957

Kalliapseudes ohtusifrons (Tanaidacea)

AM P26099

Tainisoptis fontinalis (Isopoda)

Wilson and Ponder, 1992

Stenasellus virei (Isopoda Asellota)

Magniez, 1975

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

459

Table 1. Continued

Taxa Phreatoicidea

Amphisopus annectam

Amphisopus lintoni

Coluhotelson joyneri

Colubotelson seaiii

Coluhotelson sp. 1 (Penstock Lagoon, Tas.)

Colubotelson sp. 2 (Uni. Tas.)

Crenisopus acinifer

Crenoicus buntiae

Crenoicus harrisoni

Crenoicus sp. nov.

Eophreatoicus sp. nov. 4

Eophreatoicus sp. nov. 6

Hyperoeclesipus plumosus

Hypsimetopus sp. nov. (near Zeehan, Tas.)

Mesacanthotelson setosus

Mesacanihotelson tasmaniae

Mesamphisopus ahhreviatus

Mesamplusoptis capensis

Metaphreatoicus australis

Metaphreatoicus lacustris

Metaphreatoicus sp.

Neophreatoicus assitnilis

Notamphisopus clunedinensis

NichoHsia kashiense

Onchotelson brevicauclatus

Paramphisopus paiustris

Paraphreatoicus relictus

Phreatoicoides sp. A

Phreatoicoides sp. B

Phreatoicoides gracilis

Phreatoicopsis terricola

Phreatoicopsis raffae sp. nov.

Phreatoicus orarii

Phreatoicus typicus

Phreatomerus iatipes

Pilharophreatoicus sp. 1

Pilbarophreatoicus platyarthricus

Synainphisopus ambiguus

Synamphisopus doegi sp. nov.

Uramphisopus pearsoni

Peludo paraliotus Wilson and Keable, 2002

Eremisopus beei Wilson and Keable, 2002

Platypyga suhpetrae Wilson and Keable, 2002

Naiopegia xiphagrostis gen. nov., sp. nov.

Gariwerdeus turretensis gen. nov., sp. nov.

Gariwerdeus beehivensis sp. nov.

Gariwerdeus ingletonensis sp. nov.

Source of Data

AM P61300

AM P8795

AM P8796

AM P54098

AM P54096

AM P54097

Wilson and Keable, 1999

Wilson and Ho, 1996

NMV JI3924, AM P4076, AM P4081

AM P6I301

AM P6I302

AM P54099

WAM 10665/6, AM P8799

AM P54I00

TMH G634/18979

AM P8767

TMH G68I,TMH G682

TMH G678

AM P3347

AM G5502

AM P52667

Chilton, 1894

USNM 54755, Acc.No. 66824; USNM 99567,

Acc.No. 45995

Zoological Survey of India C4516/1, C4517/1

TMH G3274

AM P44487

TMH G593/18930

QVM 10: 12377

QVM 10: 12267

AM P3348

this paper

Nicholls, 1944

AM P52733, AM P52734

AM P54102

AM P54I04

Knott and Halse, 1999

this paper

TMH G725

WAM C 25051-25052; AM P60532-60533

WAM C 25049-25050, AM P60527-60531

WAM C 25053-25054; AM P60537-60539

this paper

460

G.D.F. WILSON AND S.J. KEABLE

Abbreviations. NMV, Museum Victoria, Mel¬

bourne; AM, Australian Museum, Sydney; QVM,

Queen Victoria Museum, Launceston; SAM,

South Australian Museum, Adelaide; TMH, Tas¬

manian Museum and Art Gallery, Hobart;

USNM, United States National Museum, Wash¬

ington, DC; WAM, Western Australian Museum,

Perth; bl, body length; GPS, global position¬

ing satellite fix; ind., individuals, specimen or

specimens.

Phylogenetic analysis

Parsimony analysis of the data found 84 trees of

length 694 (see supplementary information).

When subjected to successive weighting, only

three trees result from the analysis. Figure 1

shows the strict consensus of both the parsimony

and weighted parsimony analyses. The parsimony

consensus shows large polytomies for the taxa of

the Amphisopodidae, the Western Australia and

Indian species of the Hypsimetopodidae and

various clade levels within the Phreatoicidac.

Polytomies also appear in both consenses within

the genera Mesamphisopus, Crenoicus and Gari-

werdeus gen. nov. because the analysis does not

include features that are diagnostic at a species

level within those taxa. The presence of

Platypyga suhpetrae Wilson and Keable, 2002

appears to cause most of the discordance among

' Spelaeogriphus lepidops

• Kalliapseudes obtusifmns

Tainisopus fontinalis

Stenasellus verei

■ H^medesipus plumosus

• Nichollsia kashiense

■ ^taeogriphus lepidops

• Kalliapseudes obtusifmns

Pilbarophreatoicus n.sp.

Pilbaro. platvarthricus

Hypsimetopus sp.

Phreatoicoides sp. B

Phreatoicoides sp. A

Phreatoicoides gracilis

Crenisopus acinifer

Peludo paraiiotus

Platypyga subpetrae

Eremisopus beei

Eophreatoicus sp. 4

Eophreatoicus sp. 6

Paramphisopus palustris

Phreatomerus sp.

Amphisopus annectans

Amphisopus lintoni

Phrealolcopsis terricola

Phreatoicopsis raffae n.sp.

Synamphisopus ambi^uus

Synamphisopus doegi n.sp.

E Mesamphisopus abbreviatus

Mesamphisopus n.sp.

Mesamphisopus capensis

- Notamphisopus dunedinensis

I- Uramphisopus pearsoni

L|- Onchotelson brevicaudatus

l_i— Mesacanthatelson setosus

*— Mesacanthoteison tasmaniae

Paraphreatoicus rellctus

Coluboteison sp. 1

Colubotelson sp. 2

Coluboteison joyneri

^ Coluboteison searii

E Metaphreatoicus australis

Metaphreatoicus sp.

Metaphreatoicus lacustr/s

■ Haiopegia xiphagmstis n.sp.

Gariwerdeus turretensis n.sp.

Gariwerdeus ingletonensis n.sp.

Gariwerdeus bwhivensis n.sp.

E Crenoicus buntiae

Crenoicus n. sp.

Crenoicus harrisoni

Neophreatoicus assimilis

Phreatoicus orarii

Phreatoicus typhus

j— Tainisopus fontinalis

'— Stenasellus verei

Hypsimetopus sp.

Phreatoicoides gracilis

Phreatoicoides sp. A

Phreatoicoides sp. B

Hyperoedesipus plumosus

Nichollsia kashiense

Pilbamphreatoicus n.sp.

Pilbamphreatoicus platyarthricus

Crenisopus acinifer

* Mgs amphisopus abbreviatus

■ Mesamphisopus n.sp.

~ Mesamphisopus capensis

~ Peludo paraiiotus

Eremisopus beei

Eophreatoicus sp. 4

Eophreatoicus sp. 6

Platypyga subpetrae

Phreatomerus sp.

Paramphisopus palustris

Amphisopus annectans

Amphisopus lintoni

Phreatoicopsis terricola

Phreatoicopsis raffae n.sp.

Synamphisopus ambiguus

Synamphisopus doegi n.sp.

Notamphisopus dunedinensis

T- Metaphreatoicus australis

Metaphreatoicus lacustris

~ Coluboteison Joyneri

Coluboteison searii

Coluboteison sp. 2

Coluboteison sp. 1

Paraphreatoicus relictus

Metaphreatoicus sp.

Uramphisopus pearsoni

Onchotelson brevicaudatus

Mesacanthoteison setosus

Mesacanthoteison tasmaniae

Naiopegia xiphagrostis n.sp.

Gariwerdeus turretensis n.sp.

Gariwerdeus Ingletonensis n.sp

Gariwerdeus beehivensis n.sp.

Crenoicus buntiae

Crenoicus n.sp.

Crenoicus harrisoni

Neophreatoicus assimilis

Phreatoicus orarii

Phreatoicus typicus

Figure I. A, strict consensus of parsimony analysis. B, strict consensus of successive weighted parsimony analysis.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

461

equally parsimonious trees. In an Adams consen¬

sus of the 84 trees (see supplementary infomia-

tion), Platypyga appears in a polytomy with two

resolved Amphisopodidac clades, indicating that

this species causes the collapse of the amphis-

opodid clades in the strict consensus tree. Delet¬

ing this ta,xon reduces the tree set from 84 trees to

only 9 much more resolved trees (length 679) (see

supplementary infonuation). In the trees from all

of these analyses (parsimony, weighted parsi¬

mony, Platypyga removed), general features

appear consistently - two major clades corre¬

sponding to the families Hypsimetopodidae

(sensu Wilson and Keabic, 2001) and Phreatoici-

dae, with Crenisopus Wilson and Keable, 1999

emerging on the branch between Amphis¬

opodidac and Hypsimetopodidae. Stable super¬

specific clades occur within the families:

Hypsimelopus + P/irealoicnicies;

Amphisopus + Phreatomems + Paramphisopus'^

Phreatoicopsis + Synaniphisopiis\

Eremisopiis + Eophreatoicus\

Crenoicus + New Zealand taxa Phreatoicus

and Neophreatoicus;

Gariwerdeus + Naiopegia.

The positions of these stable clades vary within

the major clades depending on the composition of

the analysis; additional species or new characters

often change the relationships. For the purposes of

this paper, however, these results are stable among

all trees. Species of Gariwerdeus gen. nov. are

monophyletic, with Naiopegia gen. nov. as its sis¬

ter group within the Phreatoicidae. The genera

Phreatoicopsis and Synamphisopus are mono¬

phyletic, and together fomi a monophyletic clade

within a less well-defined amphisopodid clade.

This latter result is surprising because Phreato¬

icopsis superficially resembles no other phrea-

toicidean genus. Nevertheless, Nicholls (1943)

originally assigned the genera Synamphisopus

and Phreatoicopsis, with Eophreatoicus Nicholls,

1926, Protamphisopus Nicholls, 1943 (a Triassic

fossil) and Uramphisopus Nicholls, 1943 to

the heterogeneous amphisopodid subfamily

"Phreatoicopsinac" [sic]. The relationships of

P. rotamphisopus are under study (research in

progress), but Eophreatoicus is the sister group to

Eremisopiis Wilson and Keabic, 2002 (fig. I A),

and clearly not as closely related to Synam¬

phisopus and Phreatoicopsis. Uramphisopus is a

member of the Phreatoicidae (Wilson and Keable,

2001). Moreover, analyses of I2S and 16S rDNA

by Wetzer et al. (20()1) also support the sister

group relationship of Phreatoicopsis and Synam¬

phisopus. Character states shared by the two

genera, other than those discussed below

(Phreatoicopsis remarks), include a large spine

on the propodal palm of the male first pereopod.

but variation among the species weakens the sup¬

port of the relationship. The rudimentary second

flagellum on the antennula is observed in both

genera (Phreatoicopsis: figs. 3C, lOF; Synam¬

phisopus: 18D), but this isopod plesiomorphy

optimises on the cladograms as a reversal sup¬

porting the sister relationship of the two genera.

Our previous analysis of the Phreatoicidea (e.g.,

Wilson and Johnson. 1999; Wilson and Keable,

2001) did not find this sister relationship because

several synapomorphies of Synamphisopus and

Phreatoicopsis were omitted from the analysis,

and one feature (depth of pleonite pleurae) was

incorrectly scored. The relative uncertainty ol

w ithin-family relationships that remain in our anal¬

yses prevents the use of Nicholls’ (1943, 1944)

subfamily categories in the current classification.

Epibionts

Practically all large specimens ol Phreatoicopsis

are found with Temnocephala (Platyhelminthes)

crawling on their stemites and ostracodes on their

pleopods, as in many Australian crayfish

(Parastacidae). All species of Gariwerdeus gen.

nov., Naiopegia gen. nov. and to a lesser extent

Synamphisopus and Phreatoicopsis have sub¬

stantial populations of protozoan epibionts grow¬

ing on their limbs and somites. In some body

positions, the protist epibiont burden is substan¬

tial (e.g., pereopods - fig. 54A, or mouthparts —

fig. 44E). Other phreatoicidcans have been

recorded with various types of sessile protists

(Kane, 1969; Clamp, 1991; Femandez-Leborans

and Tato-Porto, 2000; O’Donoghue and Adlard,

2000; Wilson and Keable, 2002). Clamp (pers.

comm.) has recorded lagenophryid peritrichs on

all phreatoicidean families: Metaphreatoicus aus¬

tralis (Chilton, 1891), Colubotelson searli

Nicholls, 1944, C. Joyneri (Nicholls, 1926),

C. chiltoni (Sheppard, 1927), Mesacanthotetson

tasmaniae (Thomson, 1894), Phreatoicoides

longicollis Nicholls, 1943, and Phreatoicopsis

terricola Spencer and Mall, 1897. Some epibionts

may have preferences for host attachment sites, as

has been obseix'ed in the unrclalcd Asellus aquati-

ctis (Cook et al., 1998), and tvso different genera

may occur on dilTerent locations on the body.

Dome-shaped Lagenophryidae peritrichs are

found on the pleopods and others on the le^,

while the stalked peritrich Vorticella (fig- 43G)

and the suctorian ciliates (fig. 49E) appear to pre¬

fer the external surfaces of the limbs and body.

How these epibionts affect their hosts is

unknown.

462

G.D.F. WILSON AND S.J. KEABLE

Systeniatics

Amphisopodidac Nicholls

Amphisopidae Nicholls, 1943: 25.

Remarks. Our analyses of character distributions

among the Phreatoicidea (results above, Wilson

and Keable, 2002) has identified an assemblage

of taxa that may be assigned to the family

Amphisopodidae. The unweighted analyses do

not provide strong support for the family, and

some taxa do not match the familial characters

discussed in Wilson and Keable, 2002. Platypyga

Wilson and Keable, 2002 is only weakly assoc¬

iated with the Amphisopodidac, although most

character weighted analyses of the data find that it

belongs to this clade (e.g., successive weighting

results outlined above). Therefore, we do not pro¬

vide a new classification or diagnosis of the

Amphisopidae at this time because the status of

this family may be questioned. Although eyes fre¬

quently are not expressed in phreatoicideans and

thus are homoplasious for phylogenetic analyses,

all species currently assigned to this family have

eyes, if only small spots as in Phreatoicopsis.

While we do not regard this presumed plesio-

morphy as significant, this feature may assist in

identification (i.e., blind species probably belong

to another family).

Phreatoicopsis Spencer and Hall

Phreatoicopsis Spencer and Hall, 1897: 12.—

Sheppard, 1927: 117.—Nicholls. 1943: 112.

Type species. Phreatoicopsis lerricola Spencer

and Hall. 1897, by monotypy.

Diagnosis. Typhlosole well developed, ventral

invagination forming double spiral in cross

section. Pleotelson posterior margin truncate, cir¬

cular in posterior view, with temiinal anus, not

reflexed; lateral lobes absent; dorsal uropodal

ridge curx ing strongly and extending posteriorly

from uropods on pleotelson margin. Antennula

article 3 rudimentary second flagellum present.

Mandible spine row on round pedunculate projec¬

tion. Maxillulu lateral lobe distal margin with

many (24-25) robust setae, distal setal row

separated by gap from other setal rows; inner lobe

narrow and tapering, with 3 pappose setae dis-

tally. Pereopod I dactylus ventral margin proxi¬

mal tooth present; merus dorsal margin projection

spine-like and pointed. Pereopods ll-VIl pro-

podus without articular plate; pereopod IV of

male simple, not prehensile; pereopods V-Vll

basis dorsal ridge not distinctly separated from

basis shaft. Pleopods protopods I-11 lateral

epipod linear; pleopod II endopod appendix mas-

culina proximal half of shaft solid and rod-like,

indented in ventral cross section, with several

large subterminal denticles around lateral to

medial and dorsal surfaces. Uropod rami distal

tips pointed.

Remarks. Phreatoicopsis species, in addition to

being among the largest of the phreatoicideans

and living in semiterrestrial habitats, have other

unusual features. Live animals have a light, some¬

times pearly cuticle, with bands or patches of

bright yellow on some dorsal surfaces, most

obviously the head. The legs of Phreatoicopsis

are unusually thin, lacking the typical amphisopo-

didan dorsal margin plates on the pcreopodal

bases, and the large pleonal pleurae possibly form

a respiratory chamber for pleopods. Setae on the

body and limbs are few, short and typically

robust. The small setae on the margins of the

pleopods are minutely plumose. The pleotelson

lacks any of the typical embellishments seen in

aquatic phreatoicideans; it is smooth, simply con¬

structed and opens posteriorly, with a fringe of

setae on the posterior margin. Interestingly, the

lateral outline of the Phreatoicopsis pleotelson

(e.g., figs 2, 9C, 16B) resembles that ol' Protam-

phisopus, whieh Nicholls (1943) placed in his

rather heterogeneous subfamily Phreatoicopsinae.

Hermaphroditism may be an important feature of

the Phreatoicopsis reproductive system, although

the reasons for the apparently varying proportions

of interse.xes remain to be determined.

As discussed above, Phreatoicopsis is the sister

group to Synamphisopus. This relationship is

based on the following putative synapomorphies:

the highly developed typhlosole in the hindgut;

an unusually high number of robust setae on

the maxillula lateral lobes (Phreatoicopsis - fig.

5B; Synamphisopus - figs 20B-C); blunt denti¬

cles on the distal tip of the appendix masculina

(Phreatoicopsis - fig. 16D; Synamphisopus - figs

23C-E, 311-J); and a pedunculate mandibular

spine row (Phreatoicopsis - figs 4D-E, H. J;

Synamphisopus - figs 19B, H. J). The spine row

structure is distinct from that of the Phreatoicidae

because the bifurcate spines are fused basally so

that the two rami of each spine appear to be

independent, giving an open appearance to the

centre of the spine row. The spine row Synam¬

phisopus is somewhat less modified tlian in

Phreatoicopsis and flattened dorsoventrally,

appearing more simlar to the typical spine row of

other amphisopodids.

Phreatoicopsis specimens from the Grampians

were previously identified as P. terricola Spencer

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

463

and Hall (first noted by Raff, 1912). Our examin¬

ation of this morphologieally conservative genus

indicates that populations from the Grampians

represent a new species, as suggested by Nicholls

(1943).

Phreatokopsis raffae sp. nov.

Figures 2-9

Phreatokopsis terricola .—Raff, 1912; 70, pi. 5.—

Nicholls, 1943: 113, figs 27, 28 (part, material from

vicinity of the Grampians is not Phreatokopsis

rerr/co/rt Spencer and Mall, 1897).

Material examined. Hololype. Victoria (Vic.), Flatrock

Crossing, Gleneig River Road, Grampians National

Park, 37°09.77'S I42''26.59'E (GPS), soil among roots

of ferns and under sphagnum moss on side of seep, col¬

lected by hand and spoon, R, Wetzer, S. Keable and

G. Wilson, 20 Sep 1999, VIC-87, NMV J40730 (male

bl 51.9 mm, ethanol preserved).

Paratypes. All lots collection details as for holotype.

AM P61250(1 male bl 32.5 mm, 2 females bl 45.7/51.4

mm, 1 juvenile female bl 29.1 mm, I hemiaphrodite

(with penes, no appendix masculina, with oostegite

buds) bl 32.7 mm, 1 indeterminate Juvenile bl 19.6

mm), AM P61436 (1 male bl 40.7 mm, dissected

for illustration, description and SEM), AM P61437

(I female bl 46.8 mm, dissected for description and

SEM).

Other material. Vic., The Grampians, no other data.

AM P6I251 (9 ind. ex NMV). Vic., The Grampians: N

oftheDivide(37°l7'S, I42“33'E),21 Dec 1934,NMV

J44871 (>30); Vic., S of the Divide (37°I7’S.

142°33'E), I. Mitchell (Stawell, Vic.) per A. Chisolm.

25 Dec 1934, NMV J44873 (>30); Vic., Bellfields,

37°I7'S, I42“33’E, J. Clark. 14 Aug 1935, NMV

J44882 (5); Vic., Swamp at head of Wannon R.

(37°19'S. 142°3rE). J. Clark. 14 Aug 1935, NMV

J44892 (7); Vic., Swamp at head of Wannon R.

(37°I9'S, 142°3rE), J. Dawson, 04 Sep 1935, NMV

J44893 (9); Vic.. N. McCance, Jul 1961, NMV J21805

(I); Vic.. Glenelu R. (37"11’S, 141 “43'E), A. Neboiss,

16 Dec 1966. NMV J44885 (1); Vic.. 37“I7'S,

I42“33'E, R. Veerman. December 1968, NMV J44898

(I); Vic., Mt William (37“I3'S, I44“48'E), C. McCub-

bin, 28 Apr 1971, NMV J44883 (1): Vic.. Glenisla

Range (37“14'S, I42“1I'E). A. A. Calder, 04 Mar

1976, NMV J44896 (3) and NMV J44895 (2); Vic., Mt

William, 100 m S. of summit, 27 Aug 1978, M.S.

Harvey, NMV J48353 (I female): Vic.. Jimmys Creek,

25 Aug 1983, J. Baldwin. NMV J48354 (2); Vic., near

Jimmys Creek and Dunkeld Road (37“22'S. I42“3I E),

R. Duggan, 26 Jan 1987, NMV J44868 (5); Vic.,

37“I7’S, 142°33'E, H. Clark, NMV J44869 (57).

Figure 2. Phreatokopsis rajfae, sp. nov. Holotype (NMV J40730), lateral view. Scale bar 5 mm.

464

G.D.F. WILSON AND SJ. KEABLE

Figure 3. Phreatoicopsis raffae, sp. nov. Paratype male (AM P61436). A, head, dorsal view. B, antennula, antenna

proximal articles, dorsal view. C, antennula article 3 rudimentary second flagellum. D, head, lateral view with

antennula and antenna proximal articles. E, antennula, distal tip. F-G, antenna flagellum, di,stal and proximal

articles. Scale bar 1 mm.

Vic., (Vial label gives locality as "From Greener,

West Australia", A second label in the vial says

"The locality of these specimens is doubtful! Prob¬

ably Western Victoria") l.R. McCann, 9 November

1982, NMV J44890 (19). No label data: J44782 (2),

J44888 (3)

Etymology^ This species is named after Janet W.

Raff, who first recorded it from the Grampians

(Raff, 1912).

Diagnosis. Pleotelson length subequal to width in

dorsal view, dorsal length 0.96 width. Left

mandible incisor process dorsal margin with

fourth sloping cusp. Maxillula lateral lobe with 4

robust setae in distal row. Pereopod VII basis

dorsal ridge with robust setae. Uropod protopod

dorsomedial ridge not projecting beyond distal

margin, linear.

Description, including all adult forms. Coloration in

life head with large patch of yellow, posterior margins

of pereonites 6-7, pleonites 1-2 or 2-3 with yellow

transverse stripe, body otherwise grayish, white pereo-

pods, tiny black eye; in 95% ethanol uniform cream,

eye black.

Head (figs 3A, D) width 0.73 pereonite 1 width. Eyes

maximum diameter 0.12 head depth.

Pleonites (fig. 2) 1^ width 0.78 composite length in

dorsal view.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

465

Figure 4. Phreatoicopsis raffae, sp. nov. Paratype male (AM P6I436). A, paragnaths. B-G, left mandible.

H-J, right mandible. Scale bar 0.5 mm.

466

G.D.F. WILSON AND S.J. KEABLE

Figure 5. Phrealoicopsis raffae, sp. nov. Paratype male (AM P61436). A-C, maxillula. D-E, maxilla.

F-G, maxilliped. Scale bar 1 mm.

Pleolelson (figs 9A, C) lateral length 0.14 body

length; depth 1.95 pereonite 7 depth; posterior margin

with 72 setae (approximately, including anterior ventral

margin near insertion of uropod); ventral margin

anterior to uropods with 6 setae.

Aniemmla (figs 3B-C) length 0.07-0.1 body length,

with 13-14 articles. Article 5 length 1.5 width. Article

6 length 1.2 width. Several liny aestheuiscs on article 11

to terminal article. Terminal article length 0.8 width.

Antenna (figs 2, 3F-G) length 0.38 body length. Flagel¬

lum length 0.43-0.62 total antenna length, with 18-22

articles.

Afont/i/ieW clypeus width 0.52 head width. Mandible

(figs 4B-J) palp length 1.12 mandible length; article 3

with 7 .setae, setae denticulate (minutely). Left spine

row with 17 spines (basally fused, therefore total

probably representing count of both sides of approx¬

imately 8 bifurcate spines), additional spines between

pedunculate projection and molar absent. Molar process

length subcqual to width; fine simple spines forming

posterior row (short). MiLxdbda (figs 5A-C) medial

lobe width 0.5 lateral lobe width; with 8 'accessory

setae’. Lateral lobe distal margin with 24 smooth robust

setae.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

467

Figure 6. Phreatoicopsis raffae, sp. nov. Paratype female (AM P61437), paratype male (AM P6I436). A-B, female

pereopod I. C-D, male pereopod I. Scale bar 1 mm.

Maxilla (figs 5D-E) medial lobe width 1,33 outer lat¬

eral lobe width. Outer lateral lobe length subequal to

inner lateral lobe (ratio 1.02). Maxilliped (figs 5F-G)

endite distal tip with appro.ximately 20 subdistal

biserratc setae on ventral surface; medial margin with 6

coupling hooks on left side; dorsal ridge with 17 large

distally not noticeably denticulate plumose setae. Palp

article 4 length 0.73 width; article 5 length 1.75 width,

length 0.75 article 4 length.

Pereopod 1 (fig. 6) dactylus length subequal to palm

length. Propodus length 0.88 width; dorsal margin with

8 minute setae (e.xcluding distal group). Propodal palm

with 6-8 broad based setae. Basis length 2.0 width,

dorsal ridge with approximately 13 minute setae scat¬

tered along ridge. Pereopod IV (figs 7A-B) length 0.38

body length. Propodus length 4.5 width, with 4 broad

based setae on ventral margin (5 submarginally). Car¬

pus length 0.16 pereopod length, with 4 broad based

setae on ventral margin in male (7 submarginally, I dis¬

tinctly larger than others). Ischium posterodisial margin

with 4 setae in male (8 submarginally). Basis length

3.25 width.

Penes length 0.37 body width at pereonite 7.

Pleopod (fig. 8) I endopod length 0.65 exopod length.

PleopodW endopod length 0.65 exopod length. Pleopod

III exopod distal article length 0.32 exopod length;

endopod length 2.04 width, 0.68 exopod length. Pleo¬

pod IV exopod length of distal article 0.33 exopod

length; endopod length 1.75 width. Pleopod V endopod

length 1.72 width. Pleopods protopod I with 16 simple

and minutely serrate setae along length of lateral mar¬

gin; protopod 11 with 8 simple and minutely serrate

setae along length of lateral margin, 2 submarginally.

Pleopod I exopod medial margin convex - divergent

from lateral margin (weakly concave proximally). Pleo¬

pod 11 endopod appendix masculina with 8 minute setae

on margin.

Llropod (figs 9B. D, E-G) total length 0.72 pleotel-

son length. Protopod length 1.89 width, 0.31-0.44 uro-

pod total length; dorsomedial ridge length 0.84 endo¬

pod length. Endopod with 3-5 robust setae. Exopod

length 0.62-0.67 endopod length, with 3 setae.

Distrlhution. Central and southern parts ol The

Grampians National Park, in wetlands or swamps

associated with the Wannon and Gleneig Rivers,

Victoria.

Remarks. The male and female character states do

not appear to be correlated with body size pos¬

sibly owing to hermaphroditism in this species, so

we have not distinguished the sexes in the above

468

G.D.F. WILSON AND S.J. KEABLE

Figure 7. Phrealoicopsis raffae, sp. nov. Paratypc male (AM P61436), paratype female (AM P61437). A, male pere-

opod IV. B, female pereopod IV. C. male pereopod Vll. D, male pleopod 11 appendix masculina. E, male pereopod

VII proximal articles, with penes. Scale bar 1 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

469

Figure 8. Phreatoicopsis raffae, sp. nov. Paratype male (AM P61436). A-E, pleopods I-V. Setae on margins are

minutely plumose. Scale bar 1 mm.

description. Species of Phreatoicopsis can be

identified using the shape of the uropod protopod

dorsomedial margin: straight or only slightly

curving dorsally and not projecting beyond inser¬

tions of rami {P. raffae, figs 9D, G) versus curv¬

ing dorsally and projecting beyond insertions of

rami {P. terricola, figs 16C, F). Robust setae on

dorsal margin on the basis of pereopod Vll in

Phreatoicopsis raffae sp. nov. (tlg.7C) as

opposed to fine setae in P. terricola will distin¬

guish adults of the two species, but this feature

may not be useful for specimens of 35 mm or less.

470

G.D.F. WILSON AND S.J. KEABLE

Figure 9. Phreatoicopsis raffae. sp. nov. Paralype male (AM P61436). A-D, pleotelson and uropod. dorsal,

posterior and lateral views. E-G, uropod ventral and medial views. Scale bar 1 mm.

Phreatoicopsis tcrricola Spencer and Hall

Figures 10-16

Phreatoicopsis lerricola SpmccT md Hall, 1897; 12,

pis. 3-4.—Raff, 1912: 70, pi.5 (part, material from

vicinity of Otway Range).—Nicholls, 1924: 98.—

Nicholls, 1926; 203.—Sheppard. 1927: 117.—Barnard,

1927: 160.—Nicholls, 1943; 113, figs 27, 28 (part,

material from vicinity of Otway Range).

Not Phreatoicopsis tcrricola. —Raff. J.W., 1912: 70.

pl.5.—^Nicholls, 1943: 113. figs 27, 28 (part, material

from vicinity of The Grampians is Phreatoicopsis raf¬

fae sp. nov.).

Type material. Probable syntypes. Victoria, “Banks of

Upper Gellibrand River, in burrows (W.II.F. Hill)”

(Spencer and Hall, 1897; 13), 38°33'S I43°39'E (esti¬

mated from map), specimens lost. Nicholls (1943)

reported examining specimens from “Museums of

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

471

Figure 10. Phreatoicopsis lenicoki. Hermaplirodite (AM P61438). A, antenna. B-C, antenna flagellum, distal and

proximal artieles. D, antennula, antenna proximal articles, dorsal view. E, antcnnula distal articles. F, antennula

article 3 rudimentary second flagellum. Scale bar I mm.

Melbourne and Adelaide” but apparent types have not

been found. Museum Victoria (G. Poore, pers. comm.)

has specimens of P. raffac sp. nov. (see above), that

account for some but not all specimens reported by

Nicholls.

Material examined. Vic.. Otway Range, 200 m west of

Benwerrin-Mount Sabine Track, 0.5 km N of Delaney

Road, 38°31.90'S I43°50.76'E (GPS), from soil 20cm

deep under fern tree root mat in bank of stream, shovel

and hand. G. Wilson, R. Wetzer and S. Keable, 25 Sep

1999. preserved in 95% ethanol, VIC-101. AM P6I252

(4 hermaphrodites bl 26 (no appendix masculina, pene

bud only on one side)/26 (with appendix masculina)/

30.5 (no appendix masculina)/40.5 (no appendix mas¬

culina) mm. 4 females bl 24.2/29.8/30.2/50.9 mm, I

juvenile female bl 24.7 mm), AM P61438 (herma¬

phrodite bl 46.6 mm, dissected for illustration, descrip¬

tion and SEM, collection details as for P61252), AM

P6I439 (hermaphrodite bl 50.2 mm partially dissected

for description and SEM, collection details as lor

P6I252); Vic., Otway Range, from 900 m SW of

Cowley Track. 38°33.36 S, 143“50.48'E (map), dug

from oval buiTows in damp sediment under tree fern

roots, no free water observed in substrate, W.F. Ponder

and G.D.F. Wilson. 15 Jul 1991, Stn VI9. AM P54I01

(4 juveniles possibly males (pene buds but no appendix

masculina) bl 16.6/18.2/25/25.9 mm, 9 males bl 21.1

(penes and appendix masculina not lully devel-

oped)/25.5/25.7/27.5/28.2/28.6/35.7/41.2 (dissected for

description)/53,8 mm, I hermaphrodite (small penes,

no appendix masculina, oostegite buds on anterior

pereopods) bl 25 mm, 2 hermaphrodites bl 36 (dissected

for description)/46.6 (no appendix masculina) mm);

Vic., near Lavers Hill, Otway Range, 38°41 S

143°24'E (estimate), pit trap, A. Fraser, 10 Feb 1972,

472

G.D.F. WILSON AND S.J. KEABLE

Figure II. Phreatoicopsis lerricola. Hermaphrodite (AM P6I439) A-B, G-l, hermaphrodite (AM P61438)

C-F. A. paragnaths. B, left mandible. C-E, left mandible. F, right mandible. G-l, right mandible. Scale bar 1 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

473

Figure 12. Phreatoicopsis leiricola. Hermaphrodite (AM P6I438) A-E, hermaphrodite (AM P6I439) F-G. A-C,

maxillula. D-E, maxilla. F-G, maxilliped. Scale bar I mm.

AM P310()4(l ind.); Vic., Forrest (38°31 S, I43°43'E),

Davey, HW. 26 Feb 1925, NMV J44880 (I); Vic.,

Upper Gellibrand Gorge, near Forrest (38°33’S,

143°39'E), ‘CWB’, 17 Dec 1946, NMV J44894 (8);

Vic., Otway. Beech Forest (38°27’S, I43°58'E),

W.II.F. Hill. l4Sep I897,NMVJ44875(4); Vic., Cape

Otway, Beech Forest (38°52'S, l43°3rE), H.P. Ash¬

worth, Apr 1897, NMV J44877 (3 degraded, possibly

dried at one time); Vic., Beech Forest (this record

included "Grampians” in phace name) W.H.F. Hill (col-

leetor of the syntypes), II Nov 1897, J44884 (16 -

tanned colour); Beech Forest NMV J44876 (1); Vic.,

Otway Ranges, G. Milledge, II Apr-14 Jun 1995;

Phillips Track 0.5 km N of Triplet Falls (38°40'S

I43°29'E), pitfall traps. G. Milledge, Nolhofagiis ctm-

iiinghaniii forest, J48355, (32); Vic., Young Creek Rd

0.4 km NW of Triplet Falls, (38“40'S 143'’29’E), pitfall

traps, Eiicah'ptus sp. forest. J48356 (6); Vic., Cobden

(38°20'S, r43°04'E), N. McCance, Sep 1963. NMV

J44886 (I); Vic., Apollo Bay (38°46'S. I43“40'E),

Field Naturalists Club of Victoria, Dec 1904, NMV

J44874 (28); Vic., Maits Rest, 10 km W of Apollo Bay,

Otway Ranges (38°45’S, I43°34'E), in Nolhofagus

cunnighumii forest, Pitfall Trap, G, Milledge, P. Lilly-

white and C. MePhee, 30 Oct 1991, NMV J4488I (2).

No collection locality data: NMV J44878 (1), NMV

J44879 (1), NMV J44848 (1 damaged specimen).

P. lerricola variant (see Remarks). Vic., Upper

474

G.D.F. WILSON AND S.J. KEABLE

Figure 13. Phreatoicopsis lerricola. Hermaphrodite (AM P6I438). A-C, pereopod 1. Scale bar 5 mm.

Gellibrand Gorge, near Forrest (38°33'S. 143'’39'E),

‘CWB’. 17 Dec 1941, NMV J44891 (22, largest is~30

mm); Vic., Forrest (38°3rS, I43°43'E). Wilhelms, Jul

1948, NMV J44887 (10, large.st is ~40 mm).

DUignosix. Pleotelson length greater than width in

dorsal view, dorsal length 1.22 width. Left manJihle

incisor process dorsal margin lacking fourth sloping

cusp. Maxillula lateral lobe with 5 robust setae in distal

row. Pereopod VII basis dorsal ridge lacking robust

setae. Uropod protopod dorsomedial ridge distinctly

projecting posteriorly beyond distal margin, concave.

Description based on hermaphrodite. Coloration in

life, head with patch of yellow, posterior margins of

pereonites 6-7. pleonites 1-3 with yellow transverse

stripes, body otherwise grayish, white pereopods, tiny

black eye; in 95% ethanol uniform cream, eye black.

Head width 0.82 pereonite 1 width. Eyes maximum

diameter 0.09 head depth.

Pleonites 1 -4 width 0.60 composite length in dorsal

view.

Pleotelson (figs 16A-C, F) lateral length 0.11 body

length; depth 1.88 pereonite 7 depth; posterior margin

with 66 .setae (approximately, including anterior ventral

margin near insertion of uropod); ventral margin ante¬

rior to uropods with 10 setae.

Antenniila (figs lOD-E. F) length 0.10-0.14 body

length, with 12-13 articles. Article 5 length 1.2 width.

Article 6 length 1.0 width. Several tiny aesthetascs on

article 9 to terminal article. Terminal article length 0.55

width. Antenna (figs 10 A-C) length 0.3 body length.

Flagellum length 0.43-0.46 total antenna length, with

24-26 articles.

Mouthfield clypeus width 0.48 head width. Mandible

(fig. 11) palp length 1.06 mandible length; article 3 with

8 setae, setae smooth. Left spine row with 21 spines

(basally fused, total probably representing count on

both sides of approximately 10 bifurcate spines), total

count including 1 spine on margin between pedunculate

projection and molar. Molar process longer than wide;

spines absent. Maxillula (figs 12A-C) medial lobe

width 0.39 lateral lobe width; with 7 "accessory setae".

Lateral lobe distal margin with 25 smooth robust setae.

Maxilla (figs 12D-E) medial lobe width 1.64 outer

lateral lobe width. Outer lateral lobe longer than inner

lateral lobe. Ma.xilliped (figs 12F-G) endite distal tip

with 38 siibdistal biserrate setae on ventral surface

(approximately); medial margin with 7 coupling hooks

on left side; dorsal ridge with 26 large distally denticu¬

late plumose setae (approximately, not noticeably

denticulate). Palp article 4 length 0.66 width; article 5

length 1.67 width, 1.0 article 4 length.

Pereopod \ (fig. 13) dactylus length 0.9 palm length.

Propodus length 0.97 width; dorsal margin with 12

setae (e.xcluding minute distal group). Propodal palm

with 7-8 broad based setae. Basis length 1.86 width,

dorsal ridge with 3 minute setae proximally. Pereopod

IV (figs 14C-D) length 0.31 body length, Propodus

length 3.0 width, with 8 broad based setae on ventral

margin. Carpus length 0.13 pereopod length, with 8

broad based setae on ventral margin, 2 distinctly larger

than others. Ischium posterodistal margin w'ith 7 setae.

Basis length 3.83 width.

Penes (lig. I4F) length 0.17 body width at pereonite 7.

Pleopods (figs 15, I6d-E) I endopod length 0.79 exo¬

pod length. Pleopod II endopod length 0.7 exopod

length. Pleopod III exopod length of distal article 0.28

exopod length; endopod length 2.21 width. 0.58 exopod

length. Pleopod IV exopod length of distal article ().29

exopod length; endopod length 1.97 width. Pleopod \'

endopod length 1.48 width. Pleopod protopods I with

10 simple and minutely serrate setae along length of lat¬

eral margin; protopod II with 18 simple and minutely

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

475

Figure 14. Phrealoicopsis teiricola. Hermaphrodite (AM P6I438) A-B, F, hermaphrodite (AM P6I439) C-E. A-B,

pereopod II. C-D, pereopod IV. E-F, pereopod VII with penes, including proximal articles, some setae missing

from basis dorsal margin. Scale bar I mm.

G.D.F. WILSON AND S.J. KEABLE

476

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

477

Figure 16. Phreotoicopsis leiricola. Hermaphrodite (AM P6I438). A-C, F, pleotelson and uropod, dorsal, lateral

and posterior views. D. pleopod II appendi.x masculina and endopod. E, pleopod I. G. uropod protopod distal

margin, ventral view. Scale bar I mm.

serrate setae along length of lateral margin. I sub-

marginally. PIcopiul I exopod medial margin straight -

divergent from lateral margin only proximally. Pleopod

II endopod appendix masculina with 16 setae on

margin.

Uropod (Txgs I6A-C. F-G) total length 0.64 pleotel¬

son length. Protopod length 2.57 width, 0.46-0.5 uro¬

pod total length; dorsomedial ridge length 1.15 endo¬

pod length. Endopod with 4-6 robust setae (including 3

small dorsolateral setae). Exopod length 0.62-0.78

endopod length; distoventral margin with 2 robust

setae.

Distribution. Wetlands of the Otway Range,

including Gellibrand Gorge and Cobden.

Victoria.

Remarks. See Remarks under P. raffae sp. nov.

for distinguishing the two known species of this

genus. Nicholls (1943) remarked upon the appar¬

ent hermaphroditism observed in this species. In

the material that we examined, the distribution of

male and female characteristics does not appear to

be correlated with body size.

478

G.D.F. WILSON AND S.J. KEABLE

Smaller specimens show some dilTerences

from adults that might cause difficulty in identifi¬

cation. In particular, the dorsomedial ridge of the

uropodal protopod of small specimens does not

project significantly beyond the distal margin of

the protopod, and may lack the ventral-most of

the three setae on the projection (figs I6F,G).

These smaller specimens, however, retain the

concave margin of the dorsomedial ridge. The

pleotelson ventral margin typically has two robust

setae anterior to the uropodal insertion, but

smaller specimens may only have a single large

seta. Two lots of specimens (NMV J44891,

J44887) retain these features in larger individuals

(up to 4cm); these have been listed separately as

variants of P. tenicola.

Synamphisopus Nicholls

Synamplmopiis Nicholls, 1943: 95.

Type species. Amphisopus amhigmis Sheard,

1936 by original designation.

Diagnosis. Typhlosole well developed, ventral

invagination forming double spiral in cross sec¬

tion. Pleotelson posterior margin cleft, refie.xed

dorsally; lateral lobes forming vertical plates; dor¬

sal uropodal ridge curv ing strongly and extending

posteriorly from uropods on pleotelson. Anten-

niila article 3 rudimentary second flagellum pre¬

sent. Mandible spine row on round pedunculate

projection. Ma.xillula lateral lobe distal margin

with 20-29 smooth robust setae, inner lobe nar¬

row and tapering. Pereopod I dactylus dorsal

margin with dense group of elongate setae. Pere-

opods propodus 11-IV without articular plate.

V-Vll with articular plate; pereopod IV se.\ually

dimorphic, prehensile in male. Pleopods II endo-

pod appendix masculina proximal half of shaft

broadly concave in ventral cross section, forming

tube at distal tip; with large subterminal denticles

around lateral to medial and dorsal surfaces.

Uropodal rami distal tips rounded.

Remarks. The synapomorphies of the clade

Phreatoicopsis + Synamphisopus are discussed

above in the Phreatoicopsis generic remarks. Fea¬

tures of Synamphi.sopus species that differ from

Phreatoicopsis include limited .sexual dimor¬

phism in the fourth pereopod, rounded tips of the

uropods, small dorsal plates of the basis on the

posterior three pereopods and shape of the pleo¬

telson. Our phylogenetic analysis identifies these

features as plesiomorphic character states within

the Amphisopodidae. The dense group of elon¬

gate setae on the medial side of the pereopod 1

dactylus, however, are unique to species of

Synamphisopus. These setae (fig. 21H) are bipin-

nate with two rows of tiny curv ed, equal length

spinules on the distal third of the setal shaft.

Although Synamphi.sopus amhiguus is a large

species, easily found and examined, it has not

been studied in detail since Sheard's (1936) orig¬

inal description and Nicholls’ (1943) treatment.

Our redescription of this species revealed a

second species of Synamphisopus from the

Grampians, which we describe below.

Synamphisopus doegi sp. nov.

Figures 17-25

Material examined. Hololype. Victoria, small tributary

of Glenclg R. crossing Sawmill Track, Grampians

National Park, 37°21.44'S 142°I7.79’E (GPS), sand

under rocks and submerged wood, hand sieves, pi 1 5.5

10.0°C, G. Wilson, R. Wetzer and S. Keable, 21 Sen

1999, VIC-96, NMV J40731 (male bl 23,3 mm, ethanol

preserved).

Paratypes. As for holotype. AM P61253 (8 males. 5

females. 1 indeterminate ind.), AM P61434 (male bl

21.4 mm. dissected for illustration. SEM and descrip

tion), AM P6I435 (female bl 17.5 mm. dissected for

SEM and description).

Other material. Vic., creek crossing Sawmill Track,

tributary of Gleneig R., Grampians, 37°20.22'S

142°19.62'E (GPS), sand under rocks and leaf litter in

stream, hand sieves. pH 6.45, 10.8°C, G. Wilson R

Wetzer and S. Keable. 21 Sep 1999, VlC-95. AM

P61254 (male, female, preserved in 95% ethanol); Vic..

Sawunill Track, Gleneig R.. Grampians, 37°21.75'S

142'’I6.90'E (map). T. Doeg, 17 Nov 1994, GrI, AM

P6I255^ (2 juvenile ind.); Vic., off Goat Road, tribu¬

tary ol Billimina Creek, Grampian.s, 37°14.22'S

I42“I9.07'E (map), T. Doeg, 15 Nov 1994, Gr49, AM

P6I257 (I juvenile ind.); Vic., 50 m below The

Fortress, Grampians, 37°I8.8() S I42°18.()0'E (esti¬

mate), J.E. Aslin, 26 Apr 1973, SAM C6027 (2

females).

Etymology. This species name honours Tim Doeg

(Flora and Fauna Branch, Department of Natural

Resources and Environment, Victoria, now at

Northcote, Victoria) who sent us fresh specimens

of Synamphisopus early in our project.

Diagnosis. Pleotelson medial dorsal ridge

smoothly arcing, low, in lateral view projeeting

ventrally to form setose lobe below level of

widely cleft medial lobe. Mandible palp article 2

with ventrolateral row of elongate setae, most

longer than distal article; spine row shaft anterior

margin with two separate dentate spines. Ma.xil-

lula medial lobe rounded and broader di.stally

than more proximally. Maxilliped palp article 5

suboval, short, length 1.6 width, lateral margin

rounded. Pereopod 1 of adult male dactylus

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

479

Figure 17. Synamphisopiis doegi, sp. nov. Holotype (NMV J4()731), lateral view. Scale bar 2 mm.

ventral margin convex, lateral face without pits

but with ventrolateral row of setal groups; pro-

podal palm heavily setose on medial margin,

lacking large projecting spine. Pereopods II-IV

basis proximal dorsal margin with row of simple

setae, shorter than setal row in male, longer than

setal row in female; male pereopod IV propodus

ventral margin linear; pereopod VII propodus dis-

tomedial margin with setae longer than articular

plate or dactylar claw, Pleopoci II appendix

masculina projecting beyond margin of proximal

exopod article, with only I flattened tubercle on

distomedial margin. Uropod protopod dis-

toventral margin with 2 robust smooth setae and 1

additional small seta.

Description based on male. Coloration in 95% ethanol

white legs, body slate gray-brown, head with black eye-

spot.

Eyes (figs 17, I8B, I) maximum diameter 0.18 head

depth.

Pleonites (fig. 17) 1-4 relative lengths unequal,

increasing in length from anterior to posterior, width

0.31 composite length in dorsal view.

Pleotelson (figs 17,25A-E) lateral length 0.90 depth;

dorsal length 1.65 width; depth 1.71 pereonite 7 depth.

Medial lobe width 0.58 pleotelson width, greatest

length 0.08 pleotelson total length.

Antenmda (figs I8C-E) length 0.07 body length,

with 9-10 artieles. Article 5 length 1,0 width. Article 6

length 1.13 width. Numerous tiny aesthetascs on article

5 to terminal article. Terminal article length 0.75 width.

Antenna (figs I8F-H) length 0.47 body length. Flagel¬

lum length 0.71 total antenna length, with 43-46

articles.

Mouthfwid clypeus width 0.52 head width. Mandible

(fig. 19) palp length 0.82 mandible length: article 3 with

14 setae; cuticular combs present; separate distal group

of setae absent; articles 1-2 with groups of long .setae

(longer than half article length) on ventral lateral

margins (along entire length of article 2). Left spine row

with 25-27 spines (appro.ximately but basally fused so

how many bifurcate unclear), additional spines between

pedunculate projection and molar absent, first spine not

separated from remaining spines. Right spine row with

19-25 spines (approximately hut basally fused so how

many bifurcate unclear), additional spines between

pedunculate projection and molar absent. Molar process

with complex sctulate spines forming posterior row.

Maxilliila (figs 20B-D) medial lobe length 0.8 lateral

lobe length, width 0.73 lateral lobe width, with 3 pap¬

pose setae; with 2 ‘accessory’ setae, one between disto-

lateral pappose setae and one between distomedial pap¬

pose setae, ‘accessory’ setae distally denticulate.

Lateral lobe distal margin with 20 smooth robust setae,

distal setal row with 5 robust setae; ventral face with 3

plumose setae. Masilla (figs 20E-G) medial lobe width

1.29 outer lateral lobe width. Outer lateral lobe width

subequal to inner lateral lobe. Maxilliped (figs 20H-I)

endite with 7 coupling hooks on right side; dorsal ridge

with 25-27 large distally denticulate plumose setae

(approximately). Palp article 4 subcircular, article 5

length 0.7 article 4 length.

Pereopod 1 (figs 21D-H) dactylus projecting beyond

palm, length 1.1 palm length; distoventral margin

480

G.D.F. WILSON AND SJ. K.EABLE

r/j

Figure 18. Synamphisopus cloegi, sp. nov. Paratype male (AM P61434), paratype female (AM P6I435). A-B, male

head, dorsal and lateral views. C, male anlennula. D, male antennula article 3 rudimentary second flagellum.

E, male antennula distal articles. F-H, male antenna. 1, female head lateral view. J-K, female antennula. Scale bar

0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

481

Figure 19. Synamphisopus doegi, sp. nov. Paratypc male (AM P6I434). A-E, right mandible. F-J, left mandible.

Scale bar 0.5 mm.

482

G.D.F. WILSON AND S.J. KEABLE

Figure 20. Synamphisoptis doegi, sp. nov. Paratype male (AM P61434). A, paragnaths. B-D, ma.xillula.

E-G, maxilla. H-l, maxilliped. Scale bar 0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

483

Figure 21. Symimpliisoptis doegi, sp. nov. Paratype female (AM P61435), paratype male (AM P6I434).

A-C, female pereopod I. D-H. male pereopod I. Scale bar I mm.

484

G.D.F. WILSON AND S.J. KEABLE

Figure 22. Synamphisopus Joegi, sp. nov. Paratype male (AM P61434), paratype female (AM P6I435). A, male

pereopod II. B-C, male pereopod IV. D, female pcrcopod IV. E-G, male pereopod VII, including pro.ximal

articles, with penes. Scale bar I mm.

spines positioned along 0.59 total length. Propodus dor¬

sal margin setae present in several groups between

proximal and distal margin. Propodal palm without

stout denticulate setae. Basis ventrodistal margin with I

elongate seta. Pereopod IV (figs 22B-C) propodus

ventral margin with 2 robust setae distinctly larger than

others.

Penes (fig. 22G) length 0.37 body width at pereonite

7, with setae on tip.

Pleopod I length O.I I body length; exopod length

2.25 width; endopod length 2.28 width, 0.82 exopod

length. Pleopod II exopod length 2.25 width; exopod

distal article length 0.37 exopod length; endopod length

I.9I width, 0.63 exopod length. Pleopod III exopod

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

485

Figure 23. Synamphisopus doegi, sp. nov. Paratype male (AM P6I434). A, pleopod I. B-E, pleopod II appendix

masculina and endopod. Scale bar 0.5 mm.

length 1.75 width; exopod di.stal article length 0.35

exopod length; endopod length 0.78 exopod length.

pleopod IV exopod length 1.6 width; endopod length

2.03 width, 0.85 exopod length. Pleopod V exopod

length 1.67 width; exopod distal article length 0.33 exo¬

pod length; endopod length 1.66 width. 0.61 exopod

length. Pleopod \ \ endopod appendix masculina with 48

setae on medial margin; length 0.59 pleopod length.

Uropod 25) total length 1.2 pleotelson length.

Protopod length 2.73 width, 0.42 uropod total length.

Endopod with 7-8 robust setae. Exopod length 0.69

endopod length; dorsal margin with 5 robust setae.

Sexual dimorphism and female characters. Antenna

length 0.4 body length; llagellum with 39-40 articles.

Pereopod I (figs 2IA-C) propodal palm with 9 stout

denticulate setae; robust simple setae basally inflated

(difficult to see behind lateral plumes of setae). Pereo¬

pod W (fig. 220) propodus ventral margin with 3 broad

based setae on ventral margin (with additional 4 sub-

marginally). Pleopod protopod II lateral margin with

8 simple setae proximally. Uropod endopod dorsal

margin with 5 robust setae.

Distribution. Glenelg R. drainage, Grampians

National Park.

Remarks. Synamphisopus doegi sp. nov. differs

from S. amhiguus in several features, making the

two species easy to distinguish. The pleotelson

medial ridge of 5. doegi extends ventrally and is

produced in a setose lobe, rather than tenninating

dorsally. The uropod protopod distoventral mar¬

gin robust setae arc smooth rather than spinose.

The appendix masculina medial margin setae tend

to form two rows basally, although a remnant (or

precursor) of these setae can be seen in the

S. ainbiguus. The pereopod I propodal palm of

S. doegi males lacks a spine. This new species

also lacks punctae on the dactylus of pereopod I

that are distinctive in S. ainbiguus. The two

species differ in the degree of setation on the

body, S. doegi generally being more setose than

S. ambiguus in homologous positions.

486 G.D.F. WILSON AND S.J. KEABLE

Figure 24. Synamphisopus doegi, sp. nov. Paratype male (AM P61434). A-E, pleopods I-V. Scale bar I mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

487

Figure 25. Synamphisopus doegi, sp. nov. Paratype male (AM P6I434). A-E, plcotelson and uropod, ventral,

dorsal, lateral and posterior views. F, uropod protopod distal margin, ventral view. G, uropod, medial view. Scale

bar 0.5 mm.

Synamphisopus ambipuiis (Sheard)

Figures 26-33

Amphisopus amhiguus Sheard, 1936; p.469, figs

1-18.

Synamphisopus amhiguus. Nicholls, 1943: 96, fig.

24.

Type maierial. flolotypc. Victoria, Fish Falls, Macken¬

zie R., Grampians, 37°07'S, I42°26'E (estimate), SAM

C 2115 (carcass), C 21 16 (slides lost).

Material examined. Scries from Vic., Grampians, col¬

lected G. Wilson, R. Wetzcr and S. Keable, Sep 1999,

preserved in 95% ethanol - top of Mackenzie Falls,

Mackenzie R., 37'’06.70'S I42°24.58'E (GPS), seep at

side of path, hand sieve, pH 6.9,8.9°C, 20 Sep. VIC-84,

AM P61258 (I ind.); Vie., base of Mackenzie Falls,

Mackenzie R.. 37°06.74'S I42°24.52'E (GPS), gravel,

sand, mud under rocks in seeps, hand sieves, 20 Sep,

VIC-85, AM P6I259 (8 ind.); Vic., base of Fish Falls.

Mackenzie R.. 37°06.59'S l42“24.0rE (GPS), sand

under roots of ferns in seeps to side of main falls near

seep sources, hand sieves, hand and spoon, pH 6.6.

10.6°C, 20 Sep, VIC-86, AM P6I260 (5 males, 3

females), AM P61440 (male bl 24.1 mm, dissected for

illustration, SEM and description, collection details as

488

G.D.F. WILSON AND S.J. KEABLE

Figure 26. Synamphisopus amhiguiis. Female (AM P6144I), male (AM P61440). A-B, female head, lateral and

ventral views. C-F, male antenna. G-H, male antennule. Scale bar 1 mm.

for P61260), AM P61441 (female bl 26.6 mm, dissected

for SEM and description, collection details as for

P61260), AM P61442 (male bl 28.2 mm, dissected for

SEM and description, collection details as for P6I260),

AM P61443 (female bl 25.5 mm, dissected for SEM

and description, collection details as for P61260); Vic..

Stony Creek below Turret Falls on Twin Falls Trail.

37°09.4rS 142°29.90'E (GPS), from gravel under

rocks, hand sieves, pH 6.7, 7.2°C, 21 Sep, VIC-89, AM

P61261 (16 ind.); Vic., Stony Creek below Turret Falls

on Twin Falls Trail, 37°07.93'S 142°30.26'E (GPS),

fern roots at stream side, hand sieves, 21 Sep, VIC-90,

AM P6I262 (I ind.); Vic., Stony Creek directly below

Turret Falls on Twin Falls Trail, 37°09.66'S

142°29.83'E (GPS), from water weed, hand sieves. 21

Sep, VIC-92, AM P6I268 (I ind. and 1 carcass); Vic.,

base of Beehive Falls near Roses Gap, 36°58.54'S

I42°27.01 'E (GPS), under rocks in pool and leaf litter

in stream, hand sieves, pH 4.6, I0.3°C, 22 Sep, VIC-97,

AM P6I269 (2 ind.). Vic., off Silverbrand Road. Stony

Creek, Grampians, 37“09.20'S 142“29.67'E (map), T.

Doeg and J. Read, 15 Nov 1994, site GRI9, AM

P54I03 (1 male bl 26.2 mm, dissected, 1 preparatory

female bl 24.9 mm, dissected); Vic., Stony Creek, near

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

489

Figure 27. Synamphisopus amhiguus. Male (AM P61440). A-C, left mandible. D-H, right mandible. Scale bar

0.5 mm.

490

G.D.F. WILSON AND S.J. KEABLE

' tw

■lr¥

Figure 28. Syiuimphisopus anihigiitis. Male (AM P61440). A, paragnaths. B-C, maxillula. D-E, maxilla.

F-H, maxilliped. I, left mandible molar. J, right mandible molar. Scale bar 0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

491

Figure 29. Symimphi.iopiix amhiguus. Male (AM P6I440), A-F, pereopod I. Scale bar I mm.

Halls Gap, Grampians. 37°09.72’S 142°29.74'E (map),

altitude 510 metres, kick sample, D. Crowthcr, 10 Dec

1998.98-220 Site 87, AM P61294 (I iiid.), AM P61295

(2 ind., collection details as for AM P6I294 but not

kick sample); Vic., 2 km from Zumsteins along road to

Halls Gap, Grampians, 37“05’S I42°25'E (esiimate).

under rocks in fine sand sprayed with water from small

trickle over cliff (on uphill side of road) onto rock ledge

below, J.H. Bradbury. 29 Mar 1995, JHB VIC#24, AM

P53150 (II ind.); Vic., The Grampians (37°I7'S,

I42°33'E). A. NcboLss, 1 Oct 1954, NMV J44897 (3,

labelled as Phrcatoicopsis leiricola): Vic., Briggs

Bluff. Mt Rosea (36“59'S, I42°28'E). I.R. McCann.

March 1957, NMV J44889 (I, labelled as Phrea-

toicopxis lerricola).

Diagnosis. Pleotelson medial dorsal ridge with

posterior obtuse angle, in lateral view projecting

only to posterior margin above level of widely

cleft medial lobe. Mandible palp article 2 with

ventrolateral setae shorter than distal article; spine

row shaft anterior margin without separate

dentate spines. Maxilliila medial lobe tapering

distally. Maxilliped palp article 5 elongate, length

2.4 width, lateral margin linear. Pereopod 1 of

adult male dactylus ventral margin sinuous, lat¬

eral face with regularly spaced setal pits; propodal

palm with setal rows on proximal medial and

lateral surfaces, with large spine midlength. Pere-

opods II IV basis proximal dorsal margin with

single robust seta; pereopod IV propodus ventral

margin convex; pereopod VI1 propodus disto-

medial margin with setae shorter than articular

plate or dactylar claw. Pleopod II appendix

492

G.D.F. WILSON AND S.J. KEABLE

Figure 30. Synamphisopus amhigiius. Female (AM P6144I). A-D, pereopod I. Scale bar I mm.

masculina not projecting beyond margin of

proximal exopod article, with 2 flattened tuber¬

cles on distomcdial margin. Uropod protopod

distoventral margin with 3 robust spinose setae.

Description hosed on male. Coloration in life white

(colourless) legs, body slate gray-brown, head with tiny

white eyespot; similar in 95% ethanol but eyespot

black.

Eyes (fig. 26A) maximum diameter 0.12 head depth.

Pleonites 1-4 relative lengths subequal, width 0.55

composite length in dorsal view.

Pleotelson (figs 33A-E, G) lateral length 0.94 depth:

dorsal length 1.46 width; depth 1.64 pereonite 7 depth.

Medial lobe width 0.54 pleotelson width, greatest

length 0.02 pleotelson total length.

Antennula (figs 26G-H) length 0.09 body length,

with 11 articles. Article 5 length 1.29 width. Article 6

length 0.75 width. Numerous tiny aesthetascs on article

8 to terminal article. Terminal article length subequal to

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

493

Figure 31. Synamphisopus amhiguus. Male (AM P61440), female (AM P61441). A-B, male pereopod II.

C-D, male pereopod IV. E, female pereopod IV. F-FI, male pereopod VII, including proximal articles, with penes.

I-J, male pleopod 11 appendix masculina and endopod. Scale bar 1 mm.

494 G.D.F. WILSON AND S.J. KEABLE

penullimale article length, 1.36 width. Antenna (figs

26C-F) length 0.51 body length. Flagellum length 0.67

total antenna length, with 54 articles (40 articles in

illustration: Sheard, 1936).

Mnulhfield c\ype\is width 0.46 head width. Mandible

(figs 27, 28I-J) palp length 0.88 mandible length,

article 3 with 23 setae, cuticular combs absent; separate

distal group of setae present; articles 1-2 with groups of

long setae (longer than half article length) on ventral

lateral margins. Left spine row with 36 spines (approx¬

imately but basally fused so how many bifurcate

unclear), first spine separated from remaining spines.

Right spine row with 33 spines (approximately, basally

fused so how many bifurcate unclear). Molar process

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK.

495

Figure 33. Synamphisoptis amhiguus. Male (AM P61440). A-C, G, pleotelson, dorsal, lateral and ventral views.

D-E, pleotelson and uropod, lateral view. F, uropod protopod distgl margin, ventral view. H, pleopod I. Scale bar

496

G.D.F. WILSON AND S.J. KF.ABLE

spines absent. Maxiltula (figs 28B-C) medial lobe

length 0.76 lateral lobe length, width 0.5 lateral lobe

width, with 4 pappose setae, with I ‘accessory’ seta

between central pappose setae, ‘accessory’ setae sim¬

ple. Lateral lobe distal margin with 29 smooth robust

setae, distal setal row with 4 robust setae; ventral face

with 4 plumose setae. Maxilla (figs 28D-E) medial lobe

width 1.86 outer lateral lobe width. Outer lateral lobe

wider than inner lateral lobe. A/av/V/iper/(figs 28F-II)

endite with 5 coupling hooks on right side; dorsal ridge

with at least 20 large distally-denticulate plumo.se setae.

Palp article 4 elongale-distally e.xpanded, article 5

length 1.09 article 4 length.

Pereopod I (figs 29A-F) dactylus only slightly

shorter than palm, length 0.94 palm length. Propodus

dorsal margin setae confined to single group at distal

margin. Propodal palm with stout denticulate serrate

setae. Basis ventrodistal margin lacking elongate setae.

Pereopod IV (figs 3IC-D) propodus ventral margin

with 3 robust setae distinctly larger than others, central

seta largest.

Penes (fig. 31II) length 0.5 body width at pereonite

7; setae absent.

Pleopods (figs 3II-J, 32) I length 0.15 body length;

exopod length 2.88 width; endopod length 2.0 width.

0.61 exopod length. Pleopod II exopod length 2.79

width; exopod distal article length 0.3 exopod length;

endopod length 2.98 width. 0.69 exopod length. Pleo¬

pod III exopod length 2.12 width; exopod distal article

length 0.33 exopod length; endopod length 0.72 exopod

length. Pleopod IV exopod length 2.4 width; endopod

length 2.34 width. 0.75 exopod length. Pleopod \ exo¬

pod length 1.83 width; exopod length of distal article

0.38 exopod length; endopod length 1.78 width, 0.67

exopod length. Pleopod II endopod appendix masculina

with 23 setae on medial margin; length 0.38 pleopod

length.

Uropod (figs 33D-E) total length 1.06 pleotelson

length. Protopod length 3.46 width, 0.38 uropod total

length. Endopod with 12 robust setae (including 5 at

distal tip). Exopod length 0.76 endopod length; dorsal

margin with 6 robu.st setae (including 3 at distal tip).

Sexual dimorphism and female characters. Antenna

length 0.45 body length; flagellum with 42 articles.

Pereopod I propodal palm with 20 stout robust conical

setae. Pereopod IV propodus ventral margin with 5-6

broad based setae on ventral margin (and additional 3-5

submarginally); robust simple .setae absent. Pleopod

protopod II lateral margin with 4 simple setae proxi-

mally. Uropod endopod dorsal margin with 13 robust

setae

Distribution. Stony Creek and Mackenzie R.

drainages, and Beehive Falls, Grampians

National Park, Victoria.

Remarks. Synamphisopus amhiguus populations

vary in at least one feature. Specimens from Fish

Falls on Mackenzie R. (the type locality) all have

3^ robust setae on each side of the cleft pleotel¬

son tip, while specimens from other locations

sometimes have more. Specimens from Stony

Creek and from Mackenzie Falls (Mackenzie R.

above Fish Falls) have 5-6 .setae in this position,

while most other specimens, including that front

Beehive Falls to the north of the type locality,

have 3-4 setae. Sheard (1936) states that the

coxae of all legs are fused, that the fourth perco-

pod is not sexually differentiated and that

appendix masculina lacks setae; our material does

not support these observations.

Phrcatoicidae Chilton

Phrcatoicidac Chilton. 1891: 151.

Diagnosis. Head tubercles absent; antennal notch

shallow, without posterior extension. Pleonite 1

pleura distinctly shallower than pleurae of

pleonites 2-5. Pleotelson posterior margin entire

reflexed dorsally, produced. Antennal article 3 '

rudimentary second (lagcllum absent. Mandible

right lacinia mobilis reduced, incorporated into

spine row, with tooth or denticulated .scale on

anterior face; spine rows linear on pedunculate

projection. Pereopod I mcrus dorsal projection

shelf-like and U-shaped. Pereopods II III propo¬

dus with articular plate on posterior side of limb;

pereopod IV sexually dimorphic, prehensile iri

adult males. Pleopod exopods II-V with lateral

proximal lobes; protopods medial margin without

coupling hooks; protopods l-II lateral epipod

absent. III V lateral epipods lobe-like. Uropod

rami distal tips pointed.

Remarks. The I’hreatoicidae includes a diverse

group of taxa, united by reduction of the right

lacinia mobilis. which is fused into the distal

member of the spine row. Additionally, the spine

rows on both mandibles arc pedunculate. The

phreatoicid spine rows are notably different from

fused spines found in the Synamphisopus +

Phreatoicopsis clade (discussed above) because

the individual bifurcate spines in the Phreatoi-

cidae are distinct and aligned parallel in a

dorsoventral plane. Other features that define the

Phrcatoicidae are homoplastic, being shared with

members of the Amphisopodidae and Hypsime-

topodidae. Material from the Grampians contains

four species of Phrcatoicidae that could not be

assigned to any existing genera. The phylogenetic

analysis shows that these species are mono-

phyletic, but we cannot diagnose this clade with

unambiguous apomorphic features, although

three of the species have several synapomorphies.

Accordingly, we introduce the genera Naio-

pegia gen. nov. and Gariwerdeus gen. nov. to

accommodate these species.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

497

Naiope^ia gen. nov.

Type Species. Naiopegia xiphagrastis sp. nov.

Ety mology. The genus name is compounded from

the Greek words "Naio", meaning to dwell or

inhabit, and "Pege" (f.) for spring.

Diagnosis. Cephalon without eyes, cuticle rugose

with cuticular hairs, with elongate setae (longer

than basal 2 articles of antennula), especially at

ventrolateral margins; clypeal notch present (but

weak). Typhlosole minimal, ventral invagination

forming laminar projection in cross section. Pleo-

telson dorsal surface rugose with cuticular hairs,

without abundant long setae; medial and lateral

lobes distinct; lateral lobes rounded; medial lobe

refle-xed, projecting beyond lateral lobes, with 4

robust setae. Antennula with 6 (rarely 7) articles,

penultimate article elongate and inflated, terminal

article shorter than broad, not compound, much

shorter than article 5. Antenna basal articles of

flagellum with dense cuticular hairs. Mandible

palp articles I and 2 with row of elongate setae

(longer than distal article); right lacinia mobilis

anterior scale with 4 robust denticles. Maxilliped

basis distal margin adjacent to palp insertion with

elongate setae (longer than palp articles 1-3).

Pereopod I not strongly dimorphic, propodus of

male only somewhat broader than female and

dorsal margin not produced pro.ximally; propodal

palm concave, spines absent, with stout conical

robust setae in male, denticulate robust setae in

female, cuticular fringe weakly developed, setal

ridge absent. Pereopods l-Vll proximal portions

(basis, merus, ischium) with numerous elongate

setae (many longer than ischium) on dorsal and

ventral margins; basis dorsal ridge in cross

section rounded on pereopod V, angular on

pereopods VI-VII. Pleopod If endopod appendix

masculina proximal half of shaft solid and rod¬

like, indented in ventral cross section, lacking

setae on lateral or medial margins. Uropod proto¬

pod distoventral margin with I robust spinosc seta

and several simple setae.

Remarks. We introduce a monotypic genus

because Naiopegia xiphagrastis gen. nov., sp.

nov. shares no unambiguous synapomorphies

with other genera of the Phreatoicidae. This

species has a typical phreatoicid body and limb

form, but no species of this family have cuticular

hairs on the basal articles of the antennae. Its

"reduced" antcnnule. another diagnostic and

apomorphic character, is broadly homoplastic,

being found in taxa outside of the Phreatoicidae,

such as Crenisopus and some species of Mesarn-

phisopus. Species of Gariwerdeus gen. nov. also

have similar six-articled antennulae (see below),

but with a differing composition of the distal two

articles. Naiopegia lacks the distinctively fomted

pleotelson of Gariwerdeus and is similar in this

regard to other phreatoicids. Of the blind, but

more typical phreatoicids, the cross sectional

shape of the appendix masculina as well as the

cuticular hairs on the antennula would separate

Naiopegia gen. nov. from Crenoicus.

Naiopegia xiphagrastis sp. nov.

Figures 34-41

Material examined. Holotype. Victoria, tiny spring-led

tributary of Stony Creek below Turret Falls, at bridge

on Twin Falls Trail. Grampians National Park,

37°07.93'S 142°30.26'E (GPS), sand with mud and

detritus in minimal seep flow among thicket of sword

grass, hand sieve, S. Keable, 21 Sep 1999. VIC-91,

NMV J40732 (male bl 12.0 mm, ethanol preserved).

Paratypes. As for holotype. AM P6127() (4 males),

AM P6I424 (I male bl 11.7 mm, dissected for illus¬

tration, SEM and description ). AM P6I425 (I female

bl 7.4 mm. dissected for SEM and description).

AM Pbl426 (male bl 11.1 mm, used to supplement

description and SEM).

Etymology’. The species name is a noun in apposi¬

tion from the Greek compound name for sword

grass, "xiphagrostis", and refers to the vegetation

found at the type locality.

Description based on male. Coloration in life with

white patches on head in eye region, gray dorsum

darker at edges; in 95% ethanol, uniform gray-white.

/toif/(figs 35B-C) length subcqual to width in dor¬

sal view; width 0.81 pereonite 1 width; surface with

dense cuticular hairs; setae sparse but forming dense

row along ventral margin.

Pereon broad, width 1.2 head width; with scattered

roughness (cuticular hairs).

Pleoniles in dorsal view 2-4 respective lengths more

than half length of pleonite 5; pleonites 1-4 relative

lengths suhequal, width 1.0 composite length in dorsal

view.

Pleotelson (figs 4IA-D) lateral length 0.12 body

length, 0.77 depth; dorsal length 1.22 width; depth 1.5

pereonite 7 depth. Dorsal surface without abundant

long setae. Medial lobe width 0.30 pleotelson width,

greatest length 0.22 pleotelson total length. Lateral

lobes in lateral view curs ing dorsally, in dorsal view

rounded, not extending posteriorly to level of medial

lobe, medial length 0.08 pleotelson total length, with 1

robust setae. Ventral margin anterior to uropods with 8

smooth .setae, posterior seta subequal to anterior adja¬

cent setae.

Antennula (figs 35B. D) length 0.07 body length,

with 6 articles. 2-3 tiny acsthetascs on articles 5-6.

Articles 5 and 6 width subequal to article 4. Antenna

(figs 35C, E, F-G) length 0.27 body length. Flagellum

length 0.66 total antenna length, with 20 articles.

498

G.D.F. WILSON AND S.J. KEABLE

Figure 34. Naiopegiaxiphagroxtis gen. nov., sp. nov. Holotype male (NMV J40732), lateral view. Scale bar 1 mm,

Moiithfield clypeus width 0.56 head width. Mandible

(fig. 36) palp length 0.74 mandible length; article 3 with

7 setae; separate distal group of setae absent; articles

1-2 with elongate setal row. Left incisor process with 2

distal cusps and 1 on dorsal margin (shoulder for fourth

cusp apparent but not forming full cusp). Left spine row

with 7 bifurcate spines, without additional spines

between pedunculate projection and molar. Right spine

row with 7 bifurcate spines in addition to lacinia

mobilis, without additional spines between pedunculate

projection and molar. Molar process longer than wide;

3^ setulate spines forming posterior row. Maxillida

(figs 37B-C) medial lobe length 0.83 lateral lobe

length; width 0.65 lateral lobe width. Lateral lobe distal

margin with 3 denticulate robust setae. 7 smooth robust

setae. Maxilla (figs 37D-F) medial lobe width 0.71

outer lateral lobe width. Maxilliped{(igs 37G- H) enditc

with 3 reeeptaculi on right side; dor.sal ridge with 10-13

large distally denticulate plumose setae (merging with

subdistal biserrate setae). Palp insertion on basis ventral

surface with 10 subdistal smooth setae (some forming

row).

Pereopod I (figs 38A-B) dactylus length 0.9 palm

length; vcntrodistal margin with row of sharp spines

along 0.37 total length. Propodus dorsal margin vt ith 21

setae in several groups between pro.ximal and distal

margin (excluding distal group); proximal region not

protruding. Propodal palm concave; stout denticulate

setae absent; with 9 stout robust simple conical setae;

elongate broad based setae absent. Basis vcntrodistal

margin with 10 elongate setae. Pereopod IV (figs

39C-D) dactylus shorter than propodal palm. Propodus

with 7 broad based setae on ventral margin, 2 distinctly

larger than others; articular plate subcqual in length to

dactylar claw. Pereopods V-VII (figs 39F-G) basis

dorsal ridge in cross section rounded on pereopod V.

angular on pereopods VI-VII.

Penes (fig. 391) length 0.25 body width at pereonite

7. distally tapering.

Pleopod I (figs 40, 3911) exopod distal margin

rounded, medial margin concave from proximal to

distal half, subparallcl to lateral margin, dorsal surface

lacking setae. Pleopod II endopod appendix maseulina

length 0.61 pleopod length, distal tip truncate; with 5

setae on margin.

Uropod (figs 4IA-E) total length 1,08 pleotel.son

length. Protopod length 0.42 uropod total length; ven¬

tral margin with anteriorly-projecting somewhat rigid

long setae, forming dense longitudinal row. setae medi¬

ally robust and distally spinose; di.stoventral margin

with I robust spinose seta and 2 simple setae. Exopod

length 0.87 endopod length.

Hexiial dimorphism, female differences. Aniennula

length 0.06 body length, with 6-7 articles, article 5

length greater than article 6. Antenna length 0.34 body

length, flagellum 0.61 total antenna length, with 16 arti¬

cles (incomplete, distal most articles missing). Peivo-

pod I (figs 38C-E) dactylus length 0.95 palm length:

vcntrodistal margin with row of 8 sharp spines, along

0.33 total length; propodal palm with 7 stout robust

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

499

Figure 35. Naiopegia xiphaffrostis gen. nov., sp. nov. Paratype male (AM P61424). A-B. head, anlennula, antenna

proximal articles, dorsal view. C, head, ventral view. D, antennula distal articles. E. antenna proximal articles,

antennula distal articles. F-G. antenna Ilagellum distal and proximal articles. Scale bar 0.5 mm.

simple conical setae; basis ventrodistal margin with 8

elongate setae. Perenpod IV (fig. 39E) propodus articu¬

lar plate shorter than dactylar claw. Uropod length 0.9

pleotelson length. 0.47 uropod total length, exopod

length 0.74 endopod length.

Distribution. Known only from small spring-fed

tributary of Stony Creek below Turret Falls, at

bridge on Twin Falls Trail, Grampians National

Park.

Remarks. This species is similar to most other

phreatoicids, save for a few distinctive features

and the lack of eyes. The elongate setae on all

limbs and the inflated, distally-broadened anten-

nular article 5 with a tiny, but distinct article 6 arc

diagnostic for this species. The antennular details

and the picsiomorphic retention of a large

reflexed medial lobe on the pleotelson also distin¬

guish it from species of Gariwerdeus gen. nov.

500

G.D.F. WILSON AND S.J. KEABLE

Figure 36. Naiopegia xiphagrostis gen. nov., sp. nov. Paratype male (AM P61426). A-C, left mandible. D-H, right

mandible. Scale bar 0.1 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

501

Figure 37. Naiopegia xiphagrostis gen. nov., .sp. nov. Paralype male (AM P6I424) A, G-H, paratype male (AM

P6I426) B-F. A, paragnaths. B-C, maxillula. D-F, maxilla. G-H, maxilliped. Scale bar 0.2 mm.

502

G.D.F. WILSON AND S.J. KEABLE

Figure 38. Naiopegki xiphagrostis gen. nov., sp. nov. f’aralype male (AM P61424), paratype female (AM P61425).

A-B, male pereopod I. C-E, female pereopod 1. Scale bar 0.5 mm.

The number of distal antennulac articles may be

variable among adults of N. xiphagrostis sp. nov.

- one female has 6 on one side and 7 on the other.

Gariwerdeus gen. nov.

Type Species. Gariwerdeus lurrelensis sp. nov.

Etymology. The Grampians (an English name

deriving from a mountain range in Scotland)

National Park is given the name “Gariwerd" by

the indigenous Jardvvadjali and Djab Wuming

people. The species of this genus appear to be

ubiquitious in streams, springs or groundwater

seeps within the Park, and thus should bear the

original name for this region. Gariwerdeus is

treated as a masculine noun.

Diagnosis. Cephalon without eyes; clypeal notch

absent. Typhlosole minimal, ventral invagination

simple (either laminar or Ti" shaped in eross sec¬

tion). Pleotelson dorsal surface with abundant

long setae and rugose with den.se cuticular hairs;

medial and lateral lobes distinct; posterior margin

rellexed dorsally, medial lobe compressed anteri¬

orly, shorter than lateral lobes; lateral lobes

acutely angular in dorsal or ventral view, in lat¬

eral view depressed ventrally, protruding poster¬

iorly from insertion of uropods. Antennula with 6

or fewer articles, distal articles inflated. Amcnna

flagellum proximal articles without dense cover

of cuticular hairs. Mandible palp article 2 with

elongate setal row (setae longer than distal

article). Pereopod 1 propodal palm concave.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

503

Figure 39. Naiopegiaxiphagroslis gen. nov., sp. nov. Paratype male (AM P6I424), paratype female (AM P61425).

A-B, male percopod II. C-D, male pereopod IV. E, female pereopod IV. F-G, male pereopod VII. II, male

pleopod II appendix masculina. I, male pereopod VII proximal articles, with penes. Scale bar 0.5 mm.

504

G.D.F. WILSON AND S.J. KEABLE

Figure 40. Naiopegia xiphagrostis gen. nov., sp. nov. Paratype male (AM P6I424). A-E, pleopods I-V. Scale bar

0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

505

Figure 41. Naiopegia xiphagrostis gen. nov., sp. nov. Paratype male (AM P6I424). A-D, pleotelson and uropod.

lateral, medial and ventral views. E, uropod protopod distoventral margin robust setae. F, pleotelson distal margin,

dorsal view. Scale bar 0.5 mm.

506

G.D.F. WILSON AND S.J. KEABLE

spines absent, with stout conical robust, elongate

or denticulate setae, setal ridge absent. Pereopods

1-Vll proximal portions (basis, merus, ischium)

with numerous elongate setae (many longer than

ischium) on dorsal and ventral margins; pereopod

Vll basis dorsal ridge in cross section angular.

PleopodW endopod appendix masculina proximal

half of shaft solid and rod-like, indented in ventral

cross section, lacking setae on lateral or medial

margins. Uropad protopod distoventral margin

with 1-2 robust spinose seta and several simple

setae.

Remarks. In addition to having a setose, light

coloured and rugose cuticle, species of Gariw-

erdetis gen. nov. can be recognised most readily

by their greatly reduced pleotelson tip, giving an

almost tnmcate appearance to the end of the body.

This feature can be seen when collecting speci¬

mens in the field. All species ot' Gariwerdeus are

also blind. The antennula is pauciarticulate, simi¬

lar to that of Naiopegia gen. nov., but the distal

articles dilTer in all of the Gariwerdeus species.

Our cursory investigation of the Grampians has

detected three species in this genus.

Gariwerdeus turretensis sp. nov.

Figures 42-49

Material e.xamiiwd. llolotype. Victoria, Stony Creek

directly above Turret Falls on Twin Falls Trail,

Grampians National Park, 37°09.62'S I42°29.82'E

(GPS), sand, silt and fern roots at edge of creek, hand

sieve, S. Keable, 21 Sep 1999, VlC-93. NMV J40733

(male bl 7.4 mm).

Paratypcs. As for holotype. AM P6I27I (20 ind.),

AM P61427 (male bl 8.6 mm, dissected for illustration,

SEM and description), AM P6I428 (female bl 8.4 mm.

dissected for SEM and description), AM P6I429 (male

bl 8.3 mm, used to supplement description and SEM).

Other material. Series from Vic.. Stony Creek,

Grampians, hand sieves, collected G. Wilson, R. Wet-

zer and S. Keable, 21 Sep 1999, preserved in 95%

ethanol - below Turret Falls on Twin Falls Trail

37“09.4rS 142‘’29.90'E (GPS), from vegetation at

edge of stream in sand, pH 6.7 (out of calibration)

7.2“C. VIC-89. AM P61273 (12 ind.); below Turret

Falls on Twin Falls Trail, 37°07.93'S I42°30.26'E

(GPS), fern roots at stream side, VIC-90. AM P6I274

(male, female); directly below Turret Falls on Twin

Falls Trail, 37°09.66 S 142‘’29.83'E (GPS), from

sphagnum at waters edge, hand sieves. VlC-92, AM

P6I272 (12 ind.) and NMV J40734 (male, 2 females).

Vic., Stony Creek, Grampians, 37°09.72'S

I42°29.74’F. (map), stream rinics, altitude 510 metres,

kick sampling, D. Crowther, 10 Dec 1998, 98-220 Site

87. AM P61257 (male).

Etymology. The name refers to Turret Falls on

Stony Creek in The Grampians National Park,

around which this species was collected.

Diagnosis. Pleotelson tip medial lobe lacking

robust sensillate setae hut with 8 fine simple

setae. Antennula article 6 inllatcd and bulbous,

temiinal two segments with one aesthetasc each,

article 5 length subequal to article 6 length.

Mandible right lacinia mobilis with 1 row of den¬

ticles. Pereopod I of male dactylus postcrodistal

margin smooth; propodus dorsal margin proxi-

mally produced, greatly expanded beyond dorsal

margin of carpus; propodal palm cuticular fringe

weakly developed. Pereopods II-IV basis dorsal

margin elongate setae in row, not clustered. Plea-

pod II endopod appendix masculina distal tip

tnmcate, with 3 setae on margin. Uropoda! proto¬

pod distoventral margin with 1 robust spinose seta

and 7 simple setae.

Figure 42. Gariwerdeus turretensis gen. nov., sp. nov. Holotype male (NMV J 40733), lateral view. Scale bar 1 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

507

Figure 43. Gariwerdens turretensis gen. nov., sp. nov. Paratype male (AM P61427) A-D, H, paratype female

(AM P6I428) E-G. A-B, H, male head, antenna, antennule, lateral, dorsal and ventral views. C-D, male antenna

flagellum, distal and proximal articles. E, female head, antenna, antennula, dorsal view. F, female antennula, distal

articles. G, ciliate protozoan epibiont (Fwhre//« sp.) on female antenna. Scale bar 0.5 mm.

Descriplion based on male. Coloralion in life trans¬

lucent with dark patches on posterior pleonites,

white dorsum anteriorly, head white in eye region; in

95% ethanol eye region lacking colour, gray-white¬

head to pereonite 6, pereonite 7 to pleotelson light

brown.

Head (fig. 43B) length subequal to width in dorsal

view; width 0.85 pereonite I width; surface smooth and

shiny (with scattered cuticular combs), dorsal .setae

sparse.

Pereon narrow, width near head width (1.18

head width); smooth. Typhlosole minimal, ventral

508

G.D.F. WILSON AND S.J. KEABLE

Figure 44. Garinerdetis turretensis gen. nov., sp. nov. Paratype male (AM P61429), paratype female (AM P61428).

A-B, male left mandible. C, male paragnaths. D, male right mandible. E-F, female right mandible palp. Scale bar

0.05 mm.

invagination forming laminar projection in cross

section.

Pleonites in dorsal view 2-3 respective lengths less

than half length of pleonitc 5. pleoniie 4 length more

than half length of pleonite 5; pleonites 1-4 relative

lengths unequal, increasing in length from anterior to

posterior, width 0.88 composite length in dorsal view.

Pleotelson (figs 49A-D, G) lateral length 0.13 body

length. 0.93 depth; dorsal length 1.3 width; depth 1.47

pereonite 7 depth. Dorsal surface with abundant long

setae. Medial lobe width 0.27 pleotelson width, greatest

length 0.04 pleotelson total length. Lateral lobes medial

length 0.04 pleotelson total length, without robust sen-

sillate setae. Ventral margin anterior to uropods with

4-6 denticulate and smooth setae (posterior seta

smooth, anterior setae weakly denticulate), posterior

seta smaller than anterior adjacent setae (thicker but

shorter).

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

509

Figure 45. Gariwerdeu.s lurretensis gen. nov., sp. nov. Paratype male (AM P6I429). A, maxillula. B-C, maxilla.

D-E, maxilliped. Scale bar 0.2 mm.

510

G.D.F. WILSON AND S.J. KEABLE

Figure 46. Gariwenleiis luiretensis gen. nov., sp. nov. Paratype male (AM P61427) A, paratype I'cmalc (AM

P61428). paratype male (AM P6I429) B. A, B, male percopod I. C-E, female pereopod I. Scale bar 0.5 mm.

Anienntila (llgs 43E-F) length 0.11 body length, with

6 articles. Single tiny aesthctasc on article 5 to terminal

article. Antenna (figs 43B-D) length 0.34 body length.

Flagellum length 0.57 total antenna length, with 16

articles.

Mouthfield clypeus broader medially, width 0.65

head width. Mandible 44) palp length 0.8 mandible

length; article 3 with 11 setae; separate distal group of

setae present; articles 1-2 w ith elongate sctal row. Left

spine row with 7-8 bifurcate spines, without additional

spines between pedunculate projection and molar.

Right spine row with 8 bifurcate spines, without addi¬

tional spines between pedtmculate projection and

molar. Molar process setulate spines forming posterior

row (2 tiny on right). Mavillida (fig. 45A) medial lobe

length 0.86 lateral lobe length; width 0.67 lateral lobe

width. Lateral lobe distal margin with 5 denticulate

robust setae, 7 smooth robtist setae. Maxilla (figs

45B-C) medial lobe width 0.56 outer lateral lobe width.

Maxilliped (figs 45D-E) endite with 2 receptaculi on

right side; dorsal ridge with 12 large distally denticulate

plumose setae. Palp insertion on basis ventral surface

without subdistal smooth setae.

Pereopod I (figs 46A-B) dactylus length 1.07 palm

length. Propodus dorsal margin .setae confined to single

group at distal margin. Propodal palm convex to

straight; stout denticulate setae absent; with 4 basally

inllated stotit robust simple setae; with 3 elongate broad

based setae. Basis ventrodistal margin with 7 elongate

setae. Pereopod IV (figs 47B-C) dactylus length sube-

qual to propodal palm. Propodus with 3 broad based

setae on ventral margin, none distinctly larger than

others; articular plate shorter than dactylar claw.

Percopods V-VII (fig. 47E) basis dorsal ridge in cross

section angular.

Penes (lig. 47F) length 0.26 body width at pereonite

7. distally tubular.

Pleopod (figs 48, 47G, 4911) I exopod distal margin

rounded; medial margin concave from proximal to dis¬

tal half, subparallcl to lateral margin; dorsal surface

lacking setae. Pleopod II endopod appendix masculina

length 0.59 pleopod length.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

511

Figure 47. Gariwenieiis turretemis gen. nov., sp. nov. Paratype male (AM P6I429), paratype female (AM P61428).

A, male pereopod II. B-C, male pereopod IV. D, female pereopod IV. E-F, male pereopod VII, including

proximal articles, with penes. G, male pleopod II appendix masculina and endopod. Scale bar 0.5 mm.

Uropod (figs 49C-D, F-G) total length 1.04 pleotel-

son length. Protopod length 0.36 uropod total length;

dorsomedial margin in dorsal view strongly concave

laterally, margin setae absent. Exopod length 0.63

endopod length.

Sexual dimorphism, female differences. Antenna length

0.31 body length, nagellum length 0.63 total antenna

length, with 17 articles. Pereopod I (figs 46C-E)

dactylus length 1.13 palm length; propodal palm with 5

stout denticulate .serrate setae, 2 stout robust conical

simple setae and 3 elongate broad based setae. Uropod

length 1.06 pleotelson length, protopod length

0.44 uropod total length, exopod length 0.8 endopod

length.

Distribution. Stony Creek, above and below

Turret Falls, Grampians National Park.

Remarks. Gariwerdeus turretensis sp. nov. is

easily distinguished from other species in the

genus and from Naiopegia xiphagrosUs sp. nov.

by the enlarged male pereopod I propodus, much

larger than males of the other species where the

first pereopods of males resemble those of

females. A short, almost truncate pleotelson with

a highly abbreviated medial lobe that lacks robust

setae, and an inflated article 6 on the antennula,

also serve to identify this species.

512

G.D.F. WILSON AND S.J. KEABLE

Figure 48. Gariwerdens turrelensis gen. nov., sp. nov. Paratype male (AM P61427). A-E, pleopods I-V. Scale bar

0.5 mm.

Garhverdeus heehivensis sp. nov.

Figures 50-57

Material examined Holotypc. Victoria, among roots of

treefem at the base of Beehive Falls, Mud Hut Creek,

near Roses Gap, Grampians National Park, 36°58.54 S

142°27.0rE (GPS), hand sieves, pH 4.6, 10.3°C, G.

Wilson, R. Wetzer and S. Keable, 22 Sep 1999, VIC-

97, NMV J40735 (male bl 7.3 mm).

Paratypes. All lots collection details as for holotypc.

AM P6I276 (39 ind., including male bl 7.8 mm, female

bl 5.4 mm used to supplement description). AM P61430

(male bl 8.7 mm. dissected for illustration, SEM .and

description), AM P6I43I (female bl 5.9 mm, dissected

for SEM and description).

Other material. Scries from Vic., Grampians, col¬

lected hand sieves, G. Wilson. R. Wetzer and S. Keable,

Sep 1999, preserved in 95% ethanol - base of Fish

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

513

[ijiilK

!!■ '

H|h iIS

Figure 49. Gariwerdeus lurrelemis gen. nov., sp. nov. Paratype male (AM P6I429), paratype female (AM P6I428).

A-D, G, male pleotelson and uropod, dorsal, vcniral and lateral views. E, suctorian ciliate protozoan epibiont on

male uropod. F, male uropod protopod distal margin, ventral view. H, female pleopod II. Scale bar 0.5 mm.

514

G.D.F. WILSON AND S.J. KEABLE

Figure 50. Gariwerdeus beehivensis gen. nov., sp. nov. Holotype male (NMV J40735), lateral view. Scale bar 1 mm.

Figure 51. Gariwerdeus beehivensis gen. nov., sp. nov. Paratype male (AM P61430), paratype female (AM

P61431). A, male head, dorsal view. B, female antennula distal articles. C, male antenna proximal articles. D, male

antenna flagellum articles. E. female antennula, dorsal view. Scale bar 0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

515

Figure 52. Gariwerdens beehivemis gen. nov., sp. nov. Paratype male (AM P6I430). A, D-E, left mandible.

B-C, right mandible. Scale bar 0.1 mm.

Falls, Mackenzie R.. 37°06.5<-rS l42“24.0rE (GPS),

from reedy pool close to main falls, pH 6.6, I0.6°C, 20

Sep, VIC-86, AM P6I277 (male, female); S side of

Goulton Gorge, 36°55.89-S 142°25.09'E (GPS), gravel

from roots of trees and grasses half way up steep water

course, pH 6.7, I4.3“C, 22 Sep, VlC-98, AM P6I278

(30 ind.); Goulton Creek on west side of PohIner Road,

36°58.35'S I42°24.83'E (GPS), roots and sediment. pH

6.1, 17.1 °C, 22 Sep, VlC-99. AM P61279 (18 ind.) and

NMV J40736 (male. 2 females).

Etymology. This species name is derived from the

type locality. Beehive Falls (on Mud Hut Creek)

in The Grampians National Park.

Diagnosis. Pleotelson medial lobe with 2 robust

smooth setae and several small simple setae.

Antennula terminal segment inflated and bulbous;

article 5 distal articulation rudimentary, length

less than article 6 length, lacking aesthetasc.

Mandible right lacinia mobilis with 1 row of

denticles. Pereopod I of male dactylus ventro-

distal margin with row of thin scale-like spines;

propodus dorsal margin not produced prox-

imally; propodal palm cuticular fringe strongly

developed, extending along length of setal row.

Pereopods II-IV basis dorsal margin elongate

setae in row, not clustered. Pleopod II endopod

appendix masculina distal tip broadly rounded,

with 3 setae on margin. Uropod distoventral

margin with 2 robust distally spinose setae,

simple setae absent.

Description based on male. Coloration in life brown

mottled (although brown colour mostly appears to be

fine sediment) with white head, white patch in eye

region.

Head (fig. 51 A) length subequal to width in dorsal

view; width 0.83 pereonite 1 width; dorsal surface with

dense cuticular hairs, setae sparse.

516

G.D.F. WILSON AND S.J. KEABLE

Figure 53. Gariwerdeus heehivemis gen. nov., sp. nov. Paratype male (AM P61430). A, paragnaths.

B-C, ma.xillula. D-E, maxilla. F-G, maxilliped. Scale bar 0.2 mm.

Pereon broad, smooth, width exceeding head

width (1.55 head width). Typhlo.sole minimal, ventral

invagination forming inverted ‘u’ shape in cross

section.

Pleoiiiles 2 length less than half length of 5 in dorsal

view, 3^ respective lengths more than half length of 5;

1^ relative lengths unequal, increasing in length from

anterior to posterior, width 1.44 composite length in

dorsal view.

Pleoiclson (Figs 57A-D) lateral length 0.1 body

length, 0.61 depth; dorsal length 1.03 width: depth 1.38

pereonite 7 depth. Dorsal surface with abundant long

setae. Medial lobe width 0.33 pleotcison width, greatest

length 0.07 pleotelson total length. Lateral lobes medial

length 0.1 pleotelson total length. Ventral margin anter¬

ior to uropods with 5-9 denticulate setae (anterior 5

setae large and robust, posterior 4 less robust, inter¬

spersed with fine setae), posterior seta subequal to

anterior adjacent setae.

AiUenmiUi (figs 5IB, E) length 0.07 body length,

with 6 articles, 1-2 tiny aesthetascs on terminal

article. Anienmi (figs 51 A. D) length 0.21 body length.

Flagellum length 0.57 total antenna length, with 12

articles.

Mouthfwld clypeus broader on left side, width 0.44

head width. Mandible (fig. 52) palp length 0.64

mandible length; article 3 with 6 setae; separate distal

group of setae present; articles 1-2 with elongate setal

row. Right incisor process with 3 ctisps. Left spine row

with 8 spines, 6 bifurcate, total count includes 2 on mar¬

gin between pedunculate projection and molar. Right

spine row with 10 spines, 8 bifurcate, total count

including 2 on margin between pedunculate projection

and molar. Molar process length subequal to width;

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

517

Figure 54. Cariweniens beehivensis gen. nov., sp. nov. Paratype male (AM P6I430), paratype female (AM

P6I43I). A-B, E, male pereopod I. C-D, female pereopod I. Scale bar 0.5 mm.

complex sclulaie spines fomiing posterior row (left) or

ciliated spines forming posterior row (right, 2). MaxU-

luhi (figs 53B-C) medial lobe length 0.82 lateral lobe

length; width 0.73 lateral lobe width. Lateral lobe distal

margin with 3 denticulate robust setae, 9 smooth robust

setae. Maxilla (figs 53D-E) medial lobe width 1.0 outer

lateral lobe width. MaxillipecI (figs 53F-G) endite with

3 receptaculi on right side; dorsal ridge with 10 large

distally denticulate plumose setae. Palp insertion on

basis ventral surface with I subdistal smooth seta.

Pereopod I (figs 54A-B. E) dactylus length subequal

to palm, length 1.03 palm length; ventrodistal margin

thin scale-like spines along 0.28 total length of margin.

Propodus dorsal margin with 10 setae in several groups

between proximal and distal margin (c.xeluding distal

group). Propodal palm sinuate; cuticular fringe weakly

developed; stout denticulate .setae absent; with 12 stout

robust simple conical setae; elongate broad based setae

absent. Basis ventrodistal margin with 9 elongate setae

in male. Pereopod IV (figs 55B-C) dactylus length

subequal to propodal palm; distal accessory claw-spines

absent. Propodus with 5 broad based setae on ventral

margin, 3 distinctly larger than others; articular plate

shorter than dactylar claw. Pereopods V-VII (fig. 55E)

basis dorsal ridge in cross section rounded on pereopod

V, angular on pereopods VI-VII.

Penes (fig. 55F) length 0.33 body width at pereonitc

7, distally tapering.

Pleopod (figs 56, 57G-I) I exopod distal margin

pointed, medial margin sinuate - divergent from lateral

margin along most of length, dorsal surface with setae.

Pleopod II endopod appendix masculina length 0.55

pleopod length.

Uropod (figs 57D-F) total length l.ll pleotelson

length. Protopod length 0.39 uropod total length; dorso-

medial margin in dorsal view parallel to lateral margin,

margin setae present distally. Exopod length 0.68 endo¬

pod length.

Sexual dimorphism, female differences. Anlenna length

0.25 body length. Flagellum iength 0.52 total antenna

length, with 11 articles. Pereopod\ (figs 54C-D) dacty¬

lus length 0.93 palm length, ventrodistal margin with 6

narrow scale-like spines, along 0.37 total length;

518

G.D.F. WILSON AND S.J. KEABLE

m ., %m

fe”*' ’"'v.

Figure 55. Gariwenkus heehivensis gen. nov., sp. nov. Paratypc male (AM P61430), paratype female

(AM P6143I). A, male pereopod 11. B-C, male pereopod IV. D, female pereopod IV. E-F, male pcreopod VII,

including proximal articles, with penes. Scale bar 0.5 mm.

propodal palm with 9 stout robust conical simple setae;

basis ventrodistal margin with 7 elongate setae. UropoJ

protopod length 0.46 uropod total length; exopod length

0.83 endopod length.

Distribution. Beehive Falls, Mud Hut and Goul-

ton Creek drainages and Fish Falls (Mackenzie

R.), Grampians National Park.

Remarks. Several characters distinguish Gariw-

erdeus heehivensis sp. nov. from other species of

the genus: the 2 inflated distal antennular seg¬

ments have only a rudimentary articulation with

article 5 lacking aesthctascs. the head is mgose

with many fine cuticular hairs, and the medial

lobe of the pleotelson has 2-4 robust setae.

The specimens from Goulton Gorge and Goul-

ton Creek (AM P61278-9) contains larger speci¬

mens than found in the types (males reaching a

length of 11.8 mm versus 8.7 mm in type mate¬

rial). These larger specimens have more promi¬

nent medial and lateral lobes on the pleotelson,

and have more robust setae on the medial lobe (4

versus 2). The same samples, however, have

specimens of similar size to, and indistinguish¬

able from, the type material. The male and female

specimens from Fish Falls (AM P61277) are of

similar size to the type material and have similar

posterior pleotelson lobes; the male from this

sample has four robust setae on the medial lobe

and the female has three.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

519

Figure 56. Gariwerdeus beehivemis gen. nov., sp. nov. Paratype male (AM P6I430). A—E, pleopods I—V. Scale bar

0.5 mm.

Gariwerdeus ingletonensis sp. nov.

Figures 58-65

Material examined. Holotype. Vic., Ingleton Spring at

picnic ground off Syphon Road, Grampians National

Park, 37°I8.17'S l42°22.0rE (GPS), in white sand at

point where spring Hows from under a rock, hand

sieves, pH 4.7, I2.7°C, G. Wilson, R. Wetzer and S.

Keable, 21 Sep 1999, VIC-94, NMV J40737 (male bl

7.2 mm).

Paratypes. All lots collection details as for holotype.

AM P61280 (26 ind., including female bl 6.6 mm used

to supplement description), AM P6I432 (male bl 8.3

mm, dissected for SEM and description), AM P61433

(female bl 6.7 mm, dissected for SEM and description),

AM P6I444 (male bl 7.9 mm, dissected for pleopod

illustrations), NMV J40738 (male, female, juvenile

female).

Etymology. As in other species of Gariwerdeus,

520

G.D.F. WILSON AND S.J. KEABLE

Figure 57. Gariwerdeus beehivensis gen. nov., sp. nov. Paralype male (AM P6I430), paratype female (AM

P6143I). A-D, male pleotelson and uropod, dorsal and lateral views. E, male uropod lateral view. F, male uropod

protopod distal margin, ventral view. G, female pleopod I. H, male pleopod I. 1, male pleopod II appendi.\

masculina and endopod. Scale bar 0.25 mm.

this species is named after the type locality. Ingle-

ton Spring, The Grampians National Park, in this

case.

Diagnosis. Pleotelson medial lobe robust sensil-

late setae absent (but with 6 fine simple setae).

Antennula with 5 free articles, articles 5 and 6

inflated, each with 1 aesthetasc, terminal segment

with unexpressed articulation. Mandible right

lacinia mobilis with 2 rows of denticles. Pereo-

pod I dactylus ventrodistal margin with thin

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

521

Figure 58. Gariwerdeus inglelonensis gen. nov., sp. nov. Holotype male (NMV J40737), lateral view.

Scale bar 1 mm.

truncate spines; propodiis dorsal margin not pro¬

duced proximally, palm cuticular fringe weakly

developed. Pereopods II IV basis dorsal margin

elongate setae divided into proximal and distal

cluster. Pleopod II cndopod appendix masculina

distal tip truncate, with 6 setae on margin. Uropod

protopod distoventral margin with 2 robust

distally spinose setae and 2 robust simple setae.

Description based on mate. Coloration in life creamy

white, head with bright white patch in eye region; in

95% ethanol, cream with lighter white patch in eye

region.

Head (fig. 59U) length shorter than width in dorsal

view; width 0.82 pereonile I width; surface with dense

cuticular hairs and scattered setae.

Pereon broad, width exceeding head width (1.32

head width); smooth. Typhlosole minimal, ventral

invagination forming laminar projection in cross

section.

Pk’onites 2-4 respective lengths more than half

length of pleonite 5 in dorsal view; pleonites 1-4

relative lengths unequal, increasing in length from

anterior to posterior, width 0.84 composite length in

dorsal view.

Pleotelson (figs 65A-B) lateral length 0. II body

length, 0.7 depth; dorsal length 1.06 width; depth 1.5

pereonitc 7 depth. Dorsal surface with abundant long

setae. Medial lobe width 0.35 pleotelson width, greatest

length 0.06 pleotelson total length. Lateral lobes medial

length 0.14 pleotelson total length; lateral lobes robust

sensillate setae absent. Ventral margin anterior to

uropods with 5 denticulate and smooth setae (posterior

setae weakly denticulate), posterior seta subequal to

anterior adjacent setae (length subequal posterior seta

slightly thicker).

Antenmda (figs 59E-I I) length 0.1 body length, with

5 articles, 2 liny aesthetascs on terminal article.

Antenna (figs 59A-B) length 0.32 body length. Flagel¬

lum length 0.61 total antenna length, with 14-15

articles.

Mouthfteki clypeus projecting anteriorly in medial

region, width 0.48 head width. Mandible (fig. 60) palp

length 0.64 mandible length; article 3 with 6 setae;

articles 1-2 with elongate setal row. Left spine row with

11 spines, 8 bifurcate, total count including 3 on margin

between pedunculate projection and molar. Right spine

row with 15 spines, 9 bifurcate, including 6 on margin

between pedunculate projection and molar. Molar pro¬

cess length subequal to width; complex sctulale spines

fomiing posterior row (both sides). Maxillula (figs

6IB-D) medial lobe length 0.85 lateral lobe length;

width 0.73 lateral lobe width. Lateral lobe distal margin

with 4 denticulate robust setae, 8 smooth robust setae.

Maxilla (figs 6IE-F) medial lobe width 0.8 outer lateral

lobe width. MaxiUiped (figs 61G-H) endite with 4

receptaculi on right side; dorsal ridge with 13 large dis¬

tally denticulate plumose setae. Palp insertion on basis

ventral surface with 9 subdislal smooth setae (forming

row).

Pereopod 1 (fig. 62A) dactylus length subequal to

palm; ventrodistal margin thin denticulate spines along

0.51 total length. Propodus dorsal margin with 5 setae

in several groups between proximal and distal margin

(excluding distal group). Propodal palm concave, with

6 stout serrate setae and 4 elongate broad based setae.

Basis ventrodistal proximal margin with 8-14 elongate

setae. Pereopod IV (fig. 63B) dactylus length subequal

to propodal palm. Propodus with 4 broad based setae on

ventral margin, I distinctly larger than others; articular

plate longer than dactylar claw. Pereopods V-Vll (fig.

63C) basis dorsal ridge in cross section angular.

522

G.D.F. WILSON AND S.J. KEABLE

Figure 59. Gariwerdeus inglelonenxis gen. nov., sp. nov. Paratype female (AM P6I433), paratype male (AM

P6I432). A-B, female antennula and antenna. C-D, male head, anlennula and antenna, dorsal view. E-F, male

antennula distal articles. G, female antennula, dorsal view. II, female antennula distal articles. Scale bar 0.5 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

523

Figure 60. Gariwenieiis inglelonensis gen. nov., sp. nov. Paratype male (AM P6I432). A-C, G, left mandible.

D-F, right mandible. Scale bar 0.1 mm.

Penes (fig. 63F) length 0.19 body width at pereonite

7, distally tapering.

PleopocI I (figs 64, 63D-E) exopod distal margin

rounded, medial margin concave from proximal to dis¬

tal half, subparallel to lateral margin, dorsal surface

lacking setae. Pleopod II endopod appendix masculina

length 0.54 pleopod length.

UropocI 65A, C-D) total length 1.22 pleotelson

length. Protopod length 0.46 uropod total length; dorso-

medial margin in dorsal view concave laterally, margin

setae present distally. Exopod length 0.8 endopod

length.

Sexual dimorphism, female differences. Antenna fiagel-

lum length 0.63 total antenna length, with 12-14 arti¬

cles. Pereopod I (figs 62B-D) dactylus ventrodistal

margin with 8 hroad cuticular fringe spines. Pereopod

IV propodus with 1 broad based seta on ventral margin.

Uropod total length 1.1 pleotelson length; exopod

length 0.89 endopod length.

Distribution. Ingleton Spring, Grampians

National Park.

Remarks. An abbreviated antennule, consisting of

only 5 segments, is the most distinctive feature of

Gariwenieus inglelonensis sp. nov. The inflated,

distally expanding distal antennular segment is a

composite of two segments with their articulation

being unexpressed, but with two separate

aesthetascs present. This morphology is similar to

524

G.D.F. WILSON AND S.J. KEABLE

4 J

Figure 61. Gariwerdeus ingletonemis gen. nov., sp. nov. Paratype male (AM P6I432). A, paragnaths.

B-D, maxillula. E-F, maxilla. G-H, maxilliped. Scale bar 0.2 mm.

NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

525

Figure 62. Gariwerdeus ingletonensis gen. nov., sp. nov. Paratype male (AM P6I432), paratype female (AM

P6I433). A, male pereopod I. B-D, female pereopod 1. Scale bar 0.5 mm.

that seen in amphisopodid species, where the

terminal antennular segment is also composite. Of

the two species of Gariwerdeus that lack robust

setae on the medial lobe of the pleotelson,

G. ingletonensis has a more rounded posterodor-

sal margin of the pleotelson in lateral view than

G. turretensis where the pleotelson appears

almost truncate in lateral view.

Acknowledgments

We thank the following people and institutions

for their part in our research. Tim Doeg sent us

specimens of Synamphisopus collected as part of

a project conducted by the Flora and Fauna

Branch, Department of the Natural Resources and

Environment, Victoria. John Clamp advised us on

protist epibionts. Regina Wetzer participated in

the field trip wherein most specimens were

collected, and is assessing DNA sequences from

various phreatoicideans. Anna Cerra dried and

mounted specimens and obtained the SEM

images. Kate Dempsey prepared the plates. Sue

Bullock inked some of our pencil drawings and

Rick Johnson assisted with managing the DELTA

database. Gary Poore and a referee for this journal

provided useful suggestions for improvement of

the manuscript. Gary also advised us of the cor¬

rect publication date for the Spencer and Hall

description of Phreatoicopsis. This research was

supported a grant from the Australian Biological

Resources Survey, and is a contribution ol the

Australian Museum Centre for Evolutionary

Research.

References

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526

G.D.F. WILSON AND S.J. KEABLE

Figure 63. Gariwerdeus ingletonensis gen. nov., sp. nov. Paralype male (AM P6I432). A, pcrcopod II. B, pereopod

IV. C, F, pereopod VII, including proximal articles, with penes. D-E, picopod II appendix masculina.

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NEW PHREATOICIDEAN CRUSTACEA FROM GRAMPIANS NATIONAL PARK

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528

G.D.F. WILSON AND S.J. KEABLE

Figure 65. Gariwerdeus inglelonensis gen. nov., sp. nov. Paratype male (AM P61432). A-B, pleotelson and uro-

pod, lateral and dorsal views. C, uropod, medial view. D, uropod protopod distal margin, ventral view.

Scale bar 0.5 mm.

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/•

‘ {

‘V

Memoirs of the Museum of Victoria 59(2): 531-540 (2002)

REDESCRIPTIONS OF ASPHALIDESMUS LEAE SILVESTRI, 1910

(CHAMBERLIN, 1920) COMB. NOV. FROM TASMANIA, AUSTRALIA (DIPLOPODA.

POLYDESMIDA: HAPLODESMIDAE)

Robert Mesibov

Queen Victoria Museum

and Art Gallery, Wellington Street, Launceston,

(e-mail: mesibov@southcom.com.au)

Tasmania 7250, Australia

Abstract

Mesibov, R., 2002. Redescriptions of

(Chamberlin, 1920) comb. nov. from Tasmania Australia ( p P

Haplodesmidae). Memoirs of Museum .l^UaUdesmus Silvestri, 1910.

Lpodesmus Chamberlin, 1920 is synonym.sed comb. nov. are

Asphalidesmus leae Silvestri, 1910 and A. panu. L . jujpgjg species appear to be

redescribed from abundant, recenOy collected material. The two milhpeae spe (

endemic to Tasmania but have widely separated ranges.

Introduction

Early last century, Filippo Silvestri and Ralph

Chamberlin independently described small

Tasmanian polydesmoid millipedes with dense

metatergal tuberculation and laterally expanded

paranota on the second tergites. Each man

established a monotypic genus for the material

he examined: Asphalidesmus Silvestri, 1910 for

a male specimen and Atopodesnnis Chamberlin,

1920 for two females. Neither Silvestri

nor Chamberlin assigned his genus to a family

and the correct taxonomic placement ol

Asphalidesmus and Alopodesmus has long been a

myriapodological puzzle.

Attems (1926) and Verhoeff (1932) placed

Asphalidesmus in Fontariidae Attems, 1926, but

Attems (1940) later regarded Asphalidesmus as a

genus of uncertain status within Vanhoetfeniidae

Attems, 1914.

Verhoeff (1932) ignored Alopodesmus entirely

because it had been described in a work (Cham¬

berlin, 1920) which lacked illustrations and was

therefore unscientific: ‘...diese Arbeit [kann)

Weder beriicksichtigt werden noch iiberhaupt als

wissenschafllich eelten’ (Verhoeff, 1932: 1981).

Attems (1926) at first placed Alopodesmus m

^’ryptodesmidae Karsch, 1879, but later consic-

cred it an uncertain genus within Oniscodesmidae

de Saussure, 1860 (Attems, 1940).

Hoffman (1980) included Alopodesmus m his

bst of Polydesmidea of uncertain status an

nily position. He placed Asphalidesmus n

lodesmidae Cook, 1896, noting that the genus

Ttainly appears to be a senior synonym of Tos-

miosLa [a Tasmanian dalodesmidj, but tte

ibability should be confirmed by the study ot

ual niLrial, to include type material of

modesmus and Lissodesmus [another Tasma-

^ d.lodesmidj, of rfiich the gonopod structure

„ill unknowrt' (Honmau. '»*<> '50),

Strrn2.toX»

d otomids. Silveslri’s descnpuoo of an

rand“»pe.r»o seemed ^

the South

=rTi«r;5o"hrrt:

“'’"‘‘it

;sion ot S miiar po y from New

SfnU) reduced Vaalogotropo-

rhoefi, ^Haplodesmidae Cook,

3 new fr,ined Haplodesminae

-ind Prosopodesminae, which had

reTSel h, Hon-man (I*

531

532

R. MESIBOV

Jeekel (1984) concluded by accepting the two

Tasmanian genera as haplodesmids, but without

subfamily placement.

The following year, Jeekel (1985) published a

key to families and non-paradoxosomatid genera

of eastern Australian Polydesmida. According to

the key. the non-sexual characters reported for

Asphalidesmus and Atopodesmus would place

these genera in llaplodesmidae. However, the

only eastern Australian genus referred to this fam¬

ily in Jeekel’s key is Agallwdesmiis Silvestri,

1910. based on Agathodesmus steeli Silvestri,

1910 from New South Wales. Agathodesmus had

not been placed in a family by Attems (1940) and

had been assigned to Dalodesmidae by HotTman

(1980).

In a later paper, Jeekel (1986) referred to his

earlier discussions of Australian llaplodesmidae

and noted that ‘In the meantime. Dr P. M. Johns,

Christchurch. New Zealand (pers. comm.)

assured me that in his opinion Asphalidesmus

Silvestri is a member of the family Dalodesmidae

rather than a haplodesmid. This leaves only

Atopodesmus Chamberlin and Agathodesmus

Silvestri as potential Australian Haplodesmidae.

Unfortunately, the male characters of these two

genera are unknown, so that as yet no certainly

can be obtained’ (Jeekel, 1986: 46). Jeekel (1986:

35) went on to describe a new haplodesmid

species from Queensland, Atopogonus hucculen-

tus, characterising its collection in 1981 as ‘the

first unquestionable record of the group from

Australia.’

In this paper 1 redescribe Asphalidesmus leae

Silvestri, 1910 and Atopodesmus parvus Cham¬

berlin, 1920. The two species have widely

separated ranges in Tasmania but are locally

abundant, and my descriptions arc based on

examination of more than 500 specimens

deposited in the Queen Victoria Museum.

Launceston, Australia. 1 do not consider that there

are sufficient differences in gonopod structure

and non-sexual details to maintain separate

genera for these Tasmanian fomts, and 1 therefore

reduce Atopodesmus to a synonym of the older

Asphalidesmus, which I redefuie below.

Haplodesmidae seem to have become a ‘tem¬

porary storage area' for genera of small poly¬

desmoids with dense metatergal tuberculation and

lateral expansion of the collum or the paranoia of

the second tergite. Until more information

becomes available about species in this group, it

seems wisest to accept Jeekel's 1984 placement

of Asphalidesmus in Haplodesmidae without

assignment to a subfamily.

Haplodesmids arc widespread in Australia and

probably species-diverse (Black, 1997). At least

one other Tasmanian species is in the Queen

Victoria Museum collection, known from two

female specimens from King Island; this elusive

species will be described when mature males are

collected.

Specimens listed under Material examined

were killed and preserved in 75-80% ethanol.

Preliminary drawings on graph paper were made

using material cleared in 60% lactic acid and

viewed at 100 or 200 x magnification through an

eyepiece graticule. A Philips Electroscan ESEM

2020 operated in high-vacuum mode was used to

examine preserved material which had been air-

dried before sputter-coating with gold. SEM

images were acquired digitally.

Note on spatial data. Universal Transverse

Mercator (UTM) grid references are the spatial

locators u.sed by most field workers to define eol-

lecting localities in Tasmania. Collecting sites for

all but a few' of the specimens listed under Mate¬

rial examined (online at ww'w.museum.vic.gov.

au/memoirs/index.html) were estimated in the

field to be within particular 100 m UTM grid

squares on 1:25000 scale maps published by the

State of Tasmania. The maximum horizontal error

in these estimates is likely to be less than 100 m.

Latitude/longitude equivalents given were cal¬

culated using GeoCalc 4.20 (GeoComp Systems,

Blackburn, Victoria) and are not as precise as the

UTM grid references.

Abbreviations. MCZ, Museum of Comparative

Zoology, Harvard University, Cambridge,

Massachusetts, USA; QVM, Queen Victoria

Museum and Art Gallery, Launceston, Tasmania,

Australia.

Order Polydesmida Leach, 1815

Suborder Polydesmidea Leach, 1815

llaplodesmidae Cook, 1895

Asphalidesmus Silvestri

Asphalidesmus Silvestri, 1910: 362.—Attems 1926:

153.—Verhoeff, 1932: 1587.—Verhoeff, 1936: 2.—

Attems, 1940: 205.—Jeekel, 1971: 313.--Hoffman,

1980: 150, 184.—Jeekel, 1984: 85.

Atopodesmus Chamberlin, 1920: 153.—Attems,

1926: 134.—Attems, 1940: 356.—Jeekel, 1971- 313 —

llolfman, 1980: 150, 186.—Jeekel, 1984: 85 (type

species: Atopodesmus parx’us Chamberlin, 1920 by

original designation), (syn. nov.).

Type species. Asphalidesmus leae Silvestri, 1910,

by original designation.

REDESCRIPTIONS OF TWO MILLIPEDES FROM TASMANIA (DIPLOPODA)

533

Included species. A. leae, A. parvus (Chamberlin,

1920) comb. nov.

Description. General appearance. Head plus 19

somites (fig. I) in both sexes. Mature males 5-6

mm long, 0.7-0.9 mm in overall midbody width;

mature females very slightly larger. Immature and

some mature individuals completely unpig-

mented, appearing white to naked eye. Most

mature (stadium VII) individuals pale yellow-

brown on collum, metatergites and preanal ring,

deepening to brown towards waist and bases of

paranota, remainder of body very pale yellow,

whole animal appearing yellow-brown to naked

eye. Most mature individuals, but no immatures,

encrusted with minute soil particles (see

Remarks).

Head with granulated vertex to level of anten¬

nal sockets, otherwise smooth (fig 2), vertigial

sulcus short and weakly impressed. Antennae

short with stout antennomeres, sixth being longest

and widest (fig. 2).

Collum almost entirely concealing head when

extended animal is viewed from above, strongly

convex and fitting head like a cap (fig. 2). Col¬

lum, metatergites and preanal ring with transverse

rows of numerous, unifomi tubercles (figs 4,6,8)

on roughened cuticular surface, each tubercle

bearing short, curved seta with a flared tip (fig. 4).

Midbody tergites of mature individuals typically

with 5-6 rows of tubercles.

Somites each with clearly defined prozonite

and metazonite and with paranota based low on

sides (fig. 3). Podous segments diminishing only

slightly in width from anterior to posterior. Pro¬

zonite surface faintly reticulated, caudal margin

of metazonite continuously fringed with minute

teeth, each with tip bent away from body at a right

angle (figs 3, 4, 6, 8).

Paranota of segment 2 expanded, extending

forward to partly cover collum and backward to

lie under the anterior edge of the paranota of seg¬

ment 3, margin divided into 7-9 seta-bearing

lobes (fig. 2). Remaining paranota (figs 6, 8)

more or less uniform in size, those of segment 18

substantially smaller. Each paranotum flexed

slightly downward (fig. 3), the posterior edge

lower than the anterior edge. Anterolateral border

of paranotum roughly arc-shaped with series of

3-4 indistinctly defined seta-bearing lobes, poste¬

rior border more or less at right angles to body

with flat, round-bordered outgrowth, here called a

paranoial lag, close to body (figs 6, 8). On most

segments few seta-bearing tubercles similar to

those on metatergite can be noted on inflated

basal portion of paranotum.

Ozoporcs on segments 5, 7, 9, 10, 12, 13,

15-17, above and distal to centre of base of para¬

notum (figs 6, 8), each ozopore opening within

short, column-like structure with well-defined,

circular, upper rim (fig. 5).

Legs short (fig. 2), the tarsus being longest

podomere. Setation nomial (no sphaerotri-

chomes). Male leg podomeres somewhat inflated

relative to those on female legs, male postfemur

and tibia proportionally longer than those of

female.

Telson with preanal ring completely covering

anal valves as viewed from above, valves opening

ventrally. Few macrosetae at apex of epiproct,

2+2 macrosetae on anal valves, 2 long macrosetae

on hypoproct.

Male with ovoid gonopod aperture, posterior

margin slightly raised (fig. 2). Gonopods in situ

with co.xae entirely contained within cavity,

telopodites reaching to anterior leg-pair of seg¬

ment 5 (fig. 2). Gonopod coxae short, tapering

towards distal end, lightly joined (not fused)

medially. Telopodite slender, setiferous on pos¬

terolateral surface from base to half to two-thirds

of length, then splitting into 2 branches: anteriorly

directed process which bends caudad, and poster¬

iorly directed process bearing end of prostatic

groove (figs 7, 9). Female with narrow, incon¬

spicuous epigynal slit; cyphopods not examined.

Juveniles very similar to adults in general appear¬

ance and details of tuberculation, but with less

prominent paranota.

Distribution and ecology. Asphalidesmus spp. are

so far known only from two widely separated

areas in Tasmania (fig. 10). Within their respec¬

tive ranges, A. leae and A.panms occur on a vari¬

ety of bedrocks and landforms in areas with

annual rainfall from c. 650 to 2000 mm (winter

maximum). Both species, however, are restricted

to cucalypt forest and Nothofagus rainforest,

within which they are found in and under wet rot¬

ting wood and in moist accumulations of leaf lit¬

ter. Asphalidesmus spp. arc gregarious and typi¬

cally occur in multi-aged groups of 10-50

individuals spread over c. 0.1 m\ They are excep¬

tionally slow-moving millipedes. When disturbed

they neither enroll, curl, walk quickly away nor

readily release their grip on the substrate. Mating

pairs are found throughout the year, although so

far only one sample {A. parvus, QVM 23:8360)

contains a pair preserved in copula.

Remarks. The differences between the gonopods

of the two known Asphalidesmus species

are large in comparison to those within many

534

R. MESIBOV

better-known polydesmoid genera. On the other

hand, the two species are virtually identical in

nonsexual morphology, share some striking

synapomorphies and are both apparently endemic

to Tasmania. Thus the gonopod differences are

likely to have evolved within a single lineage.

The Chamberlin name Alopodesmus remains

available if molecular studies of Asphalidesmus

and new studies of related Australian poly¬

desmoids justify the (re)placement of the

Tasmanian species in separate genera.

It is remarkable that body colouring and

encrusting soil matter are invariably absent in

immature stadia ot Aspliulidesmus and almost

always present in mature individuals, despite

the fact that matures and immatures live

side by side in the same microhabitats.

Is there a cuticular change in stadium VII

which allows Asphalidesmus to adsorb coloring

matter from soil and decorate itself with

particles?

Asphalidesmus leae Silvestri

Figures 1-4, 6, 7, 10

Asphalidesmus leae Silvestri. 1910: 362, fig. 8.—

Attems, 1940: 205.—Jcekel. 1971: 313.—Jeekel, 1984:

85.

Holotype. Mature male, ‘Hobart (Tasmania, A. Lea

legit)’ (Silvestri, 1920: 362), possibly in the Silvestri

type collection in Portici, not examined.

Material examined. 166 specimens in QVMAG from

80 unique localities (Fig. 10) in northern Tasmania:

Alami River. Allen Creek, Arthur River, Big Creek,

Bishops Creek, Black River. Blaekfish Creek, Bonneys

Tier, Borradale Creek, Brampton Creek. Cam River,

Camp Creek, Chasm Creek, Christmas Hills, Crayfish

Creek. Detention Falls. Devils Gate, Dial Creek,

Dynans Bridge. East Gawler River, East Ridgley, Flow-

erdale River, Frankland River, Gawler River. Gog

Range, Harris Creek, Hebe River, Holwcll Gorge,

Inglis River. Julius River, Keddies Creek, Kclcey Tier,

Kenzies Hill. Langford Creek. Lawson Plains.

Lcbrina, Library Creek, Little Claytons Rivulet,

Little Donald.son River. Lobster Rivulet. Lone Star.

Long Hill, Loyetca Peak, Lunta Tier, Meryanna,

Meunna Hills, Montagu Swamp, Mt Arthur. Mt Riana,

Paloona Dam. Peegra Road, Pioneer Link. Punchs

Terror, Retreat. Roger River West. Salmon River,

Saxons Creek. Sideling Range, Sisters Creek,

Somerset, The Clump, Trowutta Caves, Venns

Creek, Wyena, York Town Rivulet. For full

details of localities, dates, collectors, specimens and

registration numbers, see the online list at

www.museum.vic.gov. au/memoirs/index.html, or

contact the author.

Diagnosis. Differs from A. parvus as follows: (a)

anterior process of gonopod telopodite slender,

terminating at or below the level of the posterior

process in a short, caudally directed hook; (b)

paranotal tabs weakly produced or not apparent,

no more prominent than lobes on anterolateral

border of paranoia (late stadium males and

females).

Description. Paranotal tabs weakly produced

(fig. 6) and not apparent on most paranoia.

Gonopod (figs 2, 7) with telopodite divided at

about two-thirds its length into anterior and

posterior processes. Basal portion of the

telopodite fairly straight, tapering mesally, with

short, stout setae on the posterior and postero¬

lateral faces, the setiferous area terminating

just proximal to the telopodite branching point.

Posterior process directed slightly caudad

and laterad, thinning to spatulate structure,

concave mesally with coarsely toothed

terminus, small uncus arising just proximal to

concavity at about one-quarter length of branch.

Prostatic groove running along anterior face of

telopodite, curving caudad and mesad to ter¬

minate in ilattened solenomerite arising within

branch concavity, solenomerite bending laterad

and cephalad and almost extending to level of

branch tenninus. Anterior process of telopodite

slender, bending sharply caudad at about one-

third its length, tapering and temiinating in

caudally pointed hook at about level of tip of

solenomerite.

Distribution. Asphalidesmus leae occurs from

near sea level to c. 750 m in northern and

northwestern Tasmania (fig. 10). Within its

range ot c. 8000 km* A. leae seems to prefer wet

forest habitats. (See also the discussion of

Asphalidesmus distribution and ecology.)

Remarks. Remarkably little morphological varia¬

tion has been noted in A. leae. Gonopods of males

collected 200 km apart, at opposite ends of the

A. leae range, appear identical.

My identification of the QVM material with

A. leae is based on Silvestri’s text description and

gonopod drawing (fig. 7A). However, Silvestri

gives the type locality as Hobart, which is clearly

wrong (fig. 10). The collector, Arthur Lea, was

Government Entomologist of Tasmania begin¬

ning in 1899 (Marks, 1991: 207). Although Lea

was based in Hobart, he was an active field natu¬

ralist and presumably collected A. leae on a trip to

the north of the island sometime between 1899

and 1910. It is unclear how the confusion in type

locality arose.

REDESCRIPTIONS OF TWO MILLIPEDES FROM TASMANIA (DIPLOPODA)

535

Asphalidesmusparvus (Chamberlin) comb. nov.

Figures 1,2, 5, 8-10

Atopock’smus panms Chamberlin, 1920: 154.—

Altems, 1940: 357.—Jeekel, 1971: 313.—Jcekel, 1984:

85.

Hololype. mature female, ‘Tasmania (O.H. Hardy)'

(Chamberlin, 1920, p. 154), MCZ, no. 4648. Paratype:

immature female, collection details as for holotype,

MCZ. no. 4649.

Other material examined: 398 specimens in QVMAG

from 83 unique localities in south-eastern Tasmania:

Anglers Creek. Baldy Creek. Bellettcs Creek, Big Sassy

Creek, Black Hill. Blind Creek (Maria Island). Blue

Gum Spur, Browns Creek, Chauncy Vale, Counsel

Creek (Maria Island), Douglas Creek. Entrance Cave,

Espies Craig, Flash Tier, Four Mile Creek (Maria

Island), Muon River (Ars'c Road), Huon River

(Edwards Road), Huon River (Manuka Road), Lake

Sydney, Lenah Valley, Maegregor Peak. Maclaines

Creek, Mitchclmores Creek. Mother Browns Bonnet.

Mt Hobb.s. Mt Maria (Maria Island), Mt Misery, Mt

Ponsonby. Mt Rumney, Mt Tobin (Bruny Island), Mys¬

tery Creek Cave track. Ned Ryans Creek (Maria

Island), Nugent, Old Fann Road, Organ Pipes (Mt

Wellington). Paradise Gorge. Pendulum Palace (cave

PB-12, Precipitous Bluff), Phipps Creek. Pine Hut

Creek (Maria Island), Pirates Road, Ravens Hill,

Robinsons Creek (Maria Island), Sand River, Sandspit

River, Sassafras Hill, Silver Hill Road, Stormlea.

Strathblane, Tahunc Bridge, Three Thumbs, Tiger

Creek, Tinderbox, Tobys Hill. Tooms White Gum

Reserve, Waterfall Creek. Wilsons Ridge, Woodsdale,

Yarlington Tier. For full details of localities, dates, col¬

lectors, specimens and registration numbers, see the

online list at www.museum.vic.gov.au/memoirs/index.

html, or contact the author.

Diagnosis. Differs from A. leae as follows; (a)

anterior process of gonopod telopodite robust, at

least twice as long as posterior process, tenninat-

ing in a long. Ilattened, caudally directed structure

usually with 10-12 coarse teeth; (b) paranoial

tabs strongly produced in late stadium males and

females, clearly larger and more distinct than

lobes on anterolateral border of paranota.

Description. Paranotal tabs large and apparent on

most paranota (fig. 8). Gonopod (figs 2, 9) with

telopodite divided at just over half its length into

anterior and posterior processes. The basal por¬

tion of the telopodite fairly straight, slender,

tapering mesally, with short, stout setae on pos¬

terior and posterolateral faces, setiferous area ter¬

minating Just proximal to telopodite branching

point. Posterior process directed slightly caudad

and divided into lateral and mesal branches. Lat¬

eral subbranch in turn divided into distal

solenomerite and proximal process which bends

laterad, bears minute teeth and reaches about half

length of solenomerite. Mesal subbranch short,

stout, erect, terminating at level of posterior pro¬

cess on lateral subbranch. Prostatic groove run¬

ning near anteromesal face of telopodite, barely

curved, terminating near tip of slightly flattened

solenomerite. Anterior process of telopodite a

large, anteroposteriorly flattened structure, bend¬

ing sharply caudad at about two-thirds its length

and tapering into laterally flattened comb bearing

c. 10-12 coarse teeth pointed towards posterior

process. From telopodite branch point, anterior

process extending just over twice di.stance

reached by the solenomerite.

Distribution. A. parvus ranges over c. 8000 knT in

southern and south-eastern Tasmania, including

Bruny and Maria Is (fig. 10). It is common in wet

forest and in wet places (flowlines, south-facing

slopes) in dry forest. On the Tasmanian mainland

A. pan’us occurs from near sea level to c. 950 m

(on Mt Wellington, near Hobart). A. parvus has

also been collected in limestone caves in far

southern Tasmania, where it is likely to be a

troglophile or accidental. (See also the discussion

oiAsphalidesmus distribution and ecology.)

Remarks. The original description of A. parvus

includes the line ‘Caudal margin of most keels

with a single large conical tooth close to the base’

(Chamberlin, 1920; 154). A large paranotal tab is

apparent on segments 7-15 of the holotype and

segments 5-15 of the paratype. distinguishing

these specimens as A. parvus rather than A. leae.

Like A. leae. A. pamts varies very little across

its range, but in males from three sites on

Forestier Peninsula (QVM 23:41551, QVM

23:8324 and QVM 23:41528) and one on Tasman

Peninsula (QVM 23:8325) the teeth on the anter¬

ior process of the gonopod are reduced or absent.

The gonopodal teeth appear nonnal on a male

from a nearby ‘non-peninsular’ site (QVM

23:8327). The two peninsulas are already recog¬

nised as a biogeographical subregion of Tasmania

for the presence of a locally endemic snail and

millipede, and the apparent absence of at least

two other terrestrial invertebrates (Mesibov.

1996).

The type locality of A. parvus (Tasmania)

unfortunately remains inexact. However, the col¬

lecting date can be roughly bracketed in time.

G.H. Hardy worked in Western Australia before

sctx'ing as Assistant Curator of the Tasmanian

Museum in Hobart from 1913 to 1917, alter

which he studied at the University of Sydney

(Marks. 1991: 216). The Annual Report of the

536

R. MESIBOV

Director of the Museum of Comparative Zoology

at Harvard College to the President and Fellows

of Harvard College for 1914-1915 records (p. 31)

that material had been received by the MCZ dur¬

ing the year from ‘E.H. Hardy’ (sic.). Ralph

Chamberlin, who at the time was Curator of

Arachnids, Myriopods (sic.) and Worms at the

MCZ, described 14 Tasmanian myriapod species

from Hardy collections (Chamberlin, 1920) and

gives a collecting date ‘February 1915’ for one of

these, the millipede Nolodesmus scotius. Finally,

a possibly original label with the Lissodesmtis

modestus type (no. 4644) says ‘Jan 1915 Russell

Falls’. It thus seems likely that Hardy collected

A. parvus in the period from 1913 to the austral

summer of 1914-1915.

Acknowledgement.s

1 thank Dr David Steele, University of Tasmania,

for taking SEMs of Asphalidesmus specimens. At

the MCZ in October 2000, Dr Gonzalo Giribet

generously provided working space for an exam¬

ination of Chamberlin types, and Special Collec¬

tions Librarian Dana Fisher kindly found docu¬

ments for me from the Chamberlin era. For recent

A. parvus collections I thank Kevin Bonham,

a malacologist with a good eye for tiny milli¬

pedes. Finally, 1 am very grateful to Mark

Harvey, Rowland Shelley and an anonymous

referee for constructive comments on a draft of

this paper.

References

Attems, C., 1926. Myriapoda. Pp. 1-402 in: Kiikenthal,

W. and Krumbach, T.. Handbuch der Zoologie.

Eine Nalitrgeschichle der Stdmme des Tierreiches.

4(1). Progoneata. Cbilopada. Insecta 1. Waller de

Gruyter & Co.: Berlin.

Attems, C., 1940. Dar Tierreich. Myriopoda. 3. Poly-

desmoidea. III. Fam. Polydesmidae. Vanhoeffeni-

idae, Cryptodesmidae, Oniscodesmidae. Sphaero-

trichopidae, Peridoiiludesmidae. Rhaehidesmidae.

Macellolophidae, Pandirodesmidae. Walter de

Gruyter & Co.: Berlin.

Black, D.G., 1997. Diversity and biogeography of Aus¬

tralian millipedes (Diplopoda). Memoir.^ of the

Museum of Victoria 56(2): 557-561.

Chamberlin, R.V., 1920. The Myriopoda of the

Australian region. Bulletin of the Museum of

Comparative Zoology 64( 1): 1-269.

HolTman, R.L., 1980 (1979). Classification of the

Diplopoda. Musdum d’Histoire Naturclle: Geneva.

Jeekcl. C.A.W., 1971 {\97()). Nomenclator generum et

famUianim Diplopodorum: A list of the genus

and family-group names in the Class Diplopoda

from the lOth edition of Linnaeus. 1758, to the

end of 1957. Monogrqfieen van de NederUmdse

Entornologische Vereniging, No. 5. Nederlandse

Enlomologischc Vereniging: Amsterdam.

Jeekel, C.A.W.. 1984. Millipedes from Australia, 7:

The identity of the genus Lis.wde.smus Chamberlin,

with the description of four new species from Tas¬

mania (Diplopoda, Polydesmida, Dalodesmidae).

Papers and Proceedings of the Royal Society of

Tasmania 118: 85-102.

Jeekel. C.A.W., 1985. Millipedes from Australia, 9: A

new polydesmoid millipede from Queensland

(Diplopoda, Polydesmida: Dalodesmidae),

Entornologische Berichten (Amsterdam) 45-

50-55.

Jeekel, C.A.W., 1986. Millipedes from Australia, 10:

Three interesting new species and a new genus

(Diplopoda: Sphaerotheriida, Spirobolida,

Polydesmida). Beaufortia 36(3): 35-50.

Marks, E.N.. 1991. Biographical history. Pp. 198-220

in CSIRO, The Insects of Australia. A textbook for

students and research workers. Volume 1. (Second

edition). Melbourne University Press: Melbourne.

Mesibov, R., 1996. Invertebrate bioregions in 7'asma-

nia. Report to Tasmanian REA Environment and

Heritage Technical Committee. Commonwealth of

Australia and the State of Tasmania: Hobart.

(www.rfa.gov.au/rfa/tas/raa/other/inverl/index.

html)

Silvestri, F., 1910. Descrizioni preliminari di novi

generi di Diplopodi. Zoologischer Anzeiger 35-

357-364.

Verhoefi; K.W., 1932. Dr. H.G. Broun s Klas.sen und

Ordnungen des Tier-reiebs wissenschaftlich

dargestellt in Wort und Bild. 5(2)(2). Klasse

Diplopoda. Akademische Verlagsgesellschalt:

Leipzig.

Verhoeff, K.W.. 1936. Die Sphaerotrichopidae der

sudlichen Maibkugcl und ihre Beziehungen.

Zoologischer Anzeiger 114: 1-14.

REDESCRIPTIONS OF TWO MILLIPEDES FROM TASMANIA (DIPLOPODA)

537

Figure I. Whole-animal views of mature male A. leae (left; QVM 23:8371) and A. parvus (right; QVM 23.8389).

Seale bars = 1 mm.

Figure 2. Ventral views of anterior segments of mature male A. leae (left; QVM 23:8322) and A. parvus (right;

QVM 23:8360). Scale bars = 0.5 mm.

538

R. MESIBOV

Figure 3. Outline drawings of left profile (left) and anterior half-section (right) of segment 8 of mature female

A. leae (QVM 23:8381).

Figure 4. Rear portion of midbody tergite of mature Figure 5. Ozopore on segment 9 of mature male

male A. leae (QVM 23:8371); anterior is to lower left. A. parvus (QVM 23:8389). Scale bar = 50 pm.

Scale bar = 150 pm.

REDESCRIPTIONS OF TWO MILLIPEDES FROM TASMANIA (DIPLOPODA)

539

Figure 6. Paranoia of segments 9-11 of mature female

A. leae (QVM 23:41535) showing paranoial tabs (t)

and ozopores (o). Scale bar = 0.5 mm.

Figure 7. Gonopod of A. leae. (A) mesal view of left

gonopod of holotype, redrawn from Silvestri (1910);

(B) mesal view of right gonopod telopodite of male

from northwest Tasmania (QVM 23:41717). Drawings

not to same scale. For posterior view, see Fig. 2.

Figure 8. Paranoia of segments 10-12 of mature female

A. pan'us (QVM 23:8389) showing paranoial tabs (t)

and ozopores (o). Scale bar = 0.5 mm.

Figure 9. Gonopod of A. parvus. Mesal and slightly

dorsal view of right gonopod telopodite of male from

southeast Tasmania (QVM 23:24743). For posterior

view, see fig. 2.

540

R. MESIBOV

100 km

Figure 10. Tasmanian localities of A. leae (triangles) and A. parvus (squares).

Memoirs of the Museum of Victoria 59(2): 541-553 (2002)

P7CA^0G07V(/M(PYCN0G0NIDA: PYCNOGONIDAE) FROM AUSTRALIA WITH

DESCRIPTIONS OF TWO NEW SPECIES

David A. Staples

Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E,

Melbourne, Vic., 3001 Australia

(e-mail: dstaples@museum.vic.gov.au)

Abstract

Staples. D.A.. 2002. Pycmgomim (Pycnogonida: Pycnogonidae) from Australia with

descriptions of two new species. A/ewo/r.v p/'A/irscKW K/e/on'a. 59(2):541-553

Australian representatives of the genus Pycnogonum Briinnich, 1764 are reviewed.

gonum carinatum sp. nov. and P. clarki sp. nov. are described and compared with their

congeners. Pvcnogonum occa Loman, 1908 is provisionally recorded from the con men a

slope of eastern Bass Strait, greatly extending its geographic range from tropical to temperate

Australia. The range of P. monlenbeeki Stock, 1992 is extended to northern Aus ra la. ype

specimens of A aurilinealiim Plynn. 1919, A /orre.vi Clark, 1963, A liiherculatuni ar ,

and the New Zealand species A. anovigerurn Clark, 1956 are re-examined. e ma

Pycnogonum aurilinealum and glands surrounding the mouth oP ^’cnogomm

described. The arrangement of gonopores in five species is reviewed. Gland-like coxa

ings were noted in several species. Abnormal development in two specimens ot A ciaiKi sp.

nov. is recorded. A key to the seven Australian species is provided.

Intraduction

The genus Pycnogonum Brunnich, 1764 consists

of over sixty species characterized by a robust

body, well developed proboscis and complete

absence of chelifores and palps. In the absence of

ovigers in some species, eggs are cemented

directly to the ventral surface of the trunk. Pyc-

nogontim are specialized feeders, most often

recorded in association with anemones on which

they feed. Records are predominantly from shal¬

low waters but are known from depths exceeding

2000 m. This paper reviews the Australian fauna

and reports on two additional species new to

science. Of the .seven species now' recorded, five

appear to be endemic to the Australian region.

T he arrangement of gonopores in males and

females was examined and pale spots (herein pro¬

visionally referred to as coxal glands) on the

coxae noted by Bouvier, 1913 and Flynn, 1919

were recorded in several species. Gland-like

structures surrounding the mouth were also

recorded..

Measurements of the coxae can vary signifi¬

cantly depending upon which surface they are

taken. For this reason measurements have been

taken on the lateral surface that presents the

reatest length and excludes the articular mem-

rane. Similarly, the proboscis is measured later-

llv e.xcluding the articular membrane which in

5 mc cases exceeds 20% of the proboscis length

hen extended. Trunk length is measured from

iterior margin of cephalothorax to base of

jdomen. Type material lodged m Australian

istitutions was re-examined. New type material

IS been lodged in Museum Victoria Abbrevia-

3 ns are; NMV, Museum Victoria Melbourne

M Australian Museum, Sydney; Mann

^s^aVch Group of Victoria; TM, Tasmanian

useum Hobart; NTM, Northern Territory

useum! Darwin. One millimetre scale bars refer

trunk size.

PYCNOGONIDAE Wilson

Pycnogonum Brunnich, 1764

•marks The genus Pycnogonum is divided into

ee subgenera based on the characteristics

mafr oviger (Stock, 1968): Pycnogonum

iger 8-9 segmented, terminal claw present,

moviger oviger 4-7 segmented, with or with-

Kmin.I claws and Wtoniscr, w.tk™

iners. Of the seven Australian species

541

542

D.A. STAPLES

recorded, three can now be assigned to

Nulloviger and two to Retroviger. These are P.

(Nulloviger) carinatum sp.nov., P. (Nulloviger)

tuberculatum, P. (Nulloviger) moolenheeki, P.

(Retroviger) clarki sp. nov. and P. (Retroviger)

aurilineatwn. Males of P. occa Loman, 1908 and

P. torresi have not been found and cannot be

assigned to a subgenus. Hooper (1980) recorded

P. rickettsi Schmidtt, 1934 from Arrawarra Head¬

land, northern New South Wales but re-examina¬

tion of his collection found that species not to be

present. The type locality for P. rickettsi is Mon¬

terey Bay, California. Until further material

comes to hand, this species is not included as an

Australian record.

2 .

6 .

Key to Australian species of Pycnogonum

Dorsomedian processes on trunk segments 1-3 pointed, height about 3 times

basal diameter....... p Loman, 1908

Dorsomedian processes on trunk segments 1-3 blunt or rounded, height

about equal to, or less than basal diameter. 2

Proboscis with dorsomedian tubercle. "3

Proboscis without dorsomedian tubercle. 4

Probo.scis greater than 40% length of trunk. Tibia 1 greater than 60% length

of femur. Propodus 3 times as long as wide. P. tuberculatum Clark, 1963

Proboscis less than 40% length ot trunk. Tibia I less than 60% length of

femur. Propodus about 2.5 times as long as wide.

;;.;. P. moolenbeeki Stock, 1992

Post-ocular tubercle present. p carinatum sp. nov.

Post-ocular tubercle absent. 5

Tibia 2 greater than 50% of tibia 1 . Abdomen rounded!!"”!”!”".”"!"!!””!..

.. . torresi Clark. 1963

Tibia 2 very short, less than 20% of tibia 1. Abdomen truncate. 6

Trunk conspicuously reticulate, unpigmented. Femur and propodus of about

equal length, propodal and tarsal spines loosely arranged in median row.

......... P. clarki sp. nov.

Trunk shagreened, typically dark with pale dorsomedian stripe. Femur

longer than propodus, tarsal and propodal spines arranged into dense clusters

. . aurilineatum Flynn, 1919

Pycnogonum (Nulloviger) carinatum sp. nov.

Figure lA-G

Material examined Molotype. Vic. Beware Reef, near

Cape Conran (37'’49.35'S, 147°47.39'E), 5-6 m,

SCUBA, T. O’Hara and A. Plummer, 15 Apr 1998, (stn

WV 11), NMV J48800 (male, ovigerous).

Other material. Vic. Southern Port Phillip Bay (38°

17.30 S, l44°41.40'E),7m, dredge, MRG, 25 Jul 1987

(stn SPPS 5), NMV J48801 (I female, gravid). Victory

Shoal, southern Port Phillip Bay (38°I7.0()'S to 38°

16.80 S, 144° 37.70' to 144° 38.30'E). 3-6 m,

SCUBA/dredge. MRG, 30 Mar 1986, (stn SPPS 3),

NMV J48802 (1 male, ovigerous. Trunk of specimen

lost). ‘The Wall’, Port Phillip Heads, in red algae

collection, 15 m. SCUBA, J.E. Watson. 9 Jun 1984,

NMV J48803 (1 female, gravid). Victory Shoal, Port

Phillip Bay. 6 m. dredge, MRG. 30 Mar 1986, (stn

SPPS 3), NMV J48804 (protonymphon. 3 pair of legs

developed). OfTShortland’s BlufI', Port Phillip Bay (38°

1 7.20 s to 38° I6.90 S to 144° 38.80 E to 144°

40.30'E), 5-15 m. dredge, MRG, 02 Nov 1986, (stn

SPPS 4), NMV J48805 (protonymphon, 3 pair of legs

developed). Cheviot Beach, Point Nepean (38° 18'S,

144° 40 E), Phyllospora/Ecklonia habitat, 3-3.5 m,

SCUBA, T. O’Hara and A. Plummer, 31 Mar 1998

(Stn WV5). NMV J48806 (2 females). OueensclilT

Port Phillip Bay. 7 m. SCUBA, D. Staples, 20 Mar

1992, NMV J48807 (1 male?). Cape Paterson, Twin

Reefs (38° 4(TS, 145° 39 E). intertidal, G.C.B. Poore

and R.S. Wilson. 5 Mar 1982. (stn CPA 20), NMV

J48808 (1 specimen, sex indeterminate). Port Phillip

Bay ( 38° 17.00 S to 38° 16.80’S, 144° 37.70'E to 144°

38°30 E), 3-6 m. SCUBA, MRG, 21 Jan 1989 (stn

SPPS 3). NMV J48809 (1 juv). Cape Paterson. 1 km E

of Harmers Haven (38° 34'S. 145° 40"), approx, 300 m

offshore, algal turf, 5-6 m. SCUBA, R.S, Wilson and C

Larson. 6 Mar 1982. (stn CPA 15), NMV 348810 (I

female). Tas. Arched 1., (43°26.32' S I47°20.I9 E), 6

m, J. Walls, (undated), NMV J48811 (I specimen, sex

indeterminate). WA, Breaksea L, SW comer, (35°

3.90 S 118° 2.50'E). red algae between Ecklonia, 15 m,

SCUBA, G.C.B. Poore and 11.M. Lew Ton, 7 Apr 1984

(Stn SWA 14), NMV J48815 (I male?).

Diagnosis. Trunk with post-ocular tubercle,

segments 1-3 with raised transverse ridges each

with blunt dorsomedian tubercle. Ovigers absent.

AUSTRALIAN SPECIES OF PYCNOGONUM

543

544

D.A. STAPLES

both sexes. Dorsal reticulations obscure or absent.

All leg segments tuberculate dorsally, propodus

longest segment, sole spines distally cleft,

auxiliary claws absent.

Description. Integument finely granular, reticul¬

ations clearly evident vcntrally, obscure, or not

visible in dorsal view, single post-ocular tubercle

slightly lower than ocular tubercle followed by

three raised transverse ridges on trunk segments

1-3, each with blunt dorsomedian tubercle, more

prominent in some specimens than others. Lateral

processes diverging, distally separated by about

own diameter, each with raised dorsodistal nodu¬

lous ridge most prominent middorsally and pos¬

teriorly-directed ventrodistal process, most pro¬

nounced on segments 1 -2. Ocular tubercle near

anterior margin of ccphalothorax. height less than

diameter, 4 eyes lightly pigmented. Proboscis

robust, slight constriction at about one-third

length, tapering distally, without dorsal tubercles.

Oral glands present. Articular membrane between

trunk segments and at base of proboscis wide,

capable of considerable extension and retraction,

rnembrane at base of proboscis folded or collar¬

like when contracted. Abdomen not quite reach¬

ing distal margin of coxa 2 of fourth pair of legs,

directed horizontally or slightly ventrad. distally

rounded in dorsal view, bearing nodulous dor¬

sodistal elevation, anus terminal. Ovigers absent

in both sexes.

Legs: Coxa 1 of all legs with pronounced ante-

riodistal process, dorsodistal margin developed

into rugose collar, more pronounced in some

specimens but usually absent on the first pair of

legs. Coxa 2 with distal rugose collar. Coxae 1

and 2 of about equal length, longer than coxa 3,

femur longer than tibia, tibia I longer than tibia 2;

propodus longest segment. Main claw about 60%

of propodal length. Femur with prominent ventro-

proximal swelling and 2 nodulous dorsodistal

lobes, prominent spine between lobes, few

smaller spines, some cleft, scattered on all seg¬

ments. All segments tuberculate dorsally, tuber¬

cles themselves minutely granular. Tarsal and

propodal sole spines cleft at apex, number widely

variable between legs of same specimen and

between individuals, consistently less abundant

on fourth pair of legs. Sole spines range from 10-

30, about 6 spines grouped at base of main claw;

tarsal spines 7-30. Auxiliary claws absent. Male

gonoporcs tiny, on ventral surface coxa 2 of

fourth pair of legs. Female gonoporcs well

defined, on dorsodistal surface of coxa 2 of fourth

pair of legs. Variably defined coxal glands are

present on all legs of both sexes.

Measurements (mm). Flolotypc. Trunk length,

1.75; trunk width (across second lateral pro¬

cesses), 1.03; proboscis length, 0.65; abdomen,

0.04; third leg. coxa 1,0.21; coxa 2, 0.20; coxa 3,

O. 16; femur, 0.42; tibia I, 0.35; tibia 2, 0.25; tar¬

sus (ventral), 0.17; propodus, 0.50; claw; 0.28.

Distribution. Central Victorian coast, SE Tasma¬

nia and S\V Western Australia; 3-15 m depth.

Etymolog}'. The specific name alludes to the

raised transverse ridges found on the dorsal sur¬

face of the trunk segments.

Remarks. The integument has the appearance of

being embedded with what look like air bubbles

or translucent beads which appear not to commu¬

nicate with the surface. This condition may be as

in P. crosnieri Stock, 1991. described as “granu¬

lated and pitted". Pycnogonum carinatum belongs

to a group of species lacking auxiliary claws and

ovigers in both se.xcs. It shares the presence of a

single postocular tubercle with P. portus.

Barnard. 1946, P. microps, Loman, 1904,

P. angtilirostrum. Stock, 1959, P. arhustunu

Stock, 1966b, P. forte, Flynn, 1928 and P. ttimn-

losum, Loman, 1908. The new species is closest

to P. microps from which it can be distinguished

by tuberculate and nodulate leg segments, dorso¬

median trunk tubercles and distally rounded

abdomen (truncate in P. microps). P. carinatum is

smaller than material of/^. anovigemm from New

Zealand, has more prominent trunk ridges and

ocular tubercle, possesses a prominent postcular

tubercle, and shorter abdomen with terminal anus.

The anus in P. anovigeriim is on the ventrodistal

surface of the abdomen.

Pycnogonum torresi Clark

Figure IH-I

Pvciiogomim torresi Clark, 1963: 76-77, fios 37

A-D.

Material examined. Ilolotypc. Queeihsland, Murray

Island, Torres Strait, 10-15 m. C. Medley and AR

McCulloch, 1907, AM PI3689 (1 female).

Remarks. The body is straw-coloured in pre¬

served material, without markings. Propodal sole

spines are blunt, split distally; the proboscis

slightly bulbous distally; and oral glands small

and inconspicuous. Reticulation of the trunk is

more evident than illustrated by Clark. This

specimen appears to have dried out at some stage

and is in a fragile condition with some damage.

Circular markings on the dorsal surface of coxa 2

of all legs appear to support Clark’s observation

that the specimen is a female, however small

AUSTRALIAN SPECIES OF PYCNOGONUM

545

perforations in the position normally occupied

by the male gonopores on the damaged last pair

of legs may indicate the specimen is male.

Additional material is required to clarify the

arrangement of gonopores in this species.

Distribution. Known only from type locality.

Pycnogonuni (Retroviger) clarki sp. nov.

Figure 2A-<j

Pycnogonum species.—Hooper, 1980: 475

Material examined. Ilolotype. New South Wales,

Arrawarra Headland (30°I7’S, 153° 15 E), 0-2.5 m,

J.N.A. Hooper 1976, NMV J48812 (male, with slide).

Paratypes. Collected with holotype, NMV J488I7 (1

female, 1 juvenile male).

Diagnosis. Integument strongly reticulate, trunk

segments 1-3 with well-defined, rounded pro¬

cesses. Oviger male only, 7-segmented. Propodus

approximately equal to length of femur. Tibia 2

shorter than wide. Propodal sole spines cleft at

apex, loosely arranged in narrow field along

length of propodal sole. Auxiliary claws absent.

Description. Entire integument (except for pro¬

podus and ovigers) strongly reticulate, straw-

coloured in preserved state, without markings.

Trunk fully segmented, first 3 trunk segments

armed dorsodistally with well defined, rounded

process each about equal in height to the ocular

tubercle. Lateral processes narrowly .separated at

base, short, wider than long, segment 4 with low,

granular dorsodistal boss. Ocular tubercle low,

height less than basal diameter, 4 large eyes

lightly pigmented. Proboscis robust, slightly

tapered and down-curved, slight constriction at

about half length, articular membrane at base of

proboscis wide when extended. Oral glands pre¬

sent. Abdomen wedge-shaped, reaching to distal

margin of coxa I on fourth pair of legs, sw'ollen in

mid region, truncate with low' dorsodistal tuber¬

cle, anus terminal. Oviger of male 7-segmented

with strong terminal claw; last segment with 2

small ventral spines and I dorsal spine, remaining

segments devoid of spines, obscure suture lines

.separate segments 1-3. Lengths (mm) of male

oviger joints are: scg.l, 0.03; seg. 2, 0.05; seg. 3,

0.14; seg. 4, 0.15; seg. 5, 0.08; seg. 6, 0.10; seg.

7, 0.12; claw, 0.09. Oviger absent in female.

Legs: Femur and tibia 1 subequal. Coxae 1-3

of about equal length, each w'ith distal coarse

granular collar, slightly raised on coxae 1-2,

articular membrane between segments wide.

Tibia 2 as in P. aurilineatum, shorter than wide

and separated from tarsus by a suture line.

Propodal sole armed with 30-40 spines each with

a cleft apex, loosely arranged in a narrow field

along length of propodus. Ventrally, coxa 3,

femur, tibiae 1 and 2 and tarsus support groups of

similar spines in lesser numbers. Main claw about

60% of propodus. Auxiliaries absent. Female

gonopores large, well defined, situated on dorso-

posterior surface of coxae 2 of fourth pair of legs.

Male gonopores tiny, ventral, on coxa 2 of last

pair of legs. Obscure middorsal co,xal glands

present on coxa 2 of all legs in both sexes. These

spots are free of reticulations found in the

surrounding integument.

Measurements (mm). Holotype: tnink length,

1.70; trunk width (across second lateral pro¬

cesses), 1.4; proboscis length, 1.0; abdomen 0.40;

third leg, coxa 1,0.30 ; coxa 2, 0.35; coxa 3, 0.30;

femur, 0.50; tibia 1, 0.52; tibia 2, 0.17; tarsus,

0.10; propodus, 0.53; claw 0.31.

Distribution. Known only from type locality.

Etymology. This species is named for Prof W.C.

Clark in recognition of having first examined

these specimens and for his significant contribu¬

tion to our knowledge of Australian representa¬

tives of this family.

Remarks. This species is close to P. aurilin¬

eatum but is significantly smaller in overall

dimensions and bulk. Total length is less than

one-third that of (a typical) P. aurilineatum. The

conspicuous reticulations, absence of pigmen¬

tation and body markings and relative length of

the propodus (approximately equal length to the

femur, about 70 %, in P. aurilineatum) further

serve as distinguishing characters. The number

and arrangement of propodal and tarsal spines,

the number of oviger spines, and the less promi¬

nent dorsal swelling of the femur arc differences

that could be age-dependent, but when considered

in combination may serve as additional points of

difference. Both the female and juvenile arc

abnonnal specimens. The juvenile has four legs

on the left side but only three legs on the right.

The dorsal surface of trunk segment 4 of the

female is incompletely developed and the mid¬

dorsal tubercle is atrophied and offset. The dorsal

tubercle on segment 3 is low, flat and wart-like,

possibly also an abnomiality. On the ventral sur¬

face, trunk segmentation is complete. Such mor¬

phological abnonnalities are not unique but nev¬

ertheless rarely recorded. In both instances there

is no evidence of trauma. Pycnogonum guyanae

Stock, 1975 also resembles this species but can be

distinguished by the longer tibiae 2 and placement

of the female gonopores on legs 3 and 4.

Figure. 2. Pycnogonum clarki sp. nov. NMV J488I2 (all figures except E, F, of male holotype). A, trunk, dorsal

view. B, trunk lateral view. C, third leg. D, oviger. E, ventral view segment 4. NMV J48817 F, juvenile male trunk,

dorsal view. G, female trunk, dorsal view.

AUSTRALIAN SPECIES OF PYCNOGONUM

547

Pycnof’oniim occa Loman

Figure 3A-D

Pycnogonum occa Loman. 1908: 35-36, pi. 12 figs

I7I-I74.—Bouvier, 1922:116.—Stock, 1966a: 401 (in

key).—Stock. 1968: 61-62. fig 22c-e.—Child, 1988:

27.—Stock. 1997: 389-409.

Material examined Victoria, S of Point Hicks eastern

Bass Strait ( 38°19.6'S to 38°19.00'S, 149°24.3’E to

149tx)27.3'E), 930-951 m. WHOI epibcnthic sled,

rock/rubble/clay/.sand and biogenic sediments, M.

Gomon et al. on ORV Franklin. 23 Jul 1986 (sin

SLOPE 33). NMV J48813 (1 juvenile).

Remcirh. All previous records are from tropical

waters. Type material was collected from the

Ceram Sea, Indonesia, at depths of 567 and 835 m

(Loman, 1908). Subsequent records are from the

Kemiadec Trench at 2470 m (Stock, 1968), east

of Luzon Island the Philippines (^Albatross Station

5447) at 567 m (Child, 1988) and New Caledonia

at 502-570 m (Stock, 1997). Slight morphologi¬

cal variations have resulted in the cautious assign¬

ment of all records to this species and prompted

Stock (1997: 407) to comment that this species is

“apparently of variable morphology.” This uncer¬

tainty has not been assisted by records of juvenile

specimens and the apparent absence of males.

Loman (1908) did not mention gonoporcs and his

identification of his ten specimens as female was

probably based solely on the absence of ovigers.

Should ovigers be absent in both sexes he prob¬

ably would not have recognised any males had

they been present. Stock’s (1968) Kermadec

Trench specimens dilTer from other material,

notably in the prc.sence of humps on the anterior

margin of the cephalothorax, the presence of

prominent laterodistal tubercles on the first coxae,

in the larger size, in the shape of the abdomen and

posse.ssion of a heavily sclerotized integument.

These specimens were also collected from much

deeper water. I agree with Child (1988: 27) that

the assignment of the Kemiadec specimens to

P. occa is questionable. The Bass Strait specimen

is distinguished by its small size (3.18 mm total

length) and the absence of a middorsal tubercle on

the fourth mink segment, a character described by

Loman (1908) as a knot-like process and used by

Stock (1966a) in his key to separate this species

from its congeners. It agrees most closely with the

specimens collected east ot Luzon 1. (Child,

1988:27) with which it shares the more slender

proboscis and “slight incipient reticulations” in

the integument. As Child made no mention of the

absence of a dorsal tubercle on the fourth trunk

segment in the Luzon 1. specimen, it is presumed

to be present. He noted however that the trunk

tubercles were more blunt and broad compared to

other material and attributed those differences to

the immaturity of his specimen. Stock (1997) did

not comment on the morphology of the New

Caledonian record and no comparison with that

specimen can be made. Gonopores and coxal

glands not evident, either undeveloped or

obscured by the fine reticulations. Glands not evi¬

dent on the small oral surface. Ovigers are absent.

This is yet another specimen provisionally

assigned to P. occa. Figures are provided to

enable future comparison with this specimen.

DistrihuHon. This record extends the distribution

from the tropical waters of the Ceram Sea,

Indonesia; east of Luzon I. the Philippines and

New Caledonia to the temperate waters of

southern Australia; 502-2470 m depth.

Pycnogonum (Nulloviger) liiherculatum Clark

Figure 3E-G

Pycnogonum tuberculatum Clark, 1963: 11-1^, figs

38 A-D.—Stock, 1994: 68.

Material examined. Vic.WSW of Gabo L, 130 m,

trawled. K. Moller on FV Dttrraween, Dec 1929, AM

PI3690, AM PI3691 (female holotype, 2 female

paratypes). NSW. 14 mi. otT Batemans Bay. 140 m,

trawled. K. Moller on FV Durraween. no date, AM

P13693 (I female). Tas. 1.2 km E of Cape Boulander,

(42°34.'S I48°6’. E), fine bryozoan/shell. 50 m, WHOI

epibenthic sled. R. Wilson on RV Challenger, 23 Apr

1985 (sin TAS 29), NMV J488I4 (1 male).

Remarks. The Tasmanian specimen is in general

agreement with Clark’s (1963) description and

falls within the recorded depth range. Ovigers are

absent, placing this species in the subgenus

Nulloviger. The most significant omission from

Clark’s description is the presence of two heavy

dorsodistal lobes and two dorsolateral digitifonn

processes on the femora. Clark’s fig. 38B also

incorrectly shows the presence of a segmentation

line between trunk segments 3 and 4. Spines on

the tarsus and ventrodistal surface of tibia 2 are

considerably more dense and abundant than illus¬

trated by Clark. The prominent dorsal tubercle on

the femur is flanked by two low dorsolateral

bosses. Coxae 1 and 2 w'ith low distal dorsolateral

bosses. Tiny male gonopores are present on the

ventral surface of coxa 2 of the fourth pair of legs

in all the material examined. Well defined, coxal

glands arc present on the dorsodistal surface of

the second coxae of all legs. Most glands accom¬

panied by a small spine on the proximal margin.

A few spinules are scattered around the tip of the

proboscis. Oral glands are not evident. Pycno-

gomitti luherculatiim, P. noditlosutn Dohrn, 1881

548

D.A. STAPLES

Figure. 3. Pycnogomim occa Loman, 1908, NMV J488I3 (juvenile). A, trunk, dorsal view. B, trunk, lateral view.

C, proboscis and trunk anterior. D, second leg.

Pycnogonum tuberculatum Clark, 1963. NMV J48814 (all figures of male). E, third leg; F, trunk, lateral view.

G, trunk, ventral view.

AUSTRALIAN SPECIES OF PYCNOGONUM

549

and P. moolenheeki can be distinguished from

their congeners by the basal articulation of the

abdomen, possession of a single median tubercle

on the proboscis and in the general shape of the

femur. P. noduhsiim differs most significantly in

the distinct segmentation of trunk segments 3 and

4. Pvcnogoiuim tuberculatum is in close agree¬

ment with Stock’s description of P. moolenheeki

with which it shares possession of blunt propodal

spines without evidence of bifurcation, the similar

distribution of leg spines and the fusion (or partial

fusion) of trunk segments 3 and 4. Pycnogonum

tuberculatum can be distinguished from

P. moolenheeki by its larger overall size, shorter

proboscis relative to trunk length, longer tibia 2

and more slender propodus. Dorsomedian trunk

tubercles appear to be more prominent in

P. tuberculatum though this may be variable. In

the general shape of the femur, P. lobipes Stock,

1991 also agrees with these species but otherwise

differs in several respects, notably the absence of

a median process on the proboscis, the distinct

segmentation of trunk segments 3 and 4 and the

non-articulated abdomen.

Distribution. Tasmania, Bass Strait, southern

New South Wales and north of Papua New

Guinea; 1-148 m depth.

Pycnogonum (ISulloviger) moolenheeki Stock

P. moolenheeki Stock, 1992: 95-97, fig. 11.

Materiai examined. Timor Sea, Ashmore Reef, Lagoon

Patch Reef, ncarNW entrance, coralline algae and dead

coral rubble, 5 m, B.C. Russell, 23 Feb 1981, NTM N9

(1 male).

Remarks. This specimen possesses a number of

characters intermediate between P. tuberculatum

and Stock’s description of P. moolenheeki. It

agrees with P. tuberculatum in possession ot

prominent dorsomedian tubercles on trunk seg¬

ments 1-3, complete fusion of trunk segments 3

and 4 and, although the dorsal tubercle in

P. moolenheeki appears to be generally less acute,

in the shape of the femur. However the smaller

overall size (trunk length 1.70 mm, measured

from anterior margin of cephalothorax to base of

abdomen), the proportionally longer probo.scis

(42% of trunk), the shorter tibia I (53% length of

femur) and the more robust propodus align this

specimen more closely with P. moolenheeki. 1 he

tropical water habitat and depth range 0-5 m are

also in keeping with the type locality of

P. moolenheeki. Tiny male gonopores pores are

present on the ventral surface of coxa 2 ot the

fourth pair of legs. Oral glands are present. Coxal

glands are present on all legs. This record repre¬

sents a significant extension of range from the

Gulf of Oman. The status of this material should

be confirmed by re-examination of the type

material and comparison with female specimens

from the Australian region.

Pycnogonum (Retroviger) aurilineatum Flynn

Figure 4A-F1

Pycnogonum auriiineatum Flynn, 1919b: 92-95,

plate XI11, figs. 1-2; plate XIV. fig. 3.—Stock, 1973:

125.

Material examined. Holotype. Tas. Port Arthur, Mr E.

Mawle? 1918?, TM J7/13083 (1 female).

Other material. Vic. Harmers Haven, intertidal,

MRG, 18 May 1983, NMV J12936 (1 male, oviger-

ous.). Shoreham, rocky shallows, M. O’Loughlin, 13

Jan 1981. NMV J12937 (1 female). Flinders, ocean

reef, M.P. Marrow, 29 May 1979, NMV J12939 (1

female). Peterborough, intertidal. C. Handreck, 22

Feb. 1984. NMV J12941 (1 female). Cat Bay, Phillip L,

G.C.B Poore (no date). NMV J12938 (I

female).Torquay, Point Danger, R. Bum, (no date).

NMV J12940 (1 female). Lake Tyers, Red Bluff,

intertidal, C. Handreck, 12 April 1984, NMV J488I6

(I male).

Distribution Investigator Group, Great Australian

Bight, South Australia; SE Tasmania to Coffs

Harbour, New South Wales; intertidal to 23 m

depth.

Remarks. This species was described from two

female specimens. Additional material expands

our knowledge of female morphology and enables

observations of male characteristics. Flynn accu¬

rately described the appearance of the integument

as shagreened (as in rough untanned leather) and

is comparable to the illustration ol' P. elephas

(Stock, 1966a; figs 4a, c ). The integument is gen¬

erally dark brown-black with a characteristic pale

middorsal band which runs the length of the first

three trunk segments and partially on to segment

4. This character is shared with P. calculum Bam-

ber, 1995 and persists in preserved material.

Characters that otherwise distinguish these two

species are well documented (Bamber, 1995).

Distal to an annular constriction around the

propodus, the integument is pale. Raised areas

with coarse granulation’s often have a ‘burnished’

appearance, being paler in comparison to the sur¬

rounding integument. In some (recently moulted)

specimens the colour of the integument is lighter

or pale pink (Staples, 1999). The pale dorsome¬

dian band usually persists at all stages. The artic¬

ular membrane at the base of proboscis is wide,

when extended it exceeds 25% of the total

550

D.A. STAPLES

Figure. 4. Pycnogonum auritineatum Flynn, 1919. NMV JI2936 A, male third leg. B, male oviger. NMV J12940

C, female mouth showing position of oral glands. D, female, coxa 2 dorsal, detail of coxal gland in relation to gono-

pore. TM J7/13083 E, female (holotype), distal segments leg 3. NMV J12937 F, female, ventral surfaeeof proboscis

and cephalothorax showing articular membrane. NMV J48816 G, male ventral, trunk segment. NMV J12938

H, female dorsal, trunk segment 4.

AUSTRALIAN SPECIES OF PYCNOGONUM

551

proboscis length measured ventrally. Oral glands

are well defined. Swellings on the legs are vari¬

able even within the same specimen, being more

acute on some legs than others. The dorsal surface

of all leg segments is finely tuberculate with

raised areas of the trunk and legs more coarsely

granular than the remaining integument. Tiny

robust spines are scattered over the trunk and

legs. Coxa I of all legs has a distal raised shoul¬

der divided middorsally. The femur, tibia I and

propodus are slightly constricted at one-third to

half their length and encircled by what appears to

be a broad reticulation line. Dorsodistally the

femur and tibiae terminate in a bilobed process

with one or two small spines originating between

the processes; bilobed processes on tibiae 2 are

less pronounced. Ventrodistally, tibia 2 supports a

group of about six spines. Small, cleft tarsal

spines are numerous, arranged in an irregular oval

pattern on an area of light-coloured cuticle. This

gives the appearance of the spines being placed

on a ‘pad’ which may be analogous to the “shin¬

ing patch" recorded in P. aiigtiliroslrum Stock,

1959. Stock also compared that species to P. auri-

Hneatum and illustrated (fig. 9d) what appears to

be a similar arrangement of spines. The propodal

sole is lined with a dense field of tiny spines

arranged in a group which is broad distally, grad¬

ually tapering anteriorly towards the tarsus.

Female gonopores are conspicuous, placed on the

dorsodistal surface of second coxa of the fourth

pair of legs and surrounded by raised integument.

Coxal glands arc present all legs. The surface of

the gland is smooth, generally shiny and in some

instances transparent.

The smaller male is in close agreement with

female holotype. Gonopores are small and incon¬

spicuous, placed on the ventral surface of coxa 2

of the fourth pair of legs. The oviger is 7-seg-

rnented, terminating in strong, curved claw. The

segmentation line between oviger segments 2 and

3 is not well defined. In the specimens examined,

spines on the distal four oviger segments vary in

number, those oti segment seven, I or 3; segment

six, I or 2; segment five, I and segment four, 1,2

or 3. Most spines have a cleft tip.

Lengths (mm) of male oviger joints are: seg. 1,

0.13; seg. 2, 0.13; seg. 3, 0.37; seg. 4, 0.36; seg.

5, 0.20; seg. 6, 0.26; seg. 7, 0.26; claw 0.27.

Specimens have been recorded in association

with solitary corals, corallimorpharians and

anemoties (Staples, 1997). This species is in close

agreement with Pyawgonum macitigascariensis

Bouvier, 1911, redescribed by Arnaud (1971).

Arnaud neither illustrated nor discussed the

arrangement of the propodal and tarsal spines in

detail, however her illustration of the tarsus and

propodus in lateral view appear to agree with

P. aurilineatiim. The most notable distinguishing

feature appears to be the strongly reticulate

integument of P. madagascariensis and the

absence of a dorsal process on the femur which is

present but variably developed in P. aurilweatiim.

In the absence of a record of a male P. madagas¬

cariensis no further comparison can be made.

Pycnogonum anrilinealum is the most frequently

recorded representative of the genus along the

southern Australian coastline.

Remarks

Structures on the oral surface and coxae 2 have

are tentatively recognized as glands however his¬

tological examination is required to positively

establish their function.

Oral glands. Six tear drop-shaped oral glands

are recorded in Australian species of Pycno¬

gonum. These unpigmented glands are placed

around the outer margin of the mouth, one pair at

the base of each Jaw. These glands may be incon¬

spicuous, particularly in small specimens, and are

probably present in all species. The same glands

are present but previously not recorded, in the

predominantly Northern Hemisphere species

P. litlorale (Strom, 1762), atid the Antarctic

species P. gordonae Pushkin, 1984. Associations

between the Pycnogonidae and anemones are well

documented, being found inside and outside their

host. These glands that may excrete a digestive

enzyme breaking down tissue that can be readily

sucked into the proboscis.

Co.xa! glands. In his description of P. aurilin-

eatum, Flynn (1919: 93) was of the opinion that

“yellowish spots” present on the dorsal surface of

coxae 2 were referrable to the pale markings

observ'ed on Pentapycnon charcoti Bouvier,

1913, which Bouvier thought to be glands. Type

specimens of A Itibercululum and P. moolenbeeki

have been described as females based on the pres¬

ence of gonopores on the dorsal surface of coxae

2 of all legs. However, re-examination of type

material of P. tuberculatum and a specimen of

P. moolenbeeki from northern Australia, shows

that while coxa 2 of all legs does possess a dorsal

gonopore-like spot, a male gonoporc is also pre¬

sent on the ventral surface of coxa 2 legs 4.

Either, these species are hermaphroditic, or, the

dorsal spots arc as Bouvier and Flynn thought,

glands. The spots are smooth and shiny and

smaller than the typical female gonopore.

552

D.A. STAPLES

Although in some specimens these spots are more

clearly defined, there appears to be no disruption

of the integument. For these reasons 1 believe the

spots more likely to be glands than gonopores but

whether they are a form of cement gland or serve

some other function has not been determined. As

with the gonopore, each gland is usually accom¬

panied by a tiny spine on its proximal margin.

Gonopores. In the genus Pycnogonum, gono¬

pores vary in number and placement. Gonopores

have been recorded on the dorsal and ventral sur¬

face of coxa 2 of all legs, dorsal and ventral sur¬

face of the fourth pair, ventral surface of legs 3

and 4, or the ventral surface of the third pair only.

The most common arrangement is for female

gonopores to be on the dorsodistal surface of coxa

2 of leg 4 and for the smaller, less conspicuous

male gonopores to be located ventrally. These

positions are consistent with the observed repro¬

ductive strategy of P. litlorale (King, 1973: 72)

and P. auhlinealtmt (Staples, 1999: 317) in which

the gonopores of both sexes are opposed during

the transfer of eggs. The placement of the male

gonopores on the ventral surface of coxa 2 of leg

4 appears to hold true for all Australian species

but due to absence of material and possible con¬

fusion with coxal glands, the number of female

gonopores is uncertain. Gonopores in P. aurilin-

ealiim, P. clarki and P. carinalum are confined to

legs 4; the male gonopore ventral and female dor¬

sal. The possible presence of gonopores on the

ventral surface of the legs 4 in the holotype of

P. torresi raises the possibility that this specimen

also shares these characters.

Should my conclusion be correct, C. motiil-

iferum Slock, 1991 from New Caledonia appears

to be the only remaining species described as hav¬

ing female gonopores on the middorsal surface of

coxa 2 of all legs. Re-examination of the type

material is required to verify the number of gono¬

pores in that species.

Acknowledgments

I am grateful to Dr Simon Pollard and Simon

Walker of the Canterbury Museum, Christchurch,

New Zealand, for loan of the type material of

P. anovigerum; to Dr John Hooper, Prof Rod

Simpson and Zoltan Enoch for tracking down the

elusive Arrawami Headland material; to Dr Liz

Turner of the Museum and Art Galleries, Hobart,

for loan of the holotype of P. aurilinealum. Dr

Penny Berents of the Australian Museum for loan

of the holotypes of P. torresi and P. tubercul¬

atum, and Dr Richard Willan of the Northern

Territory Museum for his assistance and making

available the specimen of P. moolenbeekie. I am

particularly grateful to Tim O’Hara, Dr Jeanette

Watson, Robert Bum, Mark O’Loughlin, Clarrie

Handreck. members of the Marine Research

Group of Victoria for making their material avail¬

able. Finally to Gary Poore, Genefor Walker-

Smith and Melissa Storey for their critical reading

of the manuscript and constructive comments.

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CONTENTS

New genus and species of southern Australian and Pacific Asterinidae (Echinodermata,

Asteroidea)

P. M. O 'Loughlin . .. .277

Report on selected species of BANZARE and ANARE Holothuroidea, with reviews of

Meseres Ludwig and Heterociwumis Panning (Echinodermata)

P.M. O'Loughlin .297

A new species of Neanthes (Polychaeta: Nereididae) from southern Australia

T. Bakken .

Corystolona, a new hydroid genus (Leptolida: Leptothecatae) from southern Australia

J.E. Watson .333

Hydroids (Cnidaria: Hydrozoa) from southern Queensland

J.E. Watson .337

A revision of the Australian genus Umbilia (Gastropoda: Cypraeidae)

T.A.Dairagh .355

The Australian species of Chimarra Stephens (Trichoptera: Philopotamidae)

D.I.Cartwright .393

Five artotrogids (Crustacea: Copepoda: Siphonostomatoida) from Eastern Antarctica

R. Johnsson and C.E.F. Rocha .439

New Phreatoicidea (Crustacea: Isopoda) from Grampians National Park, with revisions of

Synamphisopus and Phreatoicopsis

G.D.F. Wilson and S.J. Keable .457

Redescriptions oi Asphalidesmus leae Silvestri, 1910 and A. parvus (Chamberlin, 1920)

comb. nov. from Tasmania, Australia (Diplopoda: Polydesmida: Haplodesmidae)

R. Mesibov .531

Pycnogonum (Pycnogonida: Pycnogonidae) from Australia with descriptions of two new species

D.A.Staples .541