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CONTENTS

Systematics and biology of Macquarie Island echinoderms

T DU O'HARA” … reve eri onee den ved han EN EEEN

New species of the water mite genus Arrenurus from eastern Australia (Acari: Hydrachnidia:

Arrenuridae)

HSM ec pee atie eke: aaa Tue ace ws Was Sk aa les Geet eee ee cnc

A second species of Atriplectides Mosely from Australia (Trichoptera: Atriplectididae)

A: Neböiss ar iode Tare pecs enen bet een warden Lk Se EEE EERE

A new genus of subcortical coccoids (Hemiptera: Coccoidea: Eriococcidae) on Eucalyptus

PJ Gullan- ©, osn Oe PE ch ns dee See Lae ae Loe gu EE, I T e a yan

A new species of Ogyris Westwood (Lepidoptera: Lycaenidae) from southern inland Australia

RiFieldnus., Sateen REE Si. ban de aen de Bae ee ener hb Aon fee RE

Replacement names for five Australian species of Lasioglossum (Chilalictus) (Hymenoptera:

Halictidae)

KE Walker ech kad aar Eet le Maat ne | aie tg als et a ETENE TEE

Designation of type species for the genera of Australian paropsine beetles (Coleoptera:

Chrysomelidae)

PG. kelly and C-A. M: Reid on. rao Binet vn 9 en LON GEN ENE

Storthyngurella, new genus of Munnopsidae (Crustacea: Isopoda), with descriptions of three

new species from deep-sea basins of the Southern Hemisphere

M: VMalyuitind: Spera pae e ocean ae tatty OL, ot AE S E eT ATVETTE

Revision of Laphystiopsidae (Crustacea: Amphipoda): new and old species from South

China Sea, southeastern Australia, Falkland Islands and western Atlantic Ocean

J: Le Barnard „arvensis e ee p a oy adr EA RE

Description of a new species of the Pacific shrimp genus Paracrangon (Crustacea: Decapoda:

Crangonidae) from southern Australia, with a key to the genus

Y. Hanamura, V. Wadley and Js Taylor rara seen Lead a A ee eee

Memoirs of Museum Victoria 57(2): 167-223 (1998)

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS

Timotuy D. O'HARA

Museum Victoria, 71 Victoria Crescent, Abbotsford, Victoria 3067, Australia

and

Zoology Department, University of Melbourne, Parkville, Victoria 3052, Australia

(tohara@mov.vic.gov.au)

Abstract

O'Hara, T.D., 1999. Systematics and biology of Macquarie Island echinoderms. Memoirs of

Museum Victoria 57: 167-223.

Fifty-two echinoderm species are recorded from off Macquarie Island and the Macquarie

Ridge in the Southern Ocean. One new asteroid Odontohenricia anarea sp. nov. and one new

holothurian Trachythyone nelladana sp. nov. are described, The asteroid genus Calvasterias is

synonymised with Anasterias. The asteroids Cycethra macquariensis and Asterina hamiltoni

are synonymised with Asterina frigida and placed in the genus Cycethra. The asteroid Cera-

master lennoxkingi is synonymised with C. patagonicus, Solaster dianae with S. notophrynus,

and Anasterias sphoerulatus with A. mawsoni. The asteroids Psilaster charcoti, Odontaster

penicillatus, Ceramaster patagonicus, Crossaster multispinus, Solaster notophrynus,

Pteraster affinis, Henricia studeri, the ophiuroid Ophioplocus incipiens, and the holothurians

Paelopatides ovalis, Svnallactes challengeri, Laetmogone sp, Taeniogyrus sp are recorded

from the island for the first time. The following species previously recorded from Macquarie

Island have been re-identified: the asteroids Odontaster aucklandensis (=O. penicillatus),

Henricia aucklandiae (=H. studeri), Henricia lukinsi (=H. obesa), Smilasterias irregularis

(=S. clarkailsa), Anasterias antarctica (=A. directa), and the ophiuroid Ophiacantha pentag-

ona (= O. vilis). The existence at Macquarie Island of the species Hymenaster sp, Goniocidaris

umbraculum and Ocnus calcareus require confirmation. The asteroids Anasterias mawsoni,

Pteraster affinis, Porania antarctica and Odontaster meridionalis are reported from the shore

around Heard Island. The ecology and relationships of echinoderms from Macquarie Island are

discussed,

Introduction

Macquarie Island (54°29’S, 158°58’E) is in the

Southern Ocean midway between Tasmania and

the Victoria Quadrant of the Antarctic continent.

The island is small (34 km long and to 5.5 km

wide) and, together with small rocky outcrops to

the north and south, lies on the central section of

the Macquarie Ridge which runs south from New

Zealand. This narrow ridge formed of Miocene

basalt forms the southeast boundary of the

Australian and Pacific tectonic plates, and

Macquarie Island is a rare example of uplifted

oceanic crust. (Williamson, 1988; Duncan and

Varne, 1988; Selkirk et al., 1990). Recent

evidence (Adamson et al., 1996) suggests that the

ridge at Macquarie Island began its major phase

of uplift approximately 5 mya and emerged

around 700—600 kya. It has remained emerged for

the six glacial/interglacial cycles since that time

and experienced only periglacial rather than

glacial activity.

The general ecology of Macquarie Island is

determined to a great extent by its geographical

position. The climate is typical of an isolated

midlatitude island, and is characterised by strong

167

winds, frequent storms and high rainfall (Streten,

1988). The water temperature usually varies from

4-7°C. The Antarctic convergence lies only

40 km to the southeast, and cold water can

occasionally reach the island causing water

temperatures to drop as low as 2.8°C (Williams,

1988). Nevertheless the island is the most south-

ern ice-free shore in the eastern section of the

Southern Ocean.

The first echinoderms from Macquarie Island

to be reported in the scientific literature were

collected by Augustus Hamilton over 2 weeks

in March 1894. Hamilton, a biologist with the

Otago University in New Zealand, travelled on

the ketch Gratitude which was supplying a

penguin oil industry on the island. He collected

“starfish, echinoderms and holothurians . . . ”

from the eastern shore (Hamilton, 1895). The

holothurians were described as a new species,

Psolus macquariensis, by Dendy (1896) while he

was Professor of Biology at the University of

New Zealand. Specimens were sent to Europe

where they were examined by Ludwig (1898) and

Perrier (1905). Ludwig (1898) transferred the

species to a new genus Pseudopsolus. One

six-armed asteroid was identified as Stichaster

168 T. D. OHARA

North Head

JN Secluded Beach

Hasselborough Bay

N Handspike Point Garden Cove

Buckles Bay

Eagle Point

Nuggets Point

Bauer Bay

160 E

Sandy Bay

New

Zealand

Macquarie }

Island

Eastern

Antarctica

Aurora Point °

Green Gorge

Macquarie Island

Lusitania Bay

0 5 km

Caroline Cove

Hurd Point

Figure 1. Macquarie Island, showing place names mentioned in the text and the position of the island in the

Southern Ocean.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 169

suteri by Benham (1909), but later provisionally

referred to Anasterias mawsoni by Koehler

(1920). Mortensen (1925) recorded a specimen

of the New Zealand holothurian Cucumaria

(= Ocnus) brevidentis var. carnlevensis collected

by A. Hamilton but suspected that the locality

label was incorrect, possibly swapped with the

label in an equally aberrant lot of Pseudopsolus

macquariensis supposedly collected by Benham

from Stewart Island, New Zealand.

Harold Hamilton, son of A. Hamilton, spent

several years surveying the biology of Macquarie

Island as part of the Australasian Antarctic

Expedition (AAE) of 1911-1914. Echino-

derms were collected from various localities

along the coast. From this material, Koehler

(1920) described five new species of asteroid:

Parastichaster _ (=Anasterias) directus, P,

mawsoni, P. sphoerulatus, Asterina hamiltoni

(=Cvcethra frigida) and Cycethra macquariensis

(=Cycethra frigida), and recorded specimens of

Sporasterias (=Anasterias) antarctica (herein

referred to Anasterias directa). Mortensen (1921)

and Koehler (1926) recorded specimens of the

echinoid Notechinus (=Pseudechinus) novae-

zealandiae. Pawson (1968b) recorded the holo-

thurians: Pseudopsolus macquariensis, Trachy-

thyone macphersonae, Pseudocnus laevigatus

and an unidentifiable species of Trochodota

(herein provisionally referred to Taeniogyrus

dunedinensis). No crinoids (A.H. Clark, 1937) or

ophiuroids (Koehler, 1922b) were collected by

the AAE expedition from Macquarie Island.

The British, Australian and New Zealand

Antarctic Research Expedition (BANZARE)

collected from several stations off Macquarie

Island during 1-5 December 1930. Echinoderms

were collected at station 81, from the shore at

Buckles Bay, and dredged from station 83, off

Lusitania Bay in 69 m. A.M. Clark (1962)

recorded two asteroids from station 81: Anaste-

rias directa, A. sphoerulata; three from station

83: Henricia obesa, Hymenaster sp., Smilasterias

sp. ef. irregularis (=S. clarkailsa); and another

two previously unrecorded specimens of Ana-

sterias mawsoni and Cycethra macquariensis

collected by the AAE. Madsen (1967) recorded

four ophiuroids from station 83: Ophiacantha

pentagona (herein referred to O. vilis), Amphiura

magellanica, Ophiura meridionalis, and

phiopyren (=Ophioleuce) regulare. No crinoids

(John, 1939), echinoids (Mortensen, 1950) or

holothurians (M. O’ Loughlin, pers. comm.) appear

to have been collected from Macquarie Island.

The Australian National Antarctic Research

Expeditions (ANARE) founded a base on

Macquarie Island in March 1948. A steady stream

of biologists has visited or stayed at the base

and systematically or sporadically collected

specimens from the shoreline. Most collected

predominantly around North Head, from Hassel-

borough to Buckles Bay, One exception was

Wim Vestjens who surveyed many sections of the

shoreline during 1961-1962. Reports based on

this material have concentrated on the coastal

ecology (Law and Burstall, 1956; Kenny and

Haysom, 1962; Bennett, 1971; Simpson, 1976),

biogeography (Edgar, 1987), reproduction (Simp-

son, 1982) and pollution (Smith and Simpson,

1995). Pawson (1962) described a new species

Trachvthvone macphersonae from a specimen

collected by Hope Macpherson in 1959, and

Pawson (1968b), reporting on some specimens

collected by Isobel Bennett in 1967, recorded

Pseudoenus laevigatus from the Island for the

first time.

The New Zealand Oceanographic Institute

(NZOI) dredged or trawled 47 benthic stations

from Macquarie Island and the surrounding

ridge between 1959 and 1965. Pawson (1968a)

recorded Psolus antarcticus, Trochodota dunedi-

nensis and Pseudechinus novaezealandiae from

off Macquarie Island, and Pseudocnus leoni-

noides and Goniocidaris umbraculum from the

northern Macquarie Ridge. McKnight (1973b,

1977) described three crinoids from this material,

Ptilocrinus sp., Metacrinus wyvillii and Comat-

ulides (= Comissia) dawsoni. McKnight (1984)

finally provided an annotated checklist of the

echinoderms from the survey. This included five

crinoids, 14 asteroids, 11 ophiuroids, two

echinoids and two holothurians.

The USS Eltanin also trawled several stations

off Macquarie, as part of a large Southern Ocean

research program. This included stations during

cruise 16 (January 1965), cruise 27 (February

1967) and cruise 34 (June 1968). To date only

data on the comatulid crinoids have been pub-

lished; Speel and Dearborn (1983) listed three

species from off Macquarie Island. The cidaroid

echinoids have been the subject of an unpublished

PhD thesis (F.J. Fell, 1976). In addition, some

asteroids, echinoids and holothurians have

been identified in the Smithsonian Institution

(D. Pawson, pers. comm.), a few of which have

been examined for this report.

The Australian Museum sponsored a general

sublittoral survey of the Island during the summer

of 1977-1978 (Lowry et al., 1978). Forty-one

SCUBA dives were conducted at six different

areas to a depth of 20 m. Several intertidal sites

were also surveyed. This material has until now

remained unidentified and unpublished. Another

survey of the benthic and pelagic fauna was

170 T. D. OHARA

conducted by ANARE between 6 and 10

December 1986 (Williams, 1988). Fish were the

predominant target but many echinoderms were

collected in the process. A new species of aste-

roid, Smilasterias clarkailsa, was described from

this material (O'Loughlin and O’Hara, 1990).

Finally, material was gathered from shallow

water (0-3 m) by the author on a short visit to the

island in November 1989.

Examination of material from these surveys

indicated that the echinoderm fauna of Macquarie

Island was imperfectly known. There are now 52

species recorded from Macquarie Island and the

surrounding ridge, up from 44 known before this

report. Two species are new and 11 species are

recorded from the island for the first time. In addi-

tion, a revision of published material indicated

that there are nine species that have either been

erroneously identified or whose presence at

Macquarie Island must be placed in doubt. The

biogeography of Macquarie Island echinoderms

has been discussed by O'Hara (1998a).

The following abbreviations are used.

Morphology: d.d. - disc diameter (ophiuroids);

dia - diameter; I/w - length over width; R - radius

from centre of disc to arm tip (asteroids);

r - radius from center to margin of disc (asteroids),

ht - height; br - breadth (all measurements include

skin covering unless stated).

Institutions: Museum Victoria, Melbourne

(NMV); Australian Museum, Sydney (AM);

Tasmanian Museum and Art Gallery, Hobart

(TM); National Institute of Water and Atmos-

pheric Research (formerly New Zealand

Oceanographic Institute), Wellington (NIWA);

the Natural History Museum, London (BMNH);

National Museum of Natural History,

Smithsonian Institution, Washington (USNM);

Zoologisk Museum, University of Copenhagen

(ZMUC).

Material examined. The bulk of material exam-

ined for this report was collected from Macquarie

Island by the 1986 ANARE Macquarie Island

biological trawl program (lodged in the NMV),

the 1977-1978 Australian Museum Macquarie

Island Expedition (AM) and various shore collec-

tions (NMV, AM, TM). The stations and material

from the 1986 ANARE cruise is listed in Table 1.

The station list (MA stations) for the 1977—1978

AM expedition (Lowry et al., 1978) is too large to

reproduce here and the material has been included

in Table 2. Historical material examined includes

specimens from the AAE (AM, BMNH), BAN-

ZARE (SAM, BMNH), NZOI (NIWA) and

Eltanin (USNM) expeditions. This material was

compared with numerous species from surround-

ing regions including Australia (AM, NMV),

New Zealand (ZMUC, AM), Kerguelen

(BMNH), Marion Island (AM), Heard Island

(BMNH, NMV), South America (BMNH,

ZMUC) and Antarctica (BMNH, AM, SAM,

NMV). Comparative material is listed separately

South Australian Museum, Adelaide (SAM);

under each species below.

Phylum Echinodermata

Class Crinoidea

Key to Macquarie Island Crinoidea

l. Stem present .s.esisrisrististisririerssisisiestetssresiisresrter tetant idt in drndenttntentertrenresrrerřsit 2

— Stem absent ....sevseriessesasossrirodregvsaissneapaaean dinado a dreri honta Fes itett ra cnaiceoneesenoeess tanen 3

2. Stem without cirri, stem terminates in a basal disk … … … … Ptilocrinus sp.*

— Stem bearing cirri at regular intervals ane Metacrinus wyvillii?

3. Usually more than ten arms ……sorvansnvsersevenenenenvenenenenvenenenennevenrvenvenenvenndens 4

Ten arms only vicccsscssessecessesssesecsseeeserecesceenessesegepeeteeaseaesetsesacecnereareresanenstasbent 5

4, Oral pinnules flexible, usually with more than 30 segments. Arms rounded

or square with smooth sides at the arm base, 11-20 arms, Cirri stout, with

fewer than 25 segments … … nn verernenvenenesnenennenenvennn Glyptometra inaequalis®

— Oral pinnule prismatic, oral pinnules with less than 15 segments. Arms

laterally compressed, with spinous median keel near the base of the arm,

10-13 arms. Cirri long and slender, with more than 25 segments

rs RN TEA. Daidalometra arachnoides?

5. Auxiliaries and second brachials with prominent proximal lobes, which

incise the neighbouring ossicles … annen onenneneerenen Florometra austini

Auxiliaries and second brachials proximally truncate … sassen verveners 6

6. Oral pinnules with short segments, as long as wide, toothed distal segments

forming a rudimentary comb_… ann eneornersenenrversenenrd Comissia dawsoni®

Oral pinnules with long and slender segments, to three times as long as wide

tn EEE EEE ENT APEL Antedonid sp.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 171

Notes on key:

a McKnight (1973b: 204; 1984: 141) recorded ossicles of a stalked crinoid, possibly of the

genus Ptilocrinus, from the southern Macquarie Ridge (NZOI stn DS, 56"40.6'S, 158°45.5'E.

1280 m). McKnight noted that it was probably related to P. antarcticus Bather, a deep-water

Southern Ocean species.

> McKnight (1973b: 202; 1984: 141) recorded a single fragmented specimen of M. wyvillii

Carpenter, 1884 from the Northern Macquarie Ridge (NZOI stn D159, 49°O1'S, 164°30°E,

741 m). This species otherwise been found in deep-water Pacific localities.

penter, 1888) from off Macquarie Island and off the northern Macquarie Ridge (682-1693 m).

This species otherwise occurs throughout the Southern and South Pacific Oceans.

d Speel and Dearborn (1983: 22-24) recorded a single specimen of D. arachnoides (A.H.

Clark, 1909) to the northeast of Macquarie Island in 1647-1665 m. This species has been

otherwise recorded from the tropical western Pacific Ocean (22-118 m) and near Auckland

Island south of New Zealand (952-1336 m).

(McKnight, 1977) from the northern Macquarie Ridge (NZOI stn D18, 52°31'S, 160°3 1E,

128 m). This species was originally described as Comatulides dawsoni but transferred to

Comissia by Hoggett and Rowe (1986: 121). This comasterid is unusual in having a central

mouth and 10 arms. It has otherwise been found from three localities on the Campbell Plateau,

south of New Zealand (128-1280 m).

f Speel and Dearborn (1983: 52) recorded four specimens of “Unidentified Antedonid species

B” from two Eltanin stations off Macquarie and Auckland Islands (750-996 m). They note that

A.M. Clark, who examined the specimens, was uncertain of their affinities within the family

Antedonidae.

Table 1. Echinoderms from ANARE Macquarie Island benthic trawl stations, December 1986

BT1, off Lusitania Bay, 54°44.0'S, 158°51.4'E, 3541 m, 6 Dec 1986

No echinoderms,

BT2, off Lusitania Bay, 54°43.5’S, 158°53.1’E, 100-105 m, 6 Dec 1986

Odontaster penicillatus, Porania antarctica, Henricia studeri, Henricia obesa, Smilasterias

clarkailsa, Amphiura magellanica, Pseudechinus novaezealandiae, Psolus neozelanicus, Pseudocnus

laevigatus, Trachythyone macphersonae

BT3, off Nuggets Point, 54°33.4'S, 158°56.9'E, 108-135 m, 8 Dec 1986

Florometra austini, Odontaster penicillatus, Ceramaster patagonicus, Henricia studeri, Henricia

obesa, Smilasterias clarkailsa, Ophiacantha vilis, Amphiura magellanica, Ophiura meridionalis,

Pseudechinus novaezealandiae, Psolus neozelanicus, Trachythyone nelladana sp. nov., Pseudocnus

laevigatus, Trachythyone macphersonae

BT4, off Lusitania Bay, 54°45.0’S, 158°52.3'E, 65-90 m, 9 Dec 1986

Odontaster penicillatus, Porania antarctica, Henricia studeri, Henricia obesa, Amphiura magellan-

ica, Ophioleuce regulare, Pseudechinus novaezealandiae, Pseudocnus laevigatus, Trachythyone

macphersonae, Taeniogyrus dunedinensis

BTS, off Lusitania Bay, 54°44.2’S, 158°52.3'E, 57-70 m, 9 Dec 1986

Henricia studeri, Henricia obesa, Amphiura magellanica, Pseudocnus laevigatus

BT6, off Lusitania Bay, 54°43.4’S, 158°52.1°E, 25-29 m, 9 Dec 1986

Psilaster charcoti, Ophioplocus incipiens, Pseudechinus novaezealandiae

BT7, off Eagle Point, 54°31.4’S, 158°50.8'E, 150-200 m, 10 Dec 1986

Amphiura magellanica

BT8, off Handspike Point, 54°29.7'S 158°51,2'E, 330-450 m, 10 Dec 1986

Henricia studeri, Solaster notophrynus., Crossaster multispinus, Amphiura magellanica, Ophiura

(Ophiuroglypha) irrorata

BT10, off Judge and Clerk Rocks, 54°23.6’S, 158°59.3’E, 100 m, 10 Dec 1986

Odontaster penicillatus., Henricia studeri, Henricia obesa, Odontohenricia anarea sp. nov.,

Ophiacantha vilis, Pseudechinus novaezealandiae

T. D. OHARA

172

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SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 173

Order Comatulida

Antedonidae

Florometra austini A.M. Clark

Plate 3k

Florometra sp.—A.M. Clark, 1966: 704,

Florometra austini A.M. Clark in A.H. Clark and

A.M. Clark, 1967: 324, fig. 16.—Speel and Dearborn,

1983: 29, fig. 3, tbl. 6— McKnight, 1984: 141.

Material examined. ANARE 1986 Expedition, BT3,

NMV F60300(1).

Description of material. Centrodorsal hemispher-

ical, 1.8 mm dia, 1.0 mm high; approximately 70

cirri (mostly broken), cirrus sockets in alternating

rows, remaining cirri with 13-16 segments,

proximal 2 segments wider than long, middle

segments 2.5 times longer than wide, slightly

waisted, distal rim slightly everted, distal seg-

ments just longer than wide, terminal claw and

opposing spine well developed. 10 arms, all

broken; 5 radials, 1 Br, very short, auxiliaries cru-

ciform, proximal lobe (synarthial tubercle) over-

lying 1Br,, second brachial also with proximal

synarthial tubercle that overlies Bry, succeeding

brachials wedge shaped, with slightly everted

distal edges. First syzygy at 3+4, second at 9+10,

length between 1Br, to second syzygy 4 mm. P}

and P, very long, to 6 mm, approx 25 segments,

proximal segments longer than wide, distal

segments small, as long as wide, with low dorsal

tubercles, pinnule appearing comb-like. Ps, the

first genital pinnule, with 12 segments, 2 proxi-

mal segments small, as long as wide, next 7 seg-

ments enlarged and widened, 2-3 times as long as

wide, bearing the gonad, distal segments slender,

with spinous distal edges. Similar genital pinnules

present until P}. Disc naked, mouth central, anal

cone eccentric, high and slender.

Colour. Macquarie specimen (in alcohol) with

white arms and centrodorsal, brown disc and

gonads.

Habitat. A.M. Clark (1966) records this species

from a bryozoa/sponge substrate.

Distribution. Cook Strait, New Zealand (192-550

m); off Campbell and Antipodes Islands (58-210

m); Macquarie Island and the Macquarie Ridge

(108-135 m).

Remarks. The cirri and pinnules P, and P, on this

specimen have far fewer segments than was

reported for the type series. Possibly this can

be attributed to the larger size of the types

(centrodorsal = 4.0-5.5 mm dia). In this species

P, is usually similar to P}, long and flagellate

with many segments. Pz is usually the first

genital pinnule, however, P, or P4 can occasion-

ally be the first to bear a gonad. This species was

previously known from Macquarie Island only

from fragments.

Class Asteroidea

Key to Macquarie Island Asteroidea

(modified from A.M. Clark, 1962)

1. 2 series of tube feet in each arm furrow. Pedicellariae rare (except

Ceramaster) sorpren eestaas

Pe EN EE 2

4 series of tube feet in each arm furrow. Straight and crossed pedicellariae

COMMON annen reneeneernevenvenvernverversenrenevenveonenenvenndeennrenvennenneneennnenn: 15

2. Marginal arm plates large and conspicuous, forming a continuous rim

around the disc and arms … nnn neneneeneesnennerneenveervenvensernsensnnnvenven 3

— Marginal arm plates small, usually inconspicuous, often similar to abactinal

E ETET 6

3 Tube feet pointed, lacking suckers … nan srnn vennen Psilaster charcoti

— Tube feet with sucking disks … nnn enen vnrneenevensenenenennarnenenvenseneneenen 4

4, Stellate form, arms long and pointed enen Pseudarchaster sp.*

— Subpentagonal forma …nnansenronrvnrnnvenseneereneeneensenenneneenneneennensenveneennnennenvennenn 5

5 Large conspicuous recurved oral spine at the apex of each AWE Yo O ssnvenaens cue

Te it rood EN ORE Pa oped Odonaster pencillatus

Several blunt tipped oral spines around the jaw margin. Pedicellariae present

with large flat valves …

Pr B roars pee Ceramaster patagonicus

6. Disc covered in a thick skin and low sparse tubules. 5 short arms. Ventral

disc surface without spines, covered in radial striations ... Porania antarctica

— Not as above

wanensvesterdenentennensensneene

AA EA TE BT: 7

174 T. D. OHARA

7. Abactinal plates with a median pillar (pedicel) which bears crown of

Spinelets: (paxillae)s, u enaere a Rete e 8

— Abactinal plates flat or slightly convex, bearing spines or granules. No

Parle. OA ET S oen AEI R E EA tr len aaa treden de dean 12

8. Abactinal surface roofed over by a membrane supported by elongate

paxillar spines forming a chamber. 5 arms, with a conspicuous webbed

actinolateral fringe Of spines … … narren eneneenenenveneenenvenenenneneeesnenvenenn. 9

— Abactinal paxillae free of a membrane covered chamber. 7—11 arms ......... 1]

9, Adambulacral spines not webbed … annen Hymenaster sp.?

— Adambulacral spines webbed ...........cccccccecsssssssesesescscseesesestscserevevacseeceeecaees 10

10. Oral spines not webbed Pteraster stellifer?

— Oral spines Webbed … ananas eneen senen Pteraster affinis

11. Abactinal paxillae with short spines and arranged in regular oblique rows.

FALIVS aa TAT E ETE N NE AA TERT Solaster notophrynus

— Abactinal paxillae with long penicillate spines and arranged in an irregular

reticulum. 9-11 arms …n eeen. Crossaster multispinus

12. Subpentagonal form. 5 short triangular arms. Dorsal side convex, ventral

LNE ENEN MIO TTT TON Cycethra frigida

— Disk small, 5 long slender subcylindrical arms. 1-2 sabre-shaped

adambulacral spines deep in arm furrow … naren eeens eneen 13

13. Large recurved oral spine at each jaw tip ...... Odontohenricia anarea sp. nov

a Several small oral spines around the jaw margin … annae neen 14

14. Abactinal spinelets small and numerous, up to 30 per plate … en.

En betes Henricia studeri

— Abactinal spinelets stout and spaced, up to 6 per plate … … … Henricia obesa

15. 2 or more spines on each adambulacral plate … aaneen 16

— l spine on each adambulacral plate … nanne. 17

16. l longitudinal dorsolateral series of spines … … Sclerasterias mollis®

— Numerous dorsolateral spinelets … … … … … … … Smilasterias clarkailsa

17. Eee a E gonnen Ee Ede endet orn hert tg te Anasterias directa

— H ATS see en tard ieit inania iih a dir E aa a Anasterias mawsoni

Notes on key:

* McKnight (1984: 141) recorded a juvenile specimen of a Pseudarchaster species from off

Macquarie Island (NZOI stn C734, 53°55'S, 158°55'E, 360 m). McKnight noted its close

similarity to P. abernethyi H.B. Fell, 1958 from New Zealand. Another species P, discus

Sladen, 1889 occurs off South America and Marion Island.

b A.M. Clark (1962: 67) recorded a juvenile Hymenaster specimen from BANZARE station

69 off Lusitania Bay, and McKnight (1984: 143) recorded a specimen of Pteraster stellifer

Sladen, 1889 from Macquarie Island (NZOI stn C734, 53°55'S, 158°55’E, 360 m). Both these

specimens are discussed under Pteraster affinis.

ern Macquarie Ridge (NZOI stn D20, 49°39.8’S, 164°02.2’E, 126 m). S mollis is otherwise

known from the South Island of New Zealand and the Chatham Islands, 22-697 m.

Order Paxillosida

Astropectinidae

Psilaster charcoti (Koehler)

Piate la

Ripaster charcoti Koehler, 1906: 4-6, pl. III figs 20,

21, 31, 32.—Koehler, 1920: 258-259, pl. LI fig. 5, pl.

LIL fig. 1, pl. LXXII fig. 1.

Psilaster charcoti—Fisher, 1940: 93-94— A.M.

Clark, 1962: 13.—H.E.S. Clark, 1963: 30-31, text-fig.

4, pl. 3 figs 7-8. —A.M. Clark, 1989: 290.

Material examined. Macquarie Island, ANARE 1986

Expedition, BT6, NMV F60274(1); Antarctica,

65° 42'S, 92° 10°E, 110 m, 21 Jan 1914, AAE stn.7,

identified by Koehler (1920), AM(1); MacRobertson

Land, 66° 45'S, 62° 03'E, 219 m, BANZARE stn 107,

K1400(2).

Comparative material examined. Psilaster acuminatus

Sladen, 1889: Australia, eastern Bass Strait, 38° 23'S,

148° 46'E, 448-460 m, 5 May 1984, NMV F82984(1).

Description of Macquarie Island material. R = 50

mm, r= 11 mm; 5 arms, flattened, tapered evenly

to narrow point, 2 arms bent back over disc. Disc

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS

10 mm

Leh Om

Plate 1. a, Psilaster charcoti, dorsal view (lower arm slightly damaged on margin), NMV F60724: b, Odon-

taster penicillatus, dorsal view, NMV F60276; c, Cycethra frigida, dorsal view (disc slightly damaged on

upper right), NMV F76227; d, Ceramaster patagonicus, dorsal view, NMV F76237; e, Porania antarctica.

dorsal view, NMV F60278; f, Crossaster multispinus, dorsal view, NMV F60291; g, Solaster notophrynus,

dorsal view (skin removed from arm on left), NMV F60290; h, Solaster notophrynus, oral view, NMV

F60290.

176 T, D. OHARA

with central epiproctal cone and exposed

madreporite. Abactinal plates in close uniform

reticulum, lobed, with narrow bars connecting up

to 6 neighbouring plates. Plates paxillate with

high narrow pedicel, widened at tip, bearing 4-9

short spinelets. Papular areas small, cach with |

papula. Marginals prominent, lateral in position,

up to 50 along arm, protrudent with fascioles

between plates, superomarginals slightly smaller

than inferomarginals, plates covered by numerous

small thin spinelets, some capitate, some serrate,

with transverse row of large flattened pointed

spines in centre of plate, up to 4 on supero-

marginals extending to next plate, up to 4 larger

spines on inferomarginals, extending in length

over 2 plates, to 3.0 mm long. Up to 5 series of

actinals on interradial area of disc, only 1 persists

beyond fifth marginal plate, to at least three-

quarters R, plates small, convex, more numerous

than adambulacrals, paxillate with 3-10 small

conical subequal spinelets, often convergent.

Adambulacrals with 3-4 long angular pointed

furrow spines, central spine longest, to 2.0 mm,

up to 6 clustered subambulacral spines, proximal

pair of plates laterally compressed with no furrow

spines. Oral plates elongated, tumid, projecting

into mouth, with 2 rows of spines on either side of

the plate, furrow row with 10-12 thick blunt

spines, innermost 1—2 enlarged, up to 6 suboral

spines. Furrow narrow, 2 series of pointed tube

feet without suckers. Ventral surface covered in

gelatinous material that obscures spines.

Reproduction. This species develops lecitho-

trophic swimming non-feeding larvae; egg size

varies from 0.77 mm (Arthur Harbour, Scotia

Arc) to 0.95 mm dia (McMurdo Sound) (Pearse

and Bosch, 1994).

Colour. Macquarie Island specimens (in alcohol)

light tan. Falkland Island specimens coloured

pink dorsally, sometimes purple in the disc centre

and yellow ventrally (Fisher, 1940). Antarctic

specimens reddish brown or brown yellow

(Koehler, 1912).

Habitat. Usually mud, sometimes with pebbles,

tock (Fisher, 1940) or sponge/bryozoa (H.E.S.

Clark, 1963).

Distribution. Circumpolar Antarctic, South

Georgia, Bouvet Island (30-3248 m); Macquarie

Island (25-29 m).

Remarks. The Macquarie specimen closely

matches the descriptions and figures of Koehler

(1906, 1920) and H.E.S Clark (1963). I have

compared it to Antarctic specimens identified by

Koehler (1920) and A.M. Clark (1962). These are

similar although larger specimens (R > 90 mm)

differ in having more numerous spines, 8-12

abactinal spinelets, 5 superomarginal spines, 5

inferomarginal spines, and subequal oral spines.

H.E.S. Clark (1963) recorded paxillate pedicellar-

iae on her Ross Sea specimens. These are absent

from the current specimens.

A similar species, P. acuminatus Sladen, 1889,

is known from off Australia, New Zealand and

South Africa. An Australian specimen in Museum

Victoria was examined for comparison. It differs

in having larger marginal plates, fewer spines on

the inferomarginals, rare or absent supero-

marginal spines, and larger abactinal paxillae.

Proserpinaster neozelanicus (Mortensen, 1925),

another superficially similar species from New

Zealand, differs in having larger marginal plates

which extend onto the dorsal surface, 2 rows of

larger spines on inferomarginal plates and 2 series

of actinal plates persisting along the arm length.

Bathybiaster loripes Sladen, 1889, a wide ranging

Southern Ocean astropectinid, has smaller

enlarged spines on the marginal plates and a mix-

ture of large and small spinelets on the abactinal

paxillae.

P. charcoti is a new record for Macquarie

Island. It is has previously been recorded only in

Antarctic waters.

Order Valvatida

Odontasteridae

Odontaster penicillatus (Philippi)

Plate 1b

Goniodiscus penicillatus Philippi, 1870: 268.

Odontaster penicillatus.—F isher, 1940: 105—109.—

A.M. Clark and Downey, 1992: 156-157, figs 24i-j,

26n-p, pl. 37b [full synonymy}— A.M. Clark, 1993:

197,

Odontaster aucklandensis—McKnight, 1984: 141

[non O. aucklandensis McKnight, 1973c].

Material examined. Macquarie Island, off Buckles Bay,

372 m, Feb 1967, NMV F60275(1). ANARE 1986

Expedition, BT2, NMV F60277(1); BT3, NMV

F60276(2); BT4, NMV F76225(2); BT10, NMV

F60313(1).

Falkland Islands, Discovery stns WS73, WS93, WS824,

WS848, 115-130 m, identified by Fisher (1940),

BMNH 1948.3.16.237,264,274,277.

Argentina, Buenos Aires, 38°25'S, 56°30'W, identified

Bernasconi, BMNH, 1960.9.1.1(1).

Comparative specimens examined. O. meridionalis

(Smith, 1876): Heard Island, Camp Beach, washed up

after storm, NMV F76234(8); Counthian Beach, 7 Jun

1951, NMV F76235(4); S.W Beach, 1 Aug 1951, NMV

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 177

F76236(6); O. penicillatus grayi (Bell, 1881): Punta

Arenas, Sandy Point, HMS Alert, 17-19 m, BMNH

(holotype), Odontaster benhami Mortensen, 1925: New

Zealand, Oamaru, 37-55 m, 1960, AM J7067(4).

Description of Macquarie Island material. R = 16

to 50 mm, r to 28 mm. Dise flat, subpentagonal

with slightly concave margins. Abactinal disc

plates tabulate, with 4-18 spinelets, peripheral

spinelets slightly more pronounced. Spinelets on

largest specimen granule-like, truncate, as wide as

high, some with angular edges; spinelets on

smaller specimens terminating in small glassy

thorns. Up to 25 plates between margin and disc

centre. Papulae present between tabulae on disc

and radial areas. Large tumid marginal plates

fringe body margin, 35 plates from arm tip to arm

tip on largest specimen, 18-19 when R = 23 mm,

with up to 300 spinelets per plate, similar in form

to abactinal spinelets, slightly longer on infero-

marginal plates. Actinal plates convex, to 10

rows, with 5-12 spinelets; spinelets near mouth

long, bluntly pointed, 3-4 times as high as wide,

up to 1.1 mm high, orientated distally; marginal

spinelets shorter, more granular and less numer-

ous. No actinal papulae. Adambulacral plates

with 6—12 spines in 2 transverse rows, 2-3 spines

nearest furrow longest, up to 1.6 mm, cylindrical

or slightly flattened, blunt tips, spines decreasing

in length away from furrow. 1 (rarely 2) large

recurved oral spines at the jaw tip, orientated dis-

tally, with pointed glassy tip, on largest specimen

4 mm long, with slightly keeled ventral midline,

up to 2 mm long when R = 23 mm. 2 series of

suckered tube feet. No pedicellariae.

Colour. Macquarie Island specimens (in alcohol)

light tan. Live Atlantic specimens greyish,

orange, yellow or brown (A.M. Clark and

Downey, 1992).

Habitat. Sand, coarse shell and pebbles, mud

(A.M. Clark and Downey, 1992); rock (Branch et

al, 1993),

Distribution. Southern South America, Falkland

Islands, Burwood Bank (8-350 m); Marion Island

(527 m); Macquarie Island (55-372 m).

Remarks. As noted in the description, the largest

Macquarie Island specimen differs from the

smaller specimens in the form of the abactinal

spinelets. There is a similar range of variation in

the abactinal spinelets on specimens of O. peni-

cillatus | have examined from South America.

The number of spinelets can reach 50 per paxillae

(BMNH, 1948.3.16.277) and the shape can vary

from squat, capitate with a thorny apex to long,

tapering and serrated. Specimens of O. meridion-

alis (Smith, 1876) from Heard Island also vary in

the form of the abactinal spinelets, with only

some having glassy tips. Conversely, the presence

of pointed glassy tips on the abactinal spine-

lets has been used to distinguish the New

Zealand species O. benhumi Mortensen, 1925

from the subantarctic New Zealand species

O. aucklandensis McKnight, 1973e.

The Macquarie specimens are provisionally

referred to the polymorphic species O, penicilla-

tus. This species varies from stellate in form with

numerous small marginal plates to subpentagonal

with fewer larger marginal plates (Fisher, 1940;

A.M. Clark and Downey, 1992). Subpentagonal

forms from the Magellanic region have been

named O. penicillatus gravi (Bell, 1881). This

subspecies differs from the Macquarie specimens

in possessing pedicellariae (Mortensen, 1925).

McKnight (1984) referred his Macquarie Island

material to O. aucklandensiy but the present

specimens (from Macquarie Island and South

America) lack the distinctive glassy granules that

occur on the abactinal tabulae of the New Zealand

species O. aucklandensis and O. benhami.

Ganeriidae

Cycethra frigida (Koehler) comb, nov.

Plate le, Figure 2

Asterina frigida Koehler, 1917: 46-48, pl. 6 figs

9-11, pl. 7 fig. 8—Madsen, 1955: 13.—A.M. Clark,

1962: 33.—Guille, 1974: 34.—Cherbonnier and Guille,

1975: 615.—A.M. Clark, 1993: 209,

Asterina hamiltoni Koehler, 1920: 133-136, pl. 35

figs 5-7, 10, pl. 36 figs 1-3, pl. 66 fig. 5.—A.M. Clark,

1962: 24— Bennett, 1971, pl. 56 fig. 4.—Rowe and

Pawson, 1977: 343.—A.M. Clark, 1993: 210.

O'Hara, 1998a: 146. [new synonymy]

Cycethra macquariensis Koehler, 1920: 139-142, pl.

34 figs 1-4, 6, 7, pl. 66 figs 5a, b- A.M. Clark, 1962:

24-25.—Rowe and Pawson, 1977: 345, Simpson,

1982: 45 — MeKnight, 1984: 142.—A.M. Clark, 1993:

200. [new synonymy]

Material examined. Macquarie Island, no date, AM

J3520(holotype of A. hamiltoni), 10 Oct 1913, AM

J3605(holotype of C. macquariensis); AAE expedition,

BMNH, 1965.8.5.27(1); Secluded Beach, 27 Nov 1989,

NMV F76232(2); NMV F76233(3); Garden Cove, 21

Aug 1952, NMV F75925(1); 31 Mar, 1962, NMV

F45113(3); 12 Aug 1962, NMV F45112(1); no date,

NMV F45008(1); 25 Nov 1989, 0.5-Im, NMV

F76229(1); NMV F76230(2); NMV F76231(3); 26 Nov

1989, NMV F76227(5); Aerial Cove, 26 Jan 1950,

NMV F75924(1); 28 Nov 1989, NMV F76228(3). AM

1977-1978 Expedition, Gorilla Head Rock (9-12 m),

MA-142(2), MA-145(1), MA-149(3); Tottan Head,

Goat Bay (9-14 m), MA-39(1), MA-41(2), MA-42(1),

MA-371(5), MA-374(8), MA-376(3); Garden Cove

(6-14 m), MA-87(1), MA-92(1), MA-124(3),

175 T. D. OHARA

Supero-

marginals

Inferomarginals

Actinals

1mm

Figure 2. Cvcethra frigida, NMV F76231,

schematic diagram of an arm cross section showing

the position of the skeletal plates and the absence of

internal struts.

MA-125(2), MA-127(3), MA-128(3), MA-350(1),

MA-379(8); Green Gorge (14-18 m), MA-245(4),

MA-251(2); Caroline Cove (8-13 m), MA-300(1),

MA-306(3); Handspike Point (intertidal), MA-136(4);

Acrial Cove (3-6 m), MA-47(3), MA-85(3), MA-98(1),

MA-100(1), MA-107(13), MA-108(8), MA-109(7),

MA-1I10(2), MA-382(8), MA-386(6), MA-387(1),

MA-388(4), MA-389(2): Anchor Rock (20 m),

MA-89(2),

Kerguelen, NW end of Long Island, BANZARE

stn 56A, identified by A.M, Clark (1962), BMNH,

1965.8,5.48(2); Jeanne d'Arc, 20 m, BANZARE stn 5,

SAM K1420(1); BANZARE collection 784, littoral,

SAM KIRIB), Gazelle Basin, 25 28 Dec 1898,

Valdivia stn 160, identified by Madsen (1955),

ZMUC(1),

Description of Macquarie Island material, R to

20 mm, r to 9 mm, disc higher than arms, ventral,

dorsal surfaces mostly flat, often small sunken

areas On dorsal interradius; 5 arms (1 specimen

with 4 arms, NMV F76233), short but distinct,

dorsally convex, ventrally flattened , wide at base,

tapering blunt, rounded tip. Madreporite halfway

to interradial margin; anus sometimes visible

centrally; 2 gonopores sometimes visible on mid-

actinal area. Abactinal plates cruciform, stellar,

triangular or irregularly-shaped, with low convex

‘Y’-shaped or broadly-cresentic thickened region

bearing spinelets, rest of plate flat, thin, imbricat-

ing with 3-5 neighbouring plates; some small

round secondary plates in papular areas. Plates

regularly arranged. Papular areas small, no

larger than plate with 1, rarely 2, papulae.

Abactinal spinelets in round or oval cluster, 4-36,

usually 10-20 per plate. Spinelets short, blunt,

terminally spiniferous, l/w = 2-2.5 (up to 3 when

cleaned of skin, with constricted waist in middle),

Spinelet clusters obscuring underlying network of

plates.

Superomarginal plates larger than abactinals,

lateral or ventral position on disc, lateral on arms,

convex, 35 plates from arm tip to arm tip on R =

17 mm, proximal plates oval, 2-2.5 times as high

as wide, orientated transversely, plates placed

obliquely near arm base and on arm, plates on arm

1-1.5 times as high as wide. Inferomarginals

oval, convex, 1—1.5 times as high as wide, slightly

oblique on arms. Marginal spinelets as abactinals,

2 transverse rows on proximal plates with up to 7

spinelets in each row, 3 rows or cluster on distal

plates. Actinals block-like, rectangular, in an

imbricating lattice, up to 10 plates from margin to

jaw; distal plates shorter; forming longitudinal

rows near the furrow, distal plates irregular or in

irregular transverse rows; 2 of the longitudinal

rows persist onto the arms, one to tip. No actinal

papulae. Actinal spinelets longer than abactinals,

1-5, usually 2-3 per plate, cylindrical, thickened,

blunt tip or tapered, 2-3 times as high as wide (4

times as high as wide when skin covering bases is

removed), up to 0.4 mm high.

Adambulacral plates rectangular, sometimes

constricted in middle, well separated from each

other. Adambulacral spines longer than actinals,

to 0.8 mm long, club-shaped, 4 times as high as

wide; 3-4, per plate, positioned irregularly on

plate, only sometimes in transverse row, usually

only | on furrow margin; subequal or furrow

spine larger. Furrow spinelets near jaw sometimes

enlarged. No superambulacral plates, no internal

extensions of abactinal or actinal plates; proximal

interradial calcified pillar supports abactinal body

wall. 3 oral spines on oral margin of plate, larger

than adambulacral spines, 1-3 suboral spines. 2

series of suckered tube feet.

Reproduction. Separate sexes, egg size in gonad

variable, to 0.8 mm dia, appears to breed through-

out the year; possibly lays egg cases (Simpson,

1982).

Colour. Macquarie Island specimens (live)

usually pale olive green on the dorsal surface

(including superomarginals), sometimes light

grey, purple or brown, arm tips paler; ventral

surface (including inferomarginals) light tan, very

light orange or cream; tube feet darker tan or

orange; preserved material is tan with darker

tube feet. Kerguelen specimens are light brown

(Cherbonnier and Guille, 1975).

Habitat. In the immediate sublittoral (0.5-2 m) it

is found on rock boulders or gutters underneath

the kelp (Durvillaea) zone, In the sublittoral (to

20 m) found mostly on rocky substrates in associ-

ation with sponges, hydroids and red algae, or

amongst Macrocystis beds. McKnight (1984) has

reported a dredged specimen from 95 m.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 179

Distribution. Kerguelen (0-20 m), Macquarie

Island (0-95 m).

Remarks. Examination of a range of specimens

indicates that Cycethra macquariensis and

Asterina hamiltoni are synonyms of Asterina

frigida. Koehler (1920) did not compare his

three species. Both Macquarie Island species

were based on single specimens. The main

differences in the descriptions are the relative

size (smaller in C, frigida), density of the

abactinal skeleton, the length of the abactinal

spinelets and the number of actinal spinelets

per plate (A.M. Clark, 1962). However, the

minor variations in the numbers of actinal

spinelets, abactinal spinelets and papulae, the

density of the skeleton, and the colour do

not form any recognisable pattern in the

current range of specimens. Moreover, the

skeleton is flexible and consequently preserved

specimens can look superficially different

depending on whether the arms are flexed

slightly downward (dorsal skeleton appearing

open, actinal spinelets spread out) or upward

(dorsal skeleton appearing robust, actinal

spinelets densely clustered). The thickness of skin

around and on the spinelets can influence the

shape and height/width ratio of the spinelets.

Finally Cherbonnier and Guille (1975) have

reported specimens from Kerguelen up to R = 22

mm, similar in size to material from Macquarie

Island.

This species belongs in the Ganeriidae as

defined by A.M. Clark (1983), having a rounded

ventrolateral arm surface, small block-like

marginals, lacking internal arm structures (fig. 2)

and in lacking pedicellariae. A.M. Clark (1983)

suggested that Cycethra should be monotypic

given the extreme polymorphism of the type

species C. verrucosa (Philippi, 1857). However, I

can find no reliable character that would distin-

guish the two species at a generic level. On the

other hand C. frigida is clearly distinguished from

the related genera Ganeria and Perknaster by the

presence in C. frigida of clusters of small

spinelets on the abactinal, marginal and actinal

plates. Specimens of C. verrucosa can vary from

having abactinal plates with paxillae to low

cresentic mounds (metapaxilliform) and marginal

plates from small and inconspicuous to block-like

and distinctive (A.M. Clark, 1983). C. frigida is

similar to specimens of C. verrucosa with meta-

paxilliform abactinals and block-like marginals.

C. verrucosa can be distinguished from C. frigida

by the greater number (to 8) of adambulacral

spinelets.

Goniasteridae

Ceramaster patagonicus Sladen

Plate Id

Ceramaster patagonicus Sladen, 1889: 269, pl. 46

figs 3, 4, pl. 49 figs 3, 4.— Rowe and Gates, 1995:

65.—A.M. Clark, 1993: 249.— Branch et al., 1993: 44.

Ceramaster grenadensis patagonicus.—A.M. Clark

and Downey, 1992: 236-237, figs 39e, f, pl. 55e-f.

Ceramaster lennoxkingi McKnight, 1973a: 178—180,

fig. 4.—McKnight, 1984: 142. [new synonymy]

Material examined. Macquarie Island, ANARE 1986

Expedition, BT3, NMV F76237(1),

Description of Macquarie Island material. R=75

mm, r= 58 mm. Body flat, pentagonal: interradial

margins slightly concave; arm tips slightly

upturned. Abactinal plates tabulate, large and

hexagonal in centre of disc, smaller and quadran-

gular interradially: plates with approximately 12

marginal and 12-15 central granules (rarely to 24

marginal and 24 central spinelets), granules squat

and polygonal. 2-jawed spatulate pedicellariae on

some abactinal, marginal and actinal plates, larger

than granules, usually widely open. Marginals

conspicuous, 28 from arm tip to tip, covered in

granules except in centre of superomarginals

where 1-2 pedicellariae usually present. Actinal

plates as abactinals, larger ones with 30 close-set

granules. Adambulacral plates with 3—4 furrow

spines and to 7 subambulacral spines in 2 longitu-

dinal rows. Oral plates with 2 rows of 5-7 small

angular spines along margin; 5 large and 5-8

smaller suboral spines. 2 rows of suckered tube

feet.

Colour. Macquarie Island specimens (in alcohol)

tan. Live Atlantic specimens are reddish orange

above, cream below (A.M. Clark and Downey,

1992).

Habitat. Sand, mud (Sladen, 1889): rock (Branch

et al., 1993).

Distribution. North Pacific, Bering Sea, Gulf of

California, Burwood Bank Antarctica, Falkland

Islands (106-192 m); Marion Island (527 m);

Macquarie Island (105-148 m); the South Island

and subantarctic islands of New Zealand

(252-1125 m); southern Australia (no depth

recorded).

Remarks. McKnight (1973a) distinguished his

new species C. lennoxkingi from C. patagonicus

by the more numerous abactinal spinelets and the

fewer subambulacral spines. However, his speci-

mens (R = 30-51 mm) were considerably smaller

180 T. D. OHARA

than the holotype of C. patagonicus (R = 68 mm)

and the distinction between the two nominal

species appears to be size related. The present

specimen conforms broadly to both descriptions.

Branch et al. (1993) figured four large oral spines

on a specimen from Marion Island, which is

possibly a different species if this characteristic is

constant.

Poraniidae

Porania antarctica Smith

Plate le

Porania antarctica Smith, 1876: 108.—Fisher, 1940:

154.— A.M, Clark, 1962: 34.—Branch et al., 1993: 46.

A.M. Clark, 1993: 232,

Porania glaber Sladen, 1889: 360-362, pl. 59 figs 1, 2.

Porania antarctica antarctica—MeKnight: 1984:

142.

Material examined, Macquarie Island, AM 1977-1978

Expedition, Caroline Cove (13-18.3 m), MA-303(1),

MA-311(1); ANARE 1986 Expedition, BT2, NMV

F60278(1); BT4, NMV F76226(1). Heard Island, Camp

Beach 16 Aug 1951, washed up after storm, NM V(6):

SW Beach, Aug 1951, NMV F75926(1).

Description of Macquarie [sland material. R to

59 mm, r to 25 mm, height to 25 mm. 5 arms,

broadly triangular. Disc high, madreporite small

at top of interradial margin. Disc and arms cov-

ered in thick skin obscuring underlying skeleton.

Dorsal spines thick, tapered to blunt tip, up to 2.2

mm high, twice as high as wide, smaller distally.

Spine distribution on arms variable, NMV

F60278, R = 25 m, with 1 carinal and 1 lateral

longitudinal row of 6 spines; R = 50 mm (MA-

303), with carinal row of 15 spines and 2 lateral

rows of 10 spines; R = 59 mm (MA-311), with

sinuous carinal row of 30 spinelets and 4—8 lateral

spinelets not organised into longitudinal rows.

Some spines scattered on disc, occasionally 2

close together. Clusters of 1—10 papulae scattered

amongst dorsal spines. Marginal spines slightly

thinner than dorsal spines, slightly flattened,

sometimes truncate, 1, sometimes 2, spines per

plate, to 2.2 mm long. Ventral disc skin with

notable radial striations (fascioles), no papulae,

no spines. 2, distally 3, furrow spines, outer spine

largest, flattened and truncate, inner spines

smaller, subcylindrical, each orientated

obliquely to furrow. 2 oral spines at jaw apex,

2-3 small, pointed spines along furrow edge,

0-1 suboral spines. 2 series of tube feet. No

pedicellariae.

Reproduction. Macquarie Island specimens with

gonads attached to the interradial septa, genital

pore situated abactinally; gonad consisting of a

dense bunch of branching tubules, to 12 mm long,

containing abundant eggs, eggs to 0.5 mm dia.

Specimens of P. antarctica from McMurdo

Sound, Antarctica, have numerous eggs

(30,000-40,000), average size 0.55 mm dia,

which develop into pelagic feeding larvae (Bosch,

1989).

Colour. Macquarie Island specimens (preserved)

are off-white with brown tube feet. Branch et al.

(1993) reported their Marion Island specimens as

being a pink colour. Fisher (1940) recorded one

live Falkland Island specimen as scarlet.

Habitat. The two Australian Museum specimens

were collected amongst Codium or red algae

adjacent to dense Macrocystis beds growing on

boulders. Branch et al. (1993) recorded Marion

Island specimens from rock, sand and sand with >

5% mud. Sladen (1889) recorded a juvenile spec-

imen from the abyssal plain near the Crozet

Islands (2976 m).

Distribution. Antarctica (12-1335 m), Magellanic

region, South America north to 35°S (18-320 m),

Prince Edward Islands (10-280 m), Crozet (2976

m), Kerguelen (shore-23 m), Heard Islands

(shore—150 m); Macquarie Island (13—105 m).

Remarks, The present specimens have strongly

developed dorsal and marginal spines similar to

those found on specimens from Marion Island

(A.M. Clark, 1962) and Kerguelen (Guille, 1974).

Several subspecies have been proposed by Fisher

(1940) and A.M. Clark (1962), however, they are

not clearly distinguished either morphologically

or geographically. There is an undescribed

species recorded from McMurdo Sound in the

Ross Sea (A.M. Clark, 1962; H.E.S Clark, 1963;

Bosch, 1989). This species is distinguished by the

smaller body size (R to 30 mm), genital pores

located on the oral surface, the relatively few eggs

(100-310), and the development of lecithotrophic

demersal larvae. Other small paucispinous speci-

mens from eastern Antarctica may prove to be a

third species (A.M. Clark, 1962). The trinomial

P. antarctica glabra Sladen, 1889 has been used

for these animals (e.g., H.E.S. Clark, 1963:

McKnight, 1976; Mein, 1992) but this is inappro-

priate as the type of P. glabra is from Kerguelen

where it falls within the range of variation shown

by specimens of P. antarctica (A.M. Clark,

1962),

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 18]

Order Velatida

Solasteridae

Solaster notophrynus Downey

Plate !g-h

Solaster notophrynus Downey, 1971: 39-42, fig.

1.—A.M. Clark and Downey, 1992: 305, pl. 73e-f.

A.M. Clark, 1996: 193.

Solaster dianae Stampanato and Jangoux in Branch

et al., 1993: 45 [in key with figures] [new synonymy].

Solaster dianei—Stampanato and Jangoux, 1993:

183,

Material examined. Macquarie Island, ANARE 1986

Expedition, BT8, NMV F60290(1);

Northwest Tasmania, 41°06'S, 143°50’E,

m, NMV F82065(1).

Comparative material examined. Solaster subarcuatus

Sladen, 1889: Antarctica, Enderby Land, 65°48'S,

53°16'E, 209-180 m, 24 Jan 1930, BANZARE stn 41,

SAM K1453(6); Australia, off Port MacDonnell,

800-1000 m, AM J17796(1).

1110-1300

Description of Macquarie Island material. R = 40

mm, r= 11 mm. 7 arms, widened at base, taper-

ing rapidly after half R to blunt, round tip;

madreporite indistinct, slight interradial sulcus.

Abactinal plates in close regular reticulum,

usually quadrilobed, imbricating with neighbour-

ing plates. Paxillae small with low widened

pedicel, to 0.6 mm high, bearing 5-15, low cylin-

drical blunt, thorny, spinelets, to 0.15 mm long.

Abactinal paxillae 0.4-0.6 mm diameter, larger in

interradial disc sulcus, to 0.8 mm diameter,

smaller adjacent to marginals. Papular areas small

with usually 1 papula, papulae present on disk

and arms. Paxillae on disc and lateral arm sur-

faces in regular oblique rows, plates on dorsal arm

surfaces less regularly arranged, sometimes

trilobed with 2 papulae between plates.

Superomarginals similar to abactinals, slightly

larger than abactinals on smaller specimen, much

smaller than inferomarginals; paxillae with up to

12 spinelets. Proximal plates alternate in position

with inferomarginals, other plates aligned trans-

versely. Inferomarginals large, 31 plates along

arm, separated by less than their height. Paxillae

with high, laterally compressed pedicels, up to 30

short conical, terminally thorny, spinelets (mostly

broken) in 3—4 transverse rows.

actinals on ventral disc area, only 1 series persists

after 14th marginal (two-thirds R), paxillae small,

with 5-8 spinelets. No actinal papulae.

Adambulacrals narrow, widely separated, 43

along arm; with 3-5 long tapered webbed furrow

Australia, off

5 series of

spines in finger-like spread, to 1.5 mm long: 6-14

long thin subambulacral spines, largest near

furrow, to 1,1 mm, subequal with furrow spines,

not webbed, some thickened at tip. First adambu-

lacral plate on smaller specimen with 5 furrow

spines and 14 subambulacral spines in 3 trans-

verse rows. Next few plates with 4 furrow spines

and 1-2 rows of subambulacrals. Plates at half R

with 3 furrow and 8 subambulacral spines in a

single transverse row. Distal plates with 3 furrow

and 6 subambulacral spines.

Oral plates widened and tumid; 10 furrow

spines, webbed at their base, innermost 2-3

enlarged and pointed, to 2.0 mm, lateral spines

interleave with spines on opposing jaw, up to 25

suboral spines on raised section of plate, inner

few enlarged, terminally thorny, rest small, coni-

cal. Oral suture wide and bare. 2 series of tube

feet.

Reproduction. Macquarie Island specimen

female; gonads small, shaped like bunch of

grapes, attached to dorsal end of interradial arc:

numerous eggs, 0.2 mm dia.

Colour. Macquarie Island specimens (in alcohol)

white. Atlantic specimens dark-reddish brown

(Downey, 1971).

Habitat. Marion specimens recorded from rock

(Branch et al., 1993)

Distribution. Northwestern Atlantic (660-1230

m), Prince Edward and Marion Islands (335-475

m), Macquarie Island (330-450 m), Australia

(1100-1300 m).

Remarks. Solaster notophrvnus is characterised

by the numerous suboral and subambulacral

spines (pl. 1h), the small dense abactinal paxillae

arranged in regular transverse and oblique series,

and the shape and number of the arms (pl. 1g).

The specimen from Australia is larger (R=70 mm)

than the Macquarie specimen but differs only in

having slightly fewer subambulacral spines. A

similar species Solaster dianae, has been recently

recorded from Marion and Prince Edward Islands

(Branch et al., 1993). A full description of this

species has not been published, however one of

the authors (S. Stampanato) has kindly supplied

Ee with a description and photos of the holotype.

S. dianae is very similar in general appearance to

S. notophrynus, the only apparent difference

being that the superomarginal plates are distinct

in S. dianae but indistinguishable from the abacti-

nal plates on the holotype of S. notophrynus. This

182 T. D. O'HARA

feature is variable on the arms of the two speci-

mens | have examined and there appears to be no

reason to retain S. dianae as a separate species.

Solaster is predominantly a northern hemi-

sphere genus. The other exceptions are S. torula-

lus Sladen, 1889, known from off New Zealand

and the Kermadec Islands in 1042-1116 m;

S. longoi Stampanato and Jangoux, 1993 from the

Enderby quarter of Antarctica; S. subarcuatus

Sladen, 1889 from Kerguelen and southern

Australia; and S. regularis Sladen, 1889 from

throughout the Southern Ocean (although not yet

from Macquarie Island). These species have

fewer spines and spinelets, and with the exception

of S. torulatus larger paxillae and more arms

(8-10). Solaster differs from the closely related

Crossaster by the smaller, denser abactinal

paxillae, arranged into oblique rows.

Crossaster multispinus H.L. Clark

Plate If

Crossaster multispinus H.L. Clark, 1916: 66, pl. 18

figs 5, 6.—II.L. Clark, 1946: 150.—Rowe and Gates,

1995: 113.

Material examined. Macquarie Island, ANARE [986

Expedition, BT8, NMV F60291(1).

Australia, Tasmania, Bruny Island, 270-314 m, AM

E5078(2 syntypes); Eastern Bass Strait, 39°11.3'S,

148°41.2'E, 440-464 m, 6 Feb 1985, NMV F82985(1):

39°3.3'S, 148°38,0/E, 435-480) m, 6 Feb 1985, NMV

F82986(1).

Description of Macquarie Island material, R = 23

mm, r= 12 mm. 10 short arms, tapered evenly to

blunt tip. Abactinal plates in irregular open retic-

ulum, paxillate with thick pedicel and bristling

crown of 15 long sharp spinelets (mostly broken),

with 3—4 narrow connecting bars to neighbouring

plates. Large papular areas with 1-2 papulae,

smaller paxillae developing within papular areas

with 1—4 spinelets. Superomarginals similar to

abactinals; inferomarginals large, 14 along arm,

laterally compressed, widely spaced, separated in

middle of arm by more than their height, pedicel

large, thick, bearing to 30 long, serrated, spinelets

(to 1.3 mm long) in 3—4 transverse rows; some

smaller paxillae interspersed. Actinal disc area

with up to 3 rows of small sparse paxillae, not

extending onto arm. Adambulacrals with 5—6

long slender tapered, slightly serrated furrow

spines (mostly broken), webbed to third of their

height, to 7 subambulacral spines in transverse

row or arc, to 1.45 mm long, decreasing in height

away from furrow, slightly larger than furrow

spines, flattened, webbed but not joined by web to

furrow spines. Oral plates widened with up to 11

webbed furrow spines, innermost 4 enlarged, up

to 10 suboral spines on raised section of plate,

increasing in size towards apex.

Colour. Macquarie Island specimen (in alcohol)

white, tube feet brown. Live colour of an

Australian specimen (NMV F82985) dull yellow-

olive dorsally with paler arm tips.

Habitat. Unknown.

New

Island

Distribution. South eastern Australia,

Zealand (90-1152 m); Macquarie

(330-450 m).

Remarks. This specimen is the first record of

Crossaster from Macquarie Island. It closely

matches the description of H.L. Clark (1916). It

was compared with syntypes of C. multispinus

and several other specimens from southern

Australia in Museum Victoria. The only slight

differences are in the length of the inferomarginal

spinelets and the number of arms, which is almost

always 11 on Australian specimens,

H.B. Fell (1958) synonymised C. multispinus

with C. japonicus (Fisher, 1911) from Japan after

examining several specimens from New Zealand.

Rowe and Gates (1995) have disagreed and

retained the name C. multispinus for Australian

specimens. The identity of the New Zealand spec-

imens is unclear, but judging from the descrip-

tions and figures of H.B. Fell (1958) and H.E.S.

Clark (1970), they appear conspecific with the

Australian and Macquarie Island species. A simi-

lar species, C. penicillatus (Sladen, 1889), is

known from off South Africa, Marion Island,

Tristan da Cunha, and Gough Island from 55-800

m (A.M. Clark and Downey, 1992). It differs in

having slightly fewer spinelets, 10 on abactinal

plates, 15 on inferomarginals, 4—5 furrow spines,

and has 8-11 (usually 10) arms. C. campbellicus

McKnight, 1973d from off New Zealand has 10

arms but differs in having fewer spinelets, only

2-5 on abactinal plates.

Pterasteridae

Pteraster affinis Smith

Pteraster affinis Smith, 1876: 108.—Koehler, 1917:

48, pl. X fig. 11—A.M,. Clark and Downey, 1992:

326.—A.M. Clark, 1996: 206.

Pteraster affinis affinis—A.M. Clark, 1962: 63, text-

figs 10f, g.

? Hymenaster sp. (juv)—A.M. Clark, 1962: 67.

Material examined, Macquarie Island, AM 1977-1978

Expedition, Caroline Cove (8 m), MA-300 AM

J22731(1); Kerguelen, Royal Sound, 2-20 m, 10 Feb

1930, BANZARE stn 49, SAM K1461(1): Bras

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 183

Bolinder, 20-30 m, 14 Feb 1930 BANZARE stn 53,

SAM K1462(2); Heard Island. Camp Beach 16 Aug

1951, washed up after storm, NMV F76183(1).

Comparative material examined. Hymenaster sp.:

Macquarie Island, Off Lusitania Bay, 54°42.58’S,

158°54.5'E, 69 m, 5 Dec 1930, BANZARE stn 83.

SAM K1469(1).

| Description of Macquarie Island material. R = 13

| mm, r = 7 mm; 5 arms, short, triangular, blunt

| tip; interradial ares rounded. Abactinal plates

paxillate; 2-6, usually 4, subequal paxillar

spinelets, thin, slightly flattened, to 1.0 mm long,

slightly serrated or widened at tip, fenestrated sur-

face. Paxillar pedicel small, twice as high as wide.

Thick, pulpy supradorsal membrane connecting

spinelets at midheight. Actinolateral fringe of

spines connected by continuous webbing along

arm, spines flat, striated, truncate tip, to 2.0 mm

long. 4 webbed adambulacral spines in transverse

row near furrow. 5 webbed marginal spines on

each oral plate, webbing not continuous over jaw

apex; | large suboral spine, distally directed,

largely covered in membrane, tricarinate near

pointed tip. 2 rows of suckered tube feet.

| Reproduction. Macquarie Island specimen

female, gonad mass small (2 mm wide) consisting

of a bunch of short transparent tubules with 2-3

eggs; approximately 30 eggs per gonad: eggs

elongate, to 0.4 mm long. Gonad is attached

near the dorsal margin of the arm, near to the

interradial arc.

Colour, (In alcohol) tan.

Habitat. The Macquarie Island specimen was

found on boulders in a dense Macrocystis bed at

8 m. Branch et al. (1993) recorded their Marion

Island specimens from rock, sand and gravel.

Distribution. Antarctica (0-603 m), Magellanic

region (0-740 m), Kerguelen (0-91 m), Heard

Island (shore), Macquarie Island (8 m), Marion

Island.

Remarks. The condition of the specimen is poor;

many spines are broken and the animal is quite

flaccid. Nevertheless, enough diagnostic charac-

ters are visible to fully identify the specimen,

| including the large erect suboral spine, and the

| number and webbing of spinelets. A.M. Clark

| (1962) recognised three subspecies of P. affinis

that surround the Southern Ocean: P. affinis

lebruni Perrier, 1891 reported from Argentina, the

Falkland/Magellan region and the Prince Edward

Islands; P. affinis aculeatus Koehler, 1920 occur-

| Ting in eastern Antarctica from Kemp Land and

| the Ross Sea; and P. affinis affinis from Kergue-

| len. However, A.M. Clark and Downey (1992)

point out the overlap between these supposed sub-

species and the close relationship of P. affinis

with P. militaris from the Arctic. The Kerguelen

specimens that I have examined are very similar

to those from Macquarie and Heard Islands, dif-

fering only in having thicker opaque webbing

between the spinelets, but this may be due to

differences in preservation,

McKnight (1984) recorded a small specimen (R

= 17 mm) of P. stellifer stellifer Sladen, 1882

from Macquarie Island. Sladen (1889) described

this species as having five marginal oral spines

not connected by webbing, a smaller proximally

directed suboral spine, and regular “stellate”

abactinal paxillae with six subequal spinelets. 1

have not been able to examine McKnight’s spec-

imen but it appears from the description to differ

from the type in having up to 7-9 spinelets on the

abactinal paxillae, the peripheral spinelets (0.75

mm long) truncate, tapering slightly and the

central spinelet long (1.5 mm) stout and non-

tapering. More specimens are required to clarify

the relationship. A.M. Clark (1962) recognised

two subspecies of P. stellifer, P. stellifer hunteri

Koehler, 1920 differing from P. stellifer stellifer

in having flared paxillar spinelets. However, sub-

sequently A.M. Clark and Downey (1992) list

P. hunteri as a synonym of P. stellifer without

comment. P, stellifer has been recorded from

southern South America and around Antarctica

(79-2084 m), including the Balleny Islands

(McKnight, 1976 as P. stellifer hunteri)

A.M. Clark (1962) has also recorded a tiny

specimen (R = 3.5 mm) of a undetermined

Hymenaster species from Macquarie Island which

I have examined. The oral and adambulacral

spines lack webbing as is characteristic of

Hymenaster. The specimen has approximately 20

abactinal paxillae with 5-8 spinelets, the middle

1-3 longest, 2-3 subambulacral spines, two oral

and one small suboral spine. The specimen is

obviously a juvenile, possibly even a juvenile of

one of the Pteraster species listed above, and its

true affinities cannot yet be determined.

Order Spinulosida

Echinasteridae

Henricia obesa (Sladen)

Plate 2a

Cribella obesa Sladen, 1889: 544-45, pl. XCVI figs

3, 4, pl. XCVII figs 5, 6.

Henricia obesa.—Fisher, 1940; 164—-A.M. Clark,

1962: 48, figs Sn, 6a-c— McKnight, 1984: 143 —

Rowe and Albertson, 1987: 190-192, figs 2a, b— A.M.

Clark and Downey, 1992; 392-393, fig. 60s, pl, 95a.—

A.M. Clark, 1996: 237,

184 T. D. OHARA

Plate 2. a, Henricia obesa, dorsal view, NMV F60281; b, Henricia studeri, dorsal view, NMV F60287;

c, Odontohenricia anarea, paratype, dorsal view, NMV F60289; d, Odontohenricia anarea, adambul-

acral and marginal spinelets, NMV F60289; e, Odontohenricia anarea, abactinal spinelets, NMV F60289;

f, Anasterias directus, dorsal view, NMV F77766; g, Anasterias dir ectus, dorsal view of arm (skin removed

on upper arm), NMV F76241; h, Anasterias directus “antarctica form”, dorsal view of arm, NMV F76242;

i, Anasterias mawsoni, dorsal view, NMV F761 70; j, Anasterias mawsoni, dorsal view of arm, NMV F76164;

k, Anasterias mawsoni “sphoerulatus form”, dorsal view of arm, NMV F76181.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 185

Henricia lukinsi— McKnight, 1984: 143 [non H.

lukinsi (Farquhar, 1898)].

Henricia sp. aff. obesa — O'Hara, 1998a: 146.

Material examined, Macquarie Island, AM 1977-1978

Expedition, Caroline Cove (8-18 m), MA-298 AM

J22732(1), MA-300 AM J22730(1), MA-303 AM

J22733(1), MA-306(1). ANARE 1986 Expedition,

BT2, NMV F60281(13); BT3, NMV F60280(3); BT4

NMV F76223(7); BTS, NMV F60282(1); BT10, NMV

F60279(2); off Lusitania Bay, 54°42.5'S, 158°54.5’E,

69 m, 5 Dec 1930, BANZARE stn 83, SAM K1449(4)

and BMNH, 1965.8,5,108-110(4): 54°23.9'S,

158°59.3'E, 183 m, NZOI stn A698, identified by

McKnight (1984) as H. lukinsi, NIWA(I).

Falkland Islands, Port William, 51°40'S, 57°50’W, 22

m, 26 Jan 1876, sand/gravel, Challenger stn 315,

BMNH 90.5.7.831-3 (holotype and paratypes of H.

obesa); Discovery stns WS 81,84, 85, 86, 93, 220,

75-151 m, identified by Fisher (1940), BMNH,

1948.3.16.467 -470(4).

Australia, New South Wales, Shoalhaven Bight, 512 m,

identified by Rowe and Albertson (1987), AM

J13288(7); Victoria, East of Gabo Island, 410-450 m,

identified by Rowe and Albertson (1987), AM

J18628(3).

Comparative material examined. H. aucklandiae

Mortensen, 1925: Auckland Island, Carnley Harbour,

29 Nov 1914, ZMUC(syntype); Figure Eight Island, 2

Dec 1914, ZMUC(syntype); Masked Island, 3 Dec

1914, ZMUC(syntype); Chatham Islands, low tide,

identified by HB. Fell (1960), BMNH,

1952.4.16.17(2). H. lukinsi (Farquhar, 1898): Auckland

Island, Carnley Harbour, Figure Eight Island, 2 Dec

1914, identified by Mortensen (1925), ZMUC(1);

Auckland Island, Laurie Harbour, Discovery Expedi-

tion, 1904, identified by A.M. Clark (1962), BMNH

1905.7.14.10(1); Campbell Island, off East Cape. 18 m,

13 Jan 1980, AM J22905(1).

Description of Macquarie Island material. R =

9-46 mm, r to 10 mm; 5 equal subcylindrical or

flattened arms (1 specimen with 6 arms NMV

F60281), sometimes inflated at base, slender

distally, tapering to round tip, tip often curled.

Abactinal plates in coarse reticulum, papular

areas largest on base of arm, to 1.5 mm dia., with

3-9 papulae, secondary plates sometimes present

in papular areas, few papulae on disc, Abactinal

plates with irregular row of l-6 short stout,

tapered or slightly capitate, terminally thorny

spinelets, 1/w = 1.5-2 (3 when skin is removed),

to 0.4 mm high. Superomarginals quadrilobed or

thomboidal with up to 6 spinelets clustered on

dorso-distal side of plate. Inferomarginals

quadrilobed, imbricating, 6-8 spinelets in irregu-

lar transverse row. Usually 1 series (sometimes

2-3 series on specimens R > 40 mm) of bar-like

intermarginals extending to quarter R with 2-4

spinelets in transverse row. Actinals square-

shaped, extending to two-thirds R, with 24

spinelets, 3-10 bar-like plates in second series

proximally, to half R on largest specimens.

Marginal and some actinal papulae present. 10

adambulacrals to 7 inferomarginals, 1 slender

spine in furrow, 1—2 large club-shaped spines

on furrow edge, 3-5 subambulacral spines in 1

(smaller specimens) or 2 (larger specimens)

rows. Oral plates each with 3-5 furrow spines,

innermost enlarged, 1-3 suboral spines. Some

actinal papulae,

Reproduction. AM Expedition specimens with

gonads consisting of several tubules with 2-3

rows of short lateral branches or lumps, attenuat-

ing is size distally; sexes separate, female gonads

with eggs of varying size classes, 0.1—1.2 mm dia,

large eggs relatively few in number.

Colour, Macquarie Island specimens (in alcohol)

dark brown, tan or off-white, with yellow-brown

adambulacral area, brown tube feet. Fisher (1940)

recorded the live colour of Falkland Islands

specimens as orange-yellow, paler distally.

Habitat. The four AM Expedition specimens

from Caroline Cove were found amongst a dense

Macrocystis bed (8-18 m). McKnight’s (1984)

record of a Macquarie Island specimen from

“0 m” at Sandy Bay was presumably a beach-

washed specimen. Falkland Islands specimens

were recorded from coarse sand, fine sand,

stones, gravel, shell (Fisher, 1940). A Marion

Island specimen has been collected from rock

(Branch et al., 1993)

Distribution. South America, Falkland Islands

(22-450 m); Marion Island (527 m); Macquarie

Island (8433 m); southeastern Australia

(73-604 m).

Remarks. | have followed A.M. Clark (1962) and

McKnight (1984) in referring these specimens to

H. obesa. This species is characterised by an open

skeleton, robust abactinal spinelets arranged in

1-2 rows along the plates, and adambulacral

spinelets also arranged in 1—2 transverse rows.

However, there are some slight differences that

distinguish the Macquarie Island material from

specimens from South America and Australia.

These latter specimens (including the types) have

consistently smaller (0.2-0.3 mm) and more

numerous abactinal spinelets. Up to 15 spinelets

occur in 2 (or even 3) rows along the abactinal

plate. Larger animals have a more open skeleton

with more secondary calcification than Macquarie

specimens of the same size. Many authors,

including Fisher (1940) and Hayashi (1941),

have commented on the extreme difficulty in

186 T. D. OHARA

identifying Henricia material and I refrain from

establishing a new species on the basis of such

small differences.

McKnight (1984) has recorded two additional

Henricia species from Macquarie Island that are

otherwise known from the subantarctic islands of

New Zealand: H. aucklandiae Mortensen, 1925

and H. lukinsi (Farquhar, 1898). I have examined

a specimen identified by McKnight as H. lukinsi

and one as M. aucklandiae. The H. lukinsi speci-

men is identical to my H. obesa material. The

specimen of H. aucklandiae has numerous

spinelets and is referable to H. studeri (see

below).

H. lukinsi material from New Zealand can be

distinguished from Æ. obesa. H. lukinsi is a small

species (R to 24 mm) with adambulacral, actinal

and marginal spinelets aligned in a prominent row

on each transverse series of plates. Other notable

features include the denticulate nature of the

abactinal spinelets, and a shallow depression in

each interradial disc area. On the other hand New

Zealand specimens of H. aucklandiae are very

similar to H. obesa differing only in having a

slightly more compact skeleton and H. aucklan-

diae appears to be no more than a regional variant

of H. obesa. Large specimens of H. aucklandiae

(R=37 mm) have 7-9 adambulacral spinelets

(0.25 mm high) in two rows, 15-18 abactinal

spinelets in clusters or in two rows, a single row

of actinal plates extending almost to the ray tip

and some intermarginal plates at the base of the

arm. Molecular or developmental studies are

required to determine whether the difference

between all these regional forms are of specific or

infraspecific rank.

Henricia studeri Perrier

Plate 2b, Figure 3a—c

Henricia studeri Perrier, 1891: 102-103, pl. 9 fig.

2.—Fisher, 1940: 163-164, pl. 11 fig. 1—A.M. Clark

and Downey, 1992: 398, fig. 60n—A.M. Clark, 1996:

242.

Henricia aucklandiae.—McKnight, 1984; 143 [non

H. aucklandiae Mortensen, 1925].

Material examined. Macquarie Island, ANARE 1986

Expedition stn BT2, NMV F60286(2); NMV

F60287(1); BT3, NMV F60285(1); BT4 NMV

F76224(1); BT8, NMV F60287(1); BT10, NMV

F60283(1); off Buckles Bay, 372 m, Feb 1967, NMV

FO0284(1); 54°59,7'S, 158°364'E, 155-198 m,

NZOI stn E236a, identified by McKnight (1984) as

H. aucklandiae, NIWA(1).

Falkland Islands, Discovery stns WS 81, 85, 86, 872,

147-151 m, identified by Fisher (1940), BMNH

1948.3,16.453-465(12).

Comparative specimens examined. H. compacta

(Sladen, 1889): West of New Zealand, 38°50'S,

169°20'E, 510 m, 23 Jun 1874, Challenger stn 166,

BMNH 90.5.7.830(holotype); Australia, off Tasmania,

42°43'S, 148°25'E, 506 m, 25 Jun 1984, NMV

F76238(1). H. abyssalis (Perrier, 1894): South Africa,

off Cape of Good Hope, 304 m, BMNH, 1903.8.1.80(2).

Description of Macquarie Island material. R =

18-38 mm, r to 7 mm, arms 5, elongate, slender,

tapered, cylindrical, slightly inflated at base,

usually curled at tip. Disc small; anus central,

noticeable; madreporite small, at interradial

margin. Abactinal plates in irregular compact

reticulum, papular areas not large, to 0.45 mm

dia., with 2-3 papulae, little secondary calcifi-

cation, no accessory plates. Plates quadrilobed or

trilobed, imbricating with 2-4 neighbouring

plates, with small cresentic central raised section,

3-30 clustered spinelets per plate, to 5 wide,

spinelets 1/w = 4—6, with 2-4 minute thorns at the

tip, to 0.2 mm high. Marginals clearly visible in

regular longitudinal rows. Superomarginals

quadrilobed or irregular, imbricating, with thick-

ened distal edge, up to 30 spinelets, similar to

abactinal spinelets. Inferomarginals larger,

quadrilobed or rhombic, with thickened proximal

edge, up to 40 spinelets (to 0.14 mm long), few

intermarginal plates present at greatest arm

breadth (fifth R), similar to superomarginals. 1

series of actinals extend the arm length, closely

imbricating, distal edge thickened, 10-20

spinelets, in clusters or in 3 transverse rows on

proximal plates. | adambulacral plate to every

inferomarginal. 1, sometimes 2, small spines in

furrow; 2, sometimes 3, large stout capitate spines

at furrow edge (to 0.55 mm long), with 7-10 sub-

ambulacral spines, in 2-3 transverse rows, spine

height decreasing away from furrow, some

thorny. Marginal and actinal papulae present even

at arm base. Oral plates each with 3-4 furrow

spines, innermost enlarged (0.6 mm long), 2

suboral spines.

Colour. Macquarie Island material (in alcohol)

off-white. A.M. Clark and Downey (1992)

recorded live colour of Atlantic specimens as

orange-yellow to light red above, dull yellow

below. Marion Island specimens are pale (Branch

et al., 1993),

Habitat. Coarse sand, shell, stones (A.M. Clark

and Downey, 1992); rock (Branch et al., 1993).

Distribution. South America, Falkland Islands

(74-430 m), Prince Edward Islands (474-527 m),

Macquarie Island (100-450 m), Marion Island.

Remarks. Henricia studeri is related to several

other Henricia species that circle the Southern

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 187

„ abactinals

superomarginals

inferomarginals

actinals

: adambulacrals

1mm

Figure 3 a-c, Henricia studeri, NMV F60287: a, detail of a jaw showing the position of the oral spinelets;

b, detail of dorsal arm surface showing arrangement of abactinal spinelets; c, ventrolateral view of the base

of an arm, showing the arrangement of marginal plates. d, Henricia compacta, NMV F72238: ventrolateral

view of the base of an arm, showing the arrangement of marginal plates. e-g, Odontohenricia anarea,

holotype, NMV F60288: e, detail of a jaw showing the position of the oral spinelets; f‚ ventrolateral view of

a section of an arm; g, detail of dorsal arm surface showing arrangement of abactinal spinelets.

188 T. D. OHARA

Hemisphere including H. compacta (Sladen,

1889), H. abvssalis (Perrier, 1894) and

H. praestens (Sladen, 1889). These species were

characterised by A.M. Clark (1962) as having a

large numbers of spinelets, two furrow spines

and a extensive row of actinal plates, usually

extending to the arm tip. The Macquarie Island

specimens conform to the descriptions given by

Perrier (1891) and A.M. Clark and Downey

(1992) for H. studeri. They are identical to

Falkland Islands material identified by Fisher

(1940),

The Macquarie Island material was compared

to the juvenile holotype of H. compacta and to

larger specimens from off southeastern Australia,

H. compacta differs from H. studeri in having a

more compact abactinal skeleton, smaller, finer

abactinal spinelets, and more subambulacral

spines. The marginal plates on H. compacta are

quite pronounced and often irregularly placed,

particularly at the base of the arms (fig 3d), there

being fewer superomarginal than inferomarginal

plates. HM. praestens, known from off Marion and

Crozet Islands, has a secondary skeletal network,

fewer dorsolateral spinelets, knob-shaped furrow

spines and is an orange colour (Branch et al.,

1993). H. abyssalis which has been found in deep

water off South Africa is a much larger species

with two series of actinal plates that continue to

the arm tip.

Odontohenricia anarea sp. nov.

Plate 2c—e, Figure 3e-g

Odontohenricia sp. nov.— O'Hara, 1998a: 146.

Material examined, Holotype, off Judge and Clerk

Rocks, 54°23.6'S, 158°59.3’E, 100 m, 10 Dee 1986, stn

BT10, NMV F60288.

Paratypes, type locality and date, NMV F60289(2).

Other material. Macquarie Island, off Lusitania Bay,

54°42.5'S, 158°54.5"E, 69 m, 5 Dec 1930, BANZARE

stn 83, SAM K1826(1).

Comparative material examined. O. clarkae Rowe and

Albertson, 1988: South Africa, 34°33'S, 18°20'E, 290

m, BMNH 1987.4.9.1.1(holotype). O. endeavouri

Rowe and Albertson, 1988: eastern Bass Strait,

38°9.1°S, 149°54.0°E, 440 m, NMV F82983(1).

Description. Holotype. R = 26 mm, r=5 mm, br

= 6 mm. Arms 5, cylindrical, slightly broadened

at base, tapering gradually to rounded tip. Disc

small, madreporite near interradial margin, anus

central. Abactinal plates in reticulum, plates

cresentic or bar-like, imbricating with 2-3 neigh-

bouring plates, which surround sunken, round,

oval or irregular papular areas, papular areas

0.7-1.3 mm dia. Raised ridge on plate bearing

combs of 2-8 spinelets (typically 3—5), in 1, occa-

sionally 2, rows. Spinelets small, slender, slightly

capitate or tapered, minutely thorny (when skin

removed), 0.3-0.5 mm high and 0.1 mm wide.

Papular areas with 1-4 papulae around rim, often

with an accessory plate in centre. Marginals

distinct in regular transverse and longitudinal

rows, to 40 along arm. Superomarginals

quadrilobed, often irregular in shape, longer than

wide, raised section on plate triangular or bar-

like, with up to 10 spinelets in 2 transverse rows.

Intermarginals bar-like, single row extending up

to 14 plates along arm (half R), up to 4 spinelets

clustered on proximal plates. Inferomarginals

larger, quadrilobed or rhomboidal, with up to 16

spinelets in several rows, to 0.4 mm high.

Actinals square or rectangular, extend to half R,

with up to 7 spinelets in 1-2 rows. Marginal and

actinal papulae present, usually 1 per area.

Adambulacrals rectangular, 9 adambulacrals to

every 7 inferomarginals, | upwardly curved spine

inside furrow, 1-2 large, cylindrical blunt spines

on furrow edge, I/w = 3 (4 when cleaned), to 0.8

mm, 5-6 subambulacral spines decreasing in

height away from furrow, arranged in 2 transverse

rows. Oral plates with large recurved spine

apically, 3.0 mm long, 1.0 mm wide, tapering to a

sharp point; 3-4 furrow spines, half as high as

apical spine, 1-3 suboral spines.

Paratypes. R = 25 and 29 mm, r to 6 mm, br to

7 mm. Largest paratype differs from holotype in

having short second row of superomarginal plates

proximally and smaller oral spines.

Reproduction. Paratypes with short branched

gonads attached at base of arms to lower dorso-

lateral surface. No eggs apparent.

Colour. (In alcohol) fawn or light orange,

ventrally paler, brown tube feet.

Habitat. Unknown.

Distribution. Macquarie Island (69-100 m).

Etymology. Latinised form of the acronym

ANARE (Australian National Antarctic Research

Expeditions), in recognition of its contribution to

this research.

Remarks. The genus Odontohenricia Rowe and

Albertson, 1988 is distinguished from Henricia

and other echinasterids by the presence of a large

recurved pointed spine at the apex of each jaw.

Four species were previously known, all

deseribed by Rowe and Albertson (1988). O.

anarea is closest in form to OQ. clarkae, known

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 189

from off South Africa. The holotype of O. clarkae

was examined for comparison. It is in poor

condition with much of the skin and spines

removed, R= 33 mm. It differs from O. anarea in

having slightly thinner arms (br = 6 mm), a finer

abactinal skeleton (papular areas typically

0,6-1,0 mm dia.), more numerous but smaller

abactinal spinelets (0.25 mm long, 0.04 mm

wide), more numerous subambulacral spines

(10-13 per plate), and a larger oral spine (to 1.2

mm).

O. hayashii from Japan differs in having thin-

ner arms and a finer skeleton (papular areas

appear to be 1.0 mm dia. in Rowe and Albertson,

1988, fig. 10), but otherwise has quite similar

abactinal spinelets and marginal and adambu-

lacral spines. O. endeavouri from off southern

Australia has a finer skeleton consisting of stellate

plates and isolated tufts of numerous small

spinelets. O. fisheri from off the North Pacific

coast of America has more numerous abactinal,

adambulacral and oral spines.

Specimens of Odontohenricia have been

misidentified as Henricia species in the past.

They are often superficially similar and, as more

specimens are collected, show the same minor

variation in skeletal and spine arrangement that

make Henricia specimens difficult to identify.

One of the current specimens was amongst nine

identified as Henricia obesa by A.M. Clark

(1962). Apart from the lack of an enlarged oral

spine, the Macquarie Island specimens of

H. obesa differ in having a more open skeleton

with fewer spinelets and spines.

Order Forcipulatida

Asteriidae

Smilasterias clarkailsa O’ Loughlin and O'Hara

Smilasterias sp. cf. irregularis —A.M. Clark, 1962:

87-88.

Smilasterias irregularis— MeKnight.

[non S. irregularis H.L. Clark, 1928].

Smilasterias clarkailsa O'Loughlin and O'Hara,

1990: 316, pl. If, g.

Material examined. Macquarie Island, off Lusitania

Bay, 54°42.7'S, 158°54.50E, 69 m, 5 Dec 1930, BAN-

ZARE stn 83, BMNH 1965.8.5.220 (paratype);

54°23.9'S, 158°59.3'E, 183 m, NZOI stn A698, identi-

fied by McKnight (1984) as S. irregularis, NIWA(I).

ANARE 1986 Expedition, BT2, NMV F53754 (holo-

type), NMV F53755(paratype); BT3, NMV

F53753(paratype). AM 1977-1978 Expedition, Garden

Cove (11-14 m), MA-379(1); Caroline Cove (13 m),

MA-311(1).

1984: 143

Comparative specimens examined. See O’ Loughlin and

O'Hara (1990),

Description of material. R = 15-75 mm, r to 13

mm; 5 subcylindrical arms, sometimes shed at

base; dise small, disc skeleton reticulate. Abac-

tinal spinelets spaced in small specimens ( R < 40

mm) or grouped in large specimens, cylindrical or

capitate, I/w = 2-2.5, Carinal plates raised,

quadrilobed, bearing 8 spinelets in a ‘V’-shape.

To 10 small, narrow dorsolateral plates

transversely between carinals and supero-

marginals, 1-3 longitudinal linkages, up to 6

spinelets per plate, up to 20 spinelets between

carinals and superomarginals. Superomarginals

cruciform, proximal plates sometimes beaded, to

8 spinelets in 1-2 transverse rows, lower spinelets

club-shaped. Inferomarginals usually forming

actinolateral margin to arm, 3-4. sometimes 5,

flattened, flaring, truncate spines. 2 actinal series

on larger specimens, first to three/quarters R,

second to quarter R, with 0-3 spines. Adambu-

lacrals with 3, sometimes 4, spines, furrow spine

rectangular and chisel-shaped, spines away from

furrow truncate and flaring,. Two pairs of oral

spines, adoral spines larger than adambulacral

spines, Dense crossed pedicellariae on abactinal,

marginal surfaces, to 3 times as numerous as

spinelets. Small lanceolate straight pedicellariae

abactinally, marginally, actinally and in furrow.

Large incipiently felipedal pedicellariae mar-

ginally, actinally and in furrow, each valve with

small notch on either side of the hooked tip,

largest on disc interradial area. 2 series of large

suckered tube feet.

Reproduction, Gonads long (to half R) with short

lateral branches or tubercules arranged in 2-3

longitudinal rows, attached at the base of arm on

the lower dorsolateral surface: sexes separate,

eggs elongate, 0.5—1.0 mm long.

Colour. Tan (in alcohol).

Habitat. The AM Expedition specimens were

found on Codium adjacent to a dense Macrocystis

bed, and on a sponge/hydroid mat under a rock

overhang.

Distribution. Macquarie Island (11-357 m);

South Tasman Rise (620-1650 m).

Remarks. The two large specimens collected by

divers on the 1977-1978 Australian Museum

Expedition (MA-311, R = 70 mm; MA-379, R =

75 mm) differ from the S. clarkailsa types (R to

38 mm) in having grouped abactinal spinelets,

more numerous spines and spinelets, a second

row of actinal plates and no beading on the super-

190 T. D. CHARA

omarginal plates. These features indicate that

S. clarkailsa is more closely related to the

subantarctic species S. scalprifera (Sladen, 1889)

than to the Australian species S. irregularis H.L.

Clark, 1928, S. clarkailsa is distinguishable from

S. scalprifera by the density of crossed pedicel-

lariae. This species has been recently collected

from seamounts south of Tasmania (O’Hara,

1998b).

Anasterias Perrier

Anasterias Perrier, 1875: §1,—Fisher, 1930: 221.—

Fisher, 1940; 231.—A.M. Clark, 1962: 93,

Sporasterias Perrier, 1894: 107 — Fisher, 1930: 2

Calvasterias Perrier, 1875: 84.—Fisher, 1930: 2

[New synonymy]

Remarks. Calvasterias was distinguished from

Anasterias by the increased calcification of the

abactinal skeleton. However, this feature varies

within and between different populations of

species in both genera. Calvas/erias includes four

species: C. asterinoides Perrier, 1875 the type

species, supposedly from the Falkland Islands,

C. stolidota Sladen, 1889 from South America,

C. laevigatus (Hutton, 1879b) and C. suteri de

Loriol, 1894 from New Zealand. | have examined

adult specimens from all four species (see under

A. directa below) and find no significant differ-

ence between the skeletons of the last three

species and the skeleton of pentamerous

Anasterias specimens from South America

(A. antarctica), Kerguelen (A. rupicola) and

Macquarie Island (A. directa). The status of the

type species is uncertain, as it is unclear as to

whether the types were collected from Torres

Strait, the Falkland Islands or the Auckland

Islands, south of New Zealand (A.M. Clark, 1962:

93-4). The only other known specimens are also

of uncertain origin (A.M. Clark, 1962). These

specimens (BMNH — 1844.5.29) are small

(R=17-23 mm) with coarse overlapping skeleton

ossicles, reduced papular areas and few carinal or

dorsolateral spines. They are similar to some

small spinous specimens of C. /aevigatus and are

possibly members of that species. The overlap-

ping skeleton is probably an artefact of collection

and preservation. There is no justification for the

retention of two genera, Calvasterias being a

junior synonym of Anasterias by page priority.

Anasterias directa Koehler

Plate 2f-h

Parastichaster directus Koehler, 1920: 97-101, pl.

xx figs 8-11, pl. xxi figs 8-12, pl. xxiii figs 1, 2, pl. Ixii

fig. 2.—Rowe and Pawson, 1977: 342.

Sporasterias directa—Fisher, 1930: 241.—Bennett,

1971, pls 34, 35, 56 fig. 2.

Anasterias directa. —A.M. Clark, 1962: 97, text-fig.

17c.—Simpson, 1982: 43-45, figs 5, 6.—McKnight,

1984: 143.

Sporasterias antarctica— Koehler, 1920: 78-79, pl.

xviii figs 1-4. [non A. antarctica (Lütken, 1857)].

Material examined. (Specimens marked * are typical of

the “antarctica” form). Macquarie Island, 10 Oct 1913,

Low tide. H, Hamilton, AM J3636(2 syntypes of

P, directus, the syntypes Koehler labelled “C” and “D”

were inadvertently destroyed in, 1939 - see Pawson and

Rowe, 1977); H. Hamilton, 1913, identified by

Koehler (1920) as $. antarctica, AM J3726(11*);

Secluded Beach, 27 Nov 1989, NMV F77766(4),

F77767(18*); Garden Cove, Dec 1959, NMV

F45010(15), F76254(30*); 15 Dec 1959, F45012(11*);

20 Dec 1961, NMV F76256(1*); 1961, NMV

F76252(2*); 31 Mar, 1962, NMV F76267(1); 12 Aug

1962, NMV F76264(4), F76249(40*); 11 Feb 1964,

NMV F76263(4), F76253(3*); 16 Jun, 1967, NMV

F76262(5); 18 Jun, 1967, NMV F76261(1*); 14 July,

1975, NMV F76266(1); 25 Nov 1989, NMV

F76274(1*); 26 Nov 1989, NMV F76273(2); Buckles

Bay, 28 Dec 1948, NMV F76248(1 juv*); 16 Jul 1949,

NMV F76247(2 juv*); 9 Feb 1950, NMV F76244(1*);

Feb 1950, NMV F76242(2*); 8 Dec 1962, NMV

F76265(2); Dec 1966, NMV F76270(1 juv); 7 Dec

1986, NMV 76272(1*); Sandy Bay, 7 Feb 1962, NMV

F76271(1 juv), F76251(3*); Green Gorge, 24 Jul 1962,

NMV F76257(4*); Lusitania Bay, 25 Jul 1962, NMV

F76250(4*); Hurd Point, 10 Jan 1962, NMV

F76260(1*); Caroline Cove, 10 Jun, 1962, NMV

F76258(6*): Aurora Point, 8-9 Feb 1962, NMV

F76268(1); Bauer Bay, 8 Oct 1962, NMV F76255(3*);

Hasselborough Bay, 19 Dec 1949, NMV F76241(1);

Aerial Cove, 2 Dec 1949, NMV F76246(1 juv*); 26 Jan

1950, NMV F76245(4 juv*); 28 Nov 1989, NMV

F77768(3), F77769(17*). AM 1977-1978 Expedition,

Gorilla Head Rock (8-12 m) MA-142(2),

MA-147(1*); Tottan Head, Goat Bay (14 m).

MA-39(1), MA-371(1); Tern Rock Bay (7 m),

MA-35(4); Garden Cove (1-10 m) MA-14(4),

MA-127(4*), MA-359(1), MA-360(1); NE Buckles

Bay (intertidal), MA-93(1); Sandy Bay (9 m),

MA-224(1); Green Gorge (int-14 m), MA-245(1),

MA-247(2), MA-253(8), MA-256(1*), MA-259(1*),

MA-280(1*), MA-281(5*); Caroline Cove (8-13 m),

MA-300(1*), MA-306(2*); Handspike Point (inter-

tidal-0.5 m), MA-136(4*), MA-138(2); Aerial Cove

(0.5-6 m), MA-46(3), MA-47(3), MA-50(3*),

MA-83(6), MA-102(2*), MA-107(3), MA-108(4*),

MA-109(1), MA-382(4), MA-383(2*), MA-384(1*),

MA-386(1), MA-388(8); Anchor Rock (13-20 m),

MA-89(1), MA-95(1).

Comparative material examined. A. antarctica (Lütken.

1857): Magellan Strait, Punta Arenas, coll. Schythe.

ZMUC(holotype); Chile, Linao, 41°50’S, 73°34 W.

BMNH 1975.11.12.6(1); Argentina, Puerto Deseado,

identified by Bernasconi, BMNH 1961.7.27.18(1);

Golfo RE Concavi, Isla Calburo, identified by

Bernasconi, BMNH 1961.7.17,19(1); Magellan Strait,

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 191

Fortescue Bay, Discovery stn 724, 0-5 m, 16 Nov 1931,

BMNH, 1948.3.16.657(10); Cape Horn, St Martins

Cove, Ross Antarctic Expedition, BMNH,

1960.4.13.1-2(4). A. minuta Perrier, 1875: Falkland

Islands, Stanley Harbour, BMNH 1960.5.17.19(1). A.

studeri Perrier, 1891: Falkland Islands, Discovery stn

WS91, 191-205 m, 8 Apr 1927, identified by Fisher

(1940), BMNH 1948.3.16.662(10). A. stolidota

(Sladen, 1889): South Chile, Messier Channel, Chal-

lenger, BMNH(holotype). C. asterinoides Perrier,

1875: Falkland Islands (or possibly Auckland Is - see

A.M, Clark, 1962), Dr J. Robertson, BMNH

1844.5.29(4). A. rupicola Verrill, 1876: Kerguelen, Port

Jeanne d’Arc, BANZARE collection 783, identified by

A.M, Clark (1962), SAM K1512(62); Greenland

Harbour, BANZARE stn 54, BMNH 1965.8.5.251-

275(24); Rivett Arm, BANZARE stn 56a, BMNH

1965.8.5.276-281(6); Marion Island, De Villier,

BMNH 1973.12.18.4-11(7); BMNH 1973.12.18.12-

15(4); Cabbage Point, G. Branch, in Macrocystis beds,

12 m, AM J18135(1); Transvaal Cove, G. Branch, in

Maerocystis beds, AM J18136(1). A. suteri (de Loriol,

1894): New Zealand, Lyttelton, Suter, BMNH

1905.1.25.2-3(2); Bodley Head, Oliver, Mortensen’s

Pacific Expedition. ZMUC(4); Banks Peninsula, Tay-

lors Mistake, E Bennett, BMNH 1952.11.18.4(2);

Stewart Island, BMNH 86.11,.18.12(1); Snares Island,

H.B. Fell, BMNH 1952.4.16,19(1); Campbell Island,

Windlass Bay, 23 Jan 1980, AM J22902(2). A. laevi-

gata (Hutton, 1879b): Auckland Island, Port Ross, 25

Noy 1914, Mortensen’s Pacific Expedition,

ZMUC(20); 26 Nov 1914(20); Figure 8 Island, 2 Dec

1914, Mortensen’s Pacific Expedition, ZMUC(4);

Carnley Harbour, Mortensen’s Pacific Expedition, 29

Nov 1914(1); Masked Island, 30 Nov 1914,

Mortensen’s Pacific Expedition, ZMUC(1); Crozier

Point, H.B. Fell, BMNH 1952.4.16.13(3); Tagua Bay,

Dawbin, Noy 1943, BMNH 1952.11.18.3(1); Passage

Inlet, Dawbin, 24 Jul, 1943. BMNH 1952.11.18.5-7(4);

Laurie Harbour, Discovery Expedition, BMNH

1905.7.14.6-9(many); Campbell Island, 26 Jan 1980,

AM J22906(2): Campbell Island, Filhol, donated Paris

Museum 1877, ZMUC(1);

Description of material, R to 60 mm, r to 17 mm.

Arms 5, dorsally convex, ventrally flat, widest at

base, tapering gradually to a blunt tip. Single

madreporite, third to half-way from centre to

interradial margin, small, sometimes inconspicu-

ous, often ringed by some spinelets. Anus incon-

spicuous, central. Arm furrows wide, 4 rows of

suckered tube feet.

Dorsal surface covered in thick pustular skin,

with clusters of papillae and scattered to clustered

low spinelets. Abactinal skeleton reticulate.

Abactinal spinelets capitate, sometimes truncate,

upper half spiniferous; of various sizes, 0.3—-0.8

mm dia, 1-2 times as high as wide. Spinelets

largest, widest on disc and proximal arm surfaces.

Carinal plates in irregular, often sinuous row;

plates square, lobed, L-shaped or irregular in

shape, imbricating. Spinelet distribution variable.

Some specimens with regular transverse arcs of 3

spinelets on succeeding carinal plates, other spec-

imens more irregular with 1-7, often 1, spinelet

on cach plate. Up to 7 dorsolateral plates linking

carinals and superomarginals in irregular trans-

verse rows, sometimes in regular rows or com-

pletely reticulate; plates bar-like or irregular; 1-3

longitudinal linkages proximally, sometimes

forming longitudinal rows. 0-1, rarely 2, spinelets

on dorsolateral plates, up to 6 present in total

between carinals and superomarginals.

Papular areas with clusters of 4—5 papulae in

centre, often forming regular longitudinal rows on

either side of superomarginal plates, otherwise

areas scattered amongst the reticulate skeleton.

1-2 actinal papulae are present between infero-

marginal plates.

Superomarginal plates longer than wide, 4-

lobed, long ventral lobe, shorter dorsal and lateral

lobes, dorsal lobe often not transversely aligned

with ventral lobe but displaced toward arm tip,

plates imbricate with surrounding dorsolateral,

superomarginal, inferomarginal plates. Supero-

marginal spinelets to 1.0 mm in height, 2—2.5

times as high as wide, 1-3 on each plate, 1-2

spinelets near dorsal lobe, 0-1 smaller spinelets

on ventral lobe, rarely some additional very small

spinelets, dorsal spinelets form regular longitudi-

nal rows extending from disc to arm tip, ventral

spinelets sometimes form a less regular and less

extensive row. Inferomarginal plates form an acti-

nolateral margin to arm, wider than long, 4-lobed,

dorsal and ventral lobes shorter than lateral lobes,

prominent oblique spine-ridge present centrally.

Inferomarginal spines thick, capitate, slightly

curved, terminally spiniferous, 2—3, rarely 4, per

plate, 2.5 mm high, 3-4 times as high as wide,

aligned obliquely. Actinal plates bar-lke, wider

than long, extend almost to arm tip, sometimes

2 series proximally, usually aligned with

inferomarginals. Actinal spines, usually 1 per

plate, sometimes 2 on proximal plates aligned

transversely, often confluent with, but smaller

than inferomarginal spines, on larger specimens

spine persists to arm tip. 8-10 adambulacral

plates to every 3 inferomarginals. Adambulacral

spines, one per plate, thinner than infero-

marginal spines, 2.5 mm high, 3—5 times as high

as wide. On each jaw 2 small oral spines, 2 larger

suboral spines, larger than nearest adambulacral

spines.

Straight pedicellariae present in arm furrows

and on jaw, valves with rounded tips, no teeth;

distribution variable, numerous to scarce. Crossed

pedicellariae present on dorsal side of infero-

marginals, near superomarginals and on lower

abactinal surface.

192 T. D. OHARA

Reproduction (from Simpson, 1982). Separate

sexes. Females brood their young from egg to

juvenile stage as a cluster under the disc. Eggs are

transferred to the brood in July and juveniles are

released in October-November. Females adopt an

arched posture to hold the brood.

Colour (live). Dorsal surface dark olive green or

green brown, sometimes mottled at arm tip,

spinclets pale, madreporite white. Ventral surface

white, tube feet and tips of adambulacral spines

tan. Juveniles pink or orange, egg mass orange.

Habitat. Shore collectors have found this species

in sheltered and exposed sites, including intertidal

rock pools, roek gutters and boulders, and

amongst Durvillaea holdfasts. Divers (to 20 m)

have also found it on sheltered and exposed

rock surfaces, and amongst Macrocystis beds.

McKnight (1984) has reported one dredged

specimen from 55 m.

Distribution. Macquarie Island (0-55 m).

Remarks. The genus Anasterias presents consid-

erable difficulties for taxonomists. Many of the

nominal species are very variable and ill-defined.

There have been many disagreements about syn-

onymies, particularly regarding Magellanic and

Falkland Islands species (Fisher, 1940; A.M.

Clark, 1962; A.M. Clark and Downey, 1992),

Macquarie Island specimens are also very vari-

able in many characters. These include the shape,

density and distribution of abactinal spinelets; the

rigidity and regularity of the abactinal skeleton;

the nature of the abactinal integument, the num-

ber of inferomarginal spinelets; and the distribu-

tion of the various pedicellariac. Some characters

such as the shape of the arm, the density of the

skeletal plates and the pustular nature of the skin

can vary with preservation. Many of the charac-

ters are difficult to quantify and do not lend them-

selves to traditional morphological analysis.

Koehler (1920) recognised two five-armed

species from Macquarie Island: Sporasterias

antarctica (Lütken,

directus Koehler, 1920. Koehler (1920) did not

directly compare the two species, but the under-

standing of the two genera at that time, indicates

that they differed in the regularity of the abactinal

skeleton and the density of the abactinal spinelets.

Fisher (1923, 1930, 1940) subsequently syn-

onymised the genera Sporasterias Perrier, 1894

and Parastichaster Koehler, 1920 with Anaste-

rias, noting the continuous variation in these

features. All subsequent five-armed specimens

collected from Macquarie Island have been

referred to the species A. directa.

1857) and Parastichaster

Two morphological extremes are recognisable

in the current large range of specimens that

correspond to Koehler’s “species”, One form

(“antarctica”) has an irregular abactinal skeleton;

a sinuous, irregular carinal series; and 1-2, occa-

sionally 3, carinal spinelets on each plate (pl. 2h).

These specimens are broadly similar to the 11

extant specimens of S. antarctica identified by

Koehler (AM J3726). The other extreme

(“directa”) has a more regular, strongly built dor-

solateral skeleton, often with regular longitudinal

rows of plates and spinelets (pl. 2g). The carinal

plates usually bear three spinelets in a transverse

arc, although plates with one or two spines are not

rare and other plates can have as many as seven.

These features are observable in specimens as

small as R = 2 mm. This form is similar to the two

extant syntypes of P. directus. However, between

these extremes there are many variations. Carinal

plates with three spinelets can be present on spec-

imens with an irregular skeleton. Sometimes the

arrangement of carinal spines can vary on the

same animal. Moreover, as listed earlier, many

other features vary in a similar fashion in both

forms. There is no consistently observable

morphological characteristic that can be used to

satisfactorily separate the two forms.

Similar morphological differences exist within

material from the Magellanic region and from the

subantarctic Islands of New Zealand. The

extremes have been generally distinguished as

species. In the South American region, strongly

calcified specimens have been distinguished as

Calvasterias stolidota Sladen, 1889 or A. varia

Philippi, 1870 from the more common A. antarc-

tica. A further form with a very weakly calcified

skeleton, represented in South America by A.

minuta Perrier, 1875, does not appear to exist at

Macquarie Island. A.M. Clark and Downey

(1992) have noted the variability of all the

Magellanic forms and record only A. antarctica

as representative of the group. Of the numerous

nominal species of Anasterias from South

America, only A. studeri appears to be readily dis-

tinguishable from A. antarctica, having numerous

long abactinal spinelets, triangular straight pedi-

cellariae, and a preference for deep water (< 100

m) habitats (Fisher, 1940; A.M. Clark, 1962:

A.M. Clark and Downey, 1992). A. antarctica is

recorded as being dark green in colour when alive

(Madsen, 1955) and broods its young in the typi-

cal arched posture. Specimens I have examined

have relatively few carinal and dorsolateral

spinelets and many specimens, unlike my

Macquarie material, have numerous crossed pedi-

cellariae on the marginal and abactinal surfaces.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 193

In the subantarctic Islands of New Zealand, two

species are recognised: 4. suteri with a regular,

strongly calcified skeleton and A. laevigata with a

more weakly calcified skeleton. A. laevigata is

similar to the Macquarie Island “antarctica”

form. Mortensen (1925) describes the abactinal

spination as being very variable, some animals

have an irregular single row of carinal spines and

1-2 irregular dorsolateral rows, others have no

carinal or dorsolateral spines at all, rarely a spec-

imen can have 2-3 carinal spines. This variation

is also present in some of the specimens I have

examined. The distribution of pedicellariae is also

very variable. Some specimens have numerous

crossed pedicellariae abactinally while others

have numbers of straight pedicellariae. Colour is

reported as dark green or grey-green with fawn

arm fips for adults, and orange for juveniles (H.B.

Fell, 1953). Brood juveniles are found between

August and November (H.B. Fell, 1953). A. suteri

on the other hand is clearly distinct having a

dense cluster of 4-8 spinelets on each carinal

plate. There is one, rarely two, longitudinal rows

of dorsolateral spines and 1-2 spines on the

superomarginal plates, The colour is fawn-brown

or grey-green; brood juveniles have been found in

October (H.B. Fell, 1953).

At Kerguelen and Marion Islands, only one

five-armed form is recognised: A. rupicola Ver-

rill, 1876, which has an irregular abactinal skele-

ton. No five-armed Anasterius species have been

found at Heard Island. Specimens of 4. rupicola

from Kerguelen are indistinguishable from the

“antarctica” specimens from Macquarie Island.

Specimens I have examined measure R=9—40 mm

and have a sinuous carinal series with 1-3

spinelets. Sometimes carinal spinelets can be

arranged in a transverse arc of three spines on at

least part of an arm, however, none approach the

regular arrangement of the “directa” specimens

from Macquarie Island, They have up to six dor-

solateral spines between the carinals and supero-

marginals, 1-2 spines on the superomarginal dor-

sal lobe and one on the ventral lobe, 2-4, usually

three, inferomarginal spines and one actinal spine.

Two specimens collected in February are arched

with traces of brood juveniles around the mouth.

Specimens of A. rupicola from Marion Island

appear to have relatively few spines. Specimens |

have examined have a single carinal spine, 2-3

dorsolateral spines between the carinals and

superomarginals, 1-2 superomarginal and 2-3

inferomarginal spines. Bernasconi (1971)

recorded numerous “thick” abactinal spines from

his Marion Island specimens, including 3-4 dor-

solateral spines transversely. The carinal “lines”

were scarcely visible. Live colour is green or

orange (Branch et al., 1993). Females start brood-

ing eggs in June to August, young are released in

December—January although remnant young con-

tinue to be present as late as March; relatively few

females brood within a population (Blankley and

Branch, 1984).

The two forms from Macquarie Island could

represent two closely related perhaps hybridising

species or one polymorphic species. However,

there appears to be no obvious geographic, bathy-

metric or ecological premating reproductive iso-

lating mechanism now operating to separate the

two forms at Macquarie Island. There is an abun-

dance of morphological intermediates. The two

forms have been regularly found from similar

sites at the same location. Their reproductive

cycles are similar. In the absence of a clear mor-

phological distinction, and without molecular evi-

dence to the contrary, I am referring all the five-

armed Macquarie Island Anasterias specimens to

the single polymorphic species: A. directa. Given

this interpretation, this species differs from

A. antarctica, A, rupicola and A. laevigata by the

greater range of variation expressed in the number

and distribution of spinelets and the restricted

range of variation in the abundance of abactinal

pedicellariae. Probably A. directa, A. rupicola and

A. laevigata should only be regarded as infra-

specific regional forms of A. antarctica, It is clear

that traditional morphological characters are

insufficient to properly differentiate the various

taxa in this genus.

Anasterias mawsoni (Koehler)

Plate 2i-k

Stichaster suteri.—Benham, 1909: 32 [non Calvaste-

rias suteri de Loriol, 1894].

Parastichaster mawsoni Koehler, 1920: 91-97, pl.

xix figs 1-8, pl. xx fig. 1, pl. xxi figs 1-6, pl. xxiii fig.

4, pl. xxiv fig. 5, pl. xxx fig. 5, pl. Ixiii fig. 2—Rowe

and Pawson, 1977: 345.

Parastichaster _ sphoerulatus Koehler, 1920:

101105, pl. xxi fig. 7, pl. xxiii figs 5-10, pl. xxiv figs

1-4, pl. Ixiii fig. 3, pl. Ixiv figs I, 2.—Rowe and

Pawson, 1977: 347. [new synonymy]

Sporasterias —mawsoni.—Fisher, 1930; 241 —

Bennett, 1971, pl. 56 fig. 1.

Sporasterias sphoerulata—Fisher, 1930: 241.—

Bennett, 1971, pl. 56 fig. 2.

Anasterias mawsoni—A.M. Clark, 1962: 96—

Simpson, 1982: 41-43, figs 3, 4.—McKnight, 1984:

143.

Anasterias _ sphoerulata— A.M.

96-97.—McKnight, 1984: 143.

Material examined. (Specimens marked * are typical of

the “sphoerulata” form). Macquarie Island, 10 Oct

Clark, 1962:

194 T. D. OHARA

1913, AM J3638(4 syntypes of P. mawsoni); Oct 1913,

AM J3637(2 syntypes of P. sphoerulatus); Secluded

Beach, 19 Jan 1965, NMV F76177(1); 27 Nov 1989,

NMV F76186(7*); Garden Cove, 7 May 1949, NMV

F76169(1); Dec 1959, NMV F45009(4 and brood); 15

Dee 1960, NMV F45013(1*); 13 Jul 1962, NMV

F76171(1*); 12 Aug 1962, NMV F76175(1), NMV

F45109(1); 11 Feb 1964, NMV F76188(2); 18 Jun

1967, NMV F76176(3); 25 Nov 1989, NMV

F76184(5*): 26 Nov 1989, NMV F76185(1*); Buckles

Bay, 28 Dec 1948, NMV F76165(2); 16 Jul 1949, NMV

F76130(2), NMV F76164(2), NMV 761621); 9 Feb

1950, NMV F76168(1), NMV F76181(2*); 9 Feb 1952,

NMV F76161(2); 8 Dec 1962, NMV F76170(7); 7 Dec

1986, NMV F76180(1); Hasselborough Bay, 31 Mar

1950, NMV F76182(1*); Aerial Cove, 28 Nov 1989,

NMV76187(6). AM 1977-1978 Expedition, Gorilla

Head Rock (9-12 m) MA-142(2), MA-145(1),

MA-146(5), MA-148(1), MA-149(11); Tottan Head,

Goat Bay (9-14 m) MA-39(1), MA-42(4*),

MA-369(1), MA-371(5), MA-374(5), MA-375(6),

MA-376(2 and brood); Tern Rock Bay (7-10 m),

MA-35(5*), MA-37(2); Garden Cove (8-14 m)

MA-14(11*), MA-15(1*), MA-87(1*), MA-91(1),

MA-123(1), MA-124(1), MA-125(2*), MA-127(7),

MA-128(6*), MA-379(4); Sandy Bay (9 m),

MA-224(1*); Green Gorge (6-18 m), MA-245(4), MA-

247(6), MA-250(1), MA-251(2), MA-269(1),

MA-275(1), MA-291(2); Caroline Cove (8-13 m),

MA-298(2), MA-300(1), MA-306(2), MA-311(8);

Aerial Cove (0.5-6 m) MA-46(4), MA-47(20),

MA-50(3), MA-56(1), MA-83(8), MA-98(2),

MA-100(3), MA-102(6), MA-107(12*), MA-108(7),

MA-109(5 and brood), MA-110(5), MA-382(>24),

MA-383(2), MA-386(17), MA-388(7), MA-380(4);

Anchor Rock (13-20 m), MA-89(3*), MA-90(6*).

Heard Island, Atlas Cove, shore, 28 Dec 1929, BAN-

ZARE stn, 19, identified by A.M. Clark (1962) as A.

sphoerulatus, SAM K1515(2*); 19 May 1949, NMV

F76125(1); 11 Sep, 1949, NMV F76127(1 juv); 16 Jan

1950, 30 m, NMV F76124(2); 3 Jul 1950, NMV

F76129(9 juv); 16 Aug 1950, NMV F73886(11); 19

Dec 1951, NMV F76128(1); 28 Jan 1952, NMV

F76179(1); 8 Feb 1952, NMV F76126(1).

Comparative specimens examined. A. perrieri (Smith,

1876): Kerguelen, Bras Bossiere, 4 m, BANZARE stn

7, SAM K1497(12); BANZARE collection 712, 9 m, 10

Nov 1930, SAM K1508(1).

Description of Macquarie Island material. R to

62 mm, r to 15 mm. Arms 6, dorsally convex,

ventrally flat, widest at base, tapering gradually to

a blunt tip. Single madreporite, half to two-thirds

from centre to interradial margin, small to large,

sometimes inconspicuous, sometimes ringed by

some spinelets. Anus inconspicuous, central. Arm

furrows wide, 4 rows of suckered tube feet.

Dorsal skin thick and pustulate. Abactinal

skeleton reticulate. Spinelets capitate, sometimes

truncate, upper half spiniferous; often polygonal

in cross section if in contact with adjacent

spinelets; size variable, 0.4-1.1 mm dia, 1-2

times as high as wide; spinelets largest, widest on

disc and proximal arm surfaces. Spinelets

arranged in rows around aboard papular areas.

Carinal plates form recognisable longitudinal

row down arm; plates irregularly quadrilobed or

rhombic, broadly imbricating. Spinelets distribu-

tion variable, from being densely clustered on

plates, with up to 13 spinelets in 5 longitudinal

rows, to scattered with as few as 2-4 discrete

spinelets. Dorsolateral skeleton variable, from

strongly calcified with block-like imbricating

plates to weakly calcified with thin bar-like or

trilobed plates; forming transverse rows of up to 7

plates between carinals and superomarginals;

forming 1-3 longitudinal rows, the row nearest

the carinals most distinct; sometimes reticulate

proximally, Dorsolateral plates with 0-3, usually

1-2, spinelets on raised section of plate: forming

uni- or biserial rows transversely, with up to

12 spinelets present between carinals and

superomarginals, forming irregular rows

longitudinally.

Papular areas with clusters of 4-8 papulae,

single actinal papulae present between

inferomarginal plates.

Superomarginal plates from as long as to longer

than wide, irregularly quadrilobed, long ventral

lobe, shorter dorsal and lateral lobes, plates imbri-

cate with surrounding plates. Superomarginal

spinelets to 1.0 mm in height, 2—2.5 times as high

as wide, 2-7, usually 2-3, per plate. Infero-

marginal plates forming actinolateral margin to

arm, as wide as long, lobed, prominent oblique

spine-ridge present centrally. Inferomarginal

spines thick, capitate, slightly curved and termi-

nally spiniferous or occasionally flattened and

flared; 2-5, usually 3-4, per plate, 2.5 mm high,

2.5-4 times as high as wide. Actinal plates bar-

like, wider than long, extend past half R, some-

times 2 series proximally, usually aligned with

inferomarginals, with one spine, often confluent

with, but smaller than inferomarginal spines. 6

adambulacral plates to every 2 inferomarginals.

One adambulacral spine per plate, thinner,

smaller than inferomarginal spines, 2.5 mm high,

4-5 times as high as wide, often club-shaped. On

each jaw 2 oral and 2 suboral spines; oral spines

often short,

Straight pedicellariae present near mouth, in

disc interradius, in arm furrows and amongst

inferomarginals; size and distribution variable,

sometime very numerous or scarce; some yalves

widened and truncate at tip; minority (typically

10 %) 3-valved, with additional small valve pre-

sent at right angles to 2 primary valves. Crossed

pedicellariae present marginally and dorsally,

sometimes in high numbers.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 195

Reproduction. Macquarie Island population with

separate sexes. Females brood juveniles under

their discs by adopting an arching posture. Broods

with up to 296 juveniles. Eggs mature in gonad

from July to December until the eggs reach 2.0

mm diameter. Eggs are usually transferred to the

brood in January-Febuary and released in

May—June (Simpson, 1982). Two specimens

in the present collection have brood juveniles

in December and January.

Colour. Live Macquarie Island specimens orange,

tan, dark brown, purple or green on dorsal sur-

face; spinelets and papulae often paler or different

colour; dark stomach caeca visible beneath skin

on smaller specimens. Ventral surface cream or

white, spines pink or grey, tube feet tan.

Habitat. Present on rock platforms under boul-

ders, or in rock gutters, and subtidally (2-20 m)

on sheltered sponges-bryozoan mats, on Codium,

or under Macrocystis. McKnight (1984) reported

several dredged specimens from 357 m.

Distribution. Heard Island (0-30 m); Macquarie

Island (0-357 m).

Remarks. The six-armed Macquarie Island speci-

mens of Anasterias are as variable as the five-

armed specimens described above under A.

directa. The dorsal skeleton varies from being

strongly calcified, with block-like plates and a

dense covering of polygonal spinelets (pl. 2k), to

a more open structure with thin bar-like plates and

relatively few discrete rounded spinelets (pl. 2j).

The former specimens were described by Koehler

(1920) as P. sphoerulatus and the latter as

P. mawsoni. However, there are numerous inter-

grades between these extremes. Koehler’s other

distinguishing character for A. sphoerulatus, the

unusual three-valved pedicellariae, are present on

most specimens. The different colour forms do

not correlate with skeletal structure or spinelet

density. Consequently, I refer A, sphoerulatus to

the synonymy of A. mawsoni.

This species is also common from the littoral of

Heard Island. Heard Island specimens show a

similar range of variation in the skeleton and

spinelets. They also have some three-valved pedi-

cellariae. A.M. Clark (1962) noted the reduced

actinal series on BANZARE specimens from

Heard Island. This is also a variable feature,

Several specimens from Heard Island (NMV

E73886) have a well developed primary actinal

series extending to the arm tip, and a second

series proximally.

The only other six-armed Anasterias species

is A. perrieri (Smith, 1876) from Kerguelen.

This species appears to occupy a similar ecologi-

cal niche at Kerguelen (McClintock, 1985) as A.

mawsoni at Macquarie Island, but it differs from

A. mawsoni in having thinner and less numerous

spinelets. Specimens of A. perrieri that I have

examined have no three-valved pedicellariae.

A, mawsoni is not merely a six-armed form of

A. directa. Besides the differing number of arms,

the two species differ in the density of the abac-

tinal spinelets, the colour, and the absence in

A. directa of three-valved pedicellariae. There is

also a time difference of four months between the

peak release of brooded young for each species

(Simpson, 1982).

Class Ophiuroidea

Key to Macquarie Island Ophiuroidea

Disc and arms covered in thick skin concealing any underlying plates or

scales unless dried. Arms cylindrical in cross section. Oral papillae and teeth

expanded with glassy toothed margin … nanne eeneerverenen Ophiomyxa sp.*

Disc and arms not obscured by skin, although cover of spines may be pre-

sent. Arms somewhat flattened in cross section … aan onvernerneeneenvervenn 2

2 squarish papillae at apex of each jaw ….sucnsarvononrnenenrnenreerenvenseenensmen sense 3

1 papilla at apex of each jaw (occasionally an additional oral papillae apical

in position, but never 2 squarish papillae) nan anene nor venernenneeseenenrnen 5

2 low oral papillae on side of each jaw in addition to apical ones, distalmost

papilla 2—3 times as long as proximal one … Amphipholis squamata?

1 distal oral papilla on each side of jaw, separated from apical one by wide

APL ANALE TE TEELEN TELENET, 4

6—7 arm spines, lowermost spine considerably longer than the upper spines.

Ventral disc surface covered in scales … Amphiura magellanica

4 subequal arm spines. Ventral disc surface naked near oral shields .............-.

EREN EMDR cca a etch PDAS Aden Amphiura cf. angularis®

Arms inserted laterally into disc and firmly fused to it. No disc spines … 6

Arms inserted ventrally under the disk, disk margin overlying base of arm … 9

196

T. D. O'HARA

6. Dorsal arm plates fragmented into 2 lateral plates and several smaller pieces

in between. No arm comb aranea Ophioplocus incipiens

T Dorsal arm plates entire. Arm comb present T Ì

Disc covered by large circular scales surrounded by smaller plates. 3 minute

arm spines, difficult to see … annen Ophiurolepis inornata%

— Nokasabogvo vti re Neal te Er SNE 8

Distal arm spines with upturned hook-like ends, Small accessory plates lat-

eral to each proximal ventral arm plate … Ophiura (Ophiuroglypha) irrorata

— Arm spines simple and pointed. No accessory ventral arm plates …

zene (eternet EE Ophiura (Ophiura) meridionalis

9. Dorsal disc plates bordered by rows of spherical granules. Disc margin bor-

dered by row of elongate granules. Arms very slender … Ophioleuce regulare

= Disc plates naked or with spines, never with granules … 10

10. Apical jaw papillae wider than long, rectangular or tricuspid. 6 arms. Asex-

ually reproduces though fission neen. Ophiactis hirta

— Apical papillae longer than wide, usually pointed. 5 arms … 1]

11. Disc stumps large, cylindrical or club shaped, regularly covered by rows of

tiny sharp thorns. Radial shields as wide or wider than long, not covered in

plates Or StUMPS … cpnpiiveevicsnetsecsevesencetacerdesetntadienetnes Ophiomitrella conferta®

— Disk spines small, irregular or conical with terminal thorns. Bar-like radial

shields largely concealed by disc plates and spines (distalmost end may be

SRPOSCA) E E E E ES E E A O AE E le 12

12. Distalmost oral papilla widened. Tentacle scales half the size of the ventral

BET plate oe a ntt Bere haidi eih er Baran Ophiacantha sollicita'

— Oral papillae subequal or proximal papilla slightly enlarged. Tentacle scale

almost as long as ventral arm plate … nn. Ophiacantha vilis

Notes on key:

è McKnight (1984: 144) recorded two specimens of an undetermined Ophiomyxa species from

Macquarie Island (183-415 m). As McKnight pointed out, they share several characteristics of

O. brevirima H.L Clark, 1915 from New Zealand, including alternating arm spines, On one 7

mm d.d. specimen I have examined (NZOI stn D6, 55°27’S, 158°31.5’E, 415 m), a large

rounded upper arm spine is present on some arm segments in addition to the 2 smaller spines.

However, unlike O. brevirima the dorsal arm plates are entire, bell-shaped and contiguous. The

material is not adequate for a complete description.

b McKnight (1984: 144) recorded one specimen of the cosmopolitan species A. squamata

(Della Chiaje, 1828) from Macquarie Island (155 m).

€ Two specimens of A. angularis Lyman, 1879 were recorded by McKnight (1984: 144) from

Macquarie Island (NZOI stn D6, 55°27'S, 158°31.5'E, 415 m). I have examined several spec-

imens and they broadly match Lyman’s (1882) description and figures (pp. 134-135, pl. XXIX

figs 1-3) except that they lack tentacle scales. The disc is naked near the oral shields and

orange gonads are visible underneath. They are much smaller (2 mm d.d.) than Lyman’s holo-

type (9 mm d.d.) and the lack of tentacle scales may be a Juvenile character or an artefact of

preservation. Without a good growth series they cannot be positively identified. A. angularis

has been recorded from Kerguelen, Heard, Marion Islands, South Africa and New Zealand.

However, not all these populations appear conspecific. Mortensen (1936) noted that specimens

from South Africa and Kerguelen are scaled on the ventral side of the disc. Branch et al. (1993)

have figured a specimen with very long arms. A thorough review is required.

d McKnight (1984: 145) recorded one specimen of the widespread deep-water species

O. inornata (Lyman, 1878) from the south of Macquarie Island (NZOI stn DS, 56°40.6'S,

158°45.5’E, 1280 m).

“McKnight (1984: 144) recorded 3 specimens of O. conferta (Koehler, 1922b) from the north-

ern Macquarie Ridge (NZOI stn D159, 49°01'S, 164°50’E, 741 m), O’Hara (1990: 299) has

recorded this species from off southeastern Australia, Antarctica and the Magellanic region

from 40-2340 m.

f McKnight (1984: 144) has recorded 28 specimens of O. sollicita Koehler, 1922b from two

NZOI stations on the northern Macquarie Ridge (NZOI stns D17 and D1 8, 52°31°S, 160°31'E,

124-128 m). The specimens from NZOI stn D17 have apparently disintegrated (P. Anderson,

pers, comm.), Three specimens from stn D18 were examined and compared to the syntypes

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 197

from off eastern Tasmania. The Macquarie Island specimens differ in having slightly smaller

disc spines (to 0,1 mm, compared to 0.15 mm for the syntypes) which are absent ventrally, and

wider ventral arm plates. More material is required to determine if the Macquarie Island

specimens represent a distinct species. O. brachygnatha H.L. Clark, 1928 from off south-

eastern Australia and New Zealand is also similar but has much thinner, moniliform arms and

smaller disc spines. O'Hara (1990; 295-296) noted the similarity of O. sollicita with the other

nominal species from the Atlantic.

Order Ophiurida

Ophiacanthidae

Ophiacantha vilis Mortensen

Plate 3a—c, Figure 4

Ophiacantha vilis Mortensen, 1924; 114—7, fig. 7—

H.B. Fell, 1952: 14.—H.B. Fell, 1958: 25.—McKnight,

1967: 308,

Ophiacantha pentagona—Madsen, 1967: 59—

McKnight, 1984: 144. [non O. pentagona Koehler,

1897]

Material examined. Macquarie Island, off Lusitania

Bay, 54°42.5'S, 158°54.5'E, 69 m, 5 Dec 1930, BAN-

ZARE stn 83, SAM K947(1). AM 1977-1978 Expedi-

tion, Tottan Head, Goat Bay (14 m), MA-371 AM

J22265(3), MA-376 AM J22266(1) and AM J22267(6);

Garden Cove (10-14 m), MA-123 AM J22258(1), MA-

124 AM J22262(1), MA-127 AM J22259(1) and AM

J22260(4), MA-128 AM J22261(1), MA-379 AM

J22263(1) and AM J22264(4); Buckles Bay (15 m),

MA-141 AM J22269(1); Caroline Cove (8 m), MA-300

AM J22268(1); Aerial Cove (6 m), MA-388 AM

J22270(1). ANARE 1986 Expedition, BT3 NMV

F60293(1), BT10 NMV F60292(1).

New Zealand, Cook Strait, 372 m, 13 Aug 1920,

ZMUC(syntype).

Comparative specimens examined. O. imago Lyman,

1878: Kerguelen, Christmas Harbour, 220 m,

Challenger, BMNH 1882.12.23.318(holotype).

Deseription of Macquarie Island material. Disc

1-7 mm d.d., arms to > 20 mm. Disc covered in

thin imbricating scales, usually bearing 1 small

conical or cylindrical stump with 4-8 terminal

thorns. Radial shields concealed. Ventral disc

stumps smaller, less numerous towards oral

shields. Oral shields trapezoid, wider than long,

distal lobe often slightly produced. Adoral shields

long, narrow, slightly curved, meeting interradi-

ally, separated radially. Apical papilla leaf-like,

pointed, or irregularly widened on larger speci-

mens; 3, rarely 4, oral papillae, innermost largest,

flattened, leaf-like, outer papillae cylindrical,

blunt. Dorsal arm plates triangular with convex

distal edge, proximal plates just contiguous on

larger specimens, other plates separate. Second

ventral arm plate triangular, other plates roughly

pentagonal with convex notched distal edge,

proximal plates wider than long, plates separate

throughout. 6, rarely 5 or 7, thick, often finely ser-

rated, bluntly pointed arm spines, uppermost

spine longest, on proximal plates enlarged to 1.5

times length of second spine, almost meeting over

dorsal midline, lower spines progressively

shorter, thinner. Tentacle scale large, almost as

long as ventral arm plate, flat, oval or truncate,

sometimes terminally thorny.

Reproduction. Viviparous, two specimens (5 mm

d.d, NMV F60292; 5.5 mm d.d., AM J22262)

with a juvenile in several bursae, other specimens

with large orange eggs.

Colour. Dorsal disc and arm surfaces brown or

tan, paler ventrally, white arm spines (in alcohol).

Habitat. Diver-collected specimens from Mac-

quarie Island have been found on rocks amongst

sponges, red algae, bryozoans and hydroids; one

specimen was found on coarse sand.

Distribution. Macquarie Island (6-433 m); New

Zealand (90-1090 m); Chatham Rise (241-251 m).

Remarks. Close examination of the Macquarie

Island specimens has revealed that they are closer

in form to O. vilis of New Zealand rather than the

Indo-Pacific species, O. pentagona, to which they

have been previously referred. O. pentagona (as

described and figured by Koehler 1897 and,

1922a) has minute tentacle scales and slender disc

spines with 3-4 long sharp terminal thorns

(Koehler, 1922a, pl. 93, fig. 5). In contrast, com-

parison of a syntype of O. vilis (4 mm d.d.) with

Macquarie Island specimens of similar size,

revealed no significant morphological differ-

ences. The syntype of O. vilis closely agrees with

Mortensen’s (1924) description and figures,

except that the radial shields are widely separated

and disc spines relatively larger than shown on

the figures. Photographs of the syntype are repro-

duced here (pl. 3b, c). The tentacle scales are

large, often as long as, or longer than, the ventral

arm plate; there are 6-7 finely serrated proximal

arm spines and the disc spines are short stumps

with 3—4 terminal thorns. There is much variation

in the shape of the oral shields in the Macquarie

material, particularly in the degree of prominence

of the distal lobe, H.B. Fell (1952) has reported a

similar variation in New Zealand specimens.

198 T. D. O'HARA

Plate 3. a, Ophiacantha vilis, dorsal view, NMV F60293; b, Ophiacantha vilis, syntype, dorsal view of disc,

ZMUC; c, Ophiacantha vilis, syntype, oral view of disc, ZMUC; d, Amphiura magellanica, dorsal view

(several arms broken), NMV F60294; e, Amphiura magellanica, ventral view (one arm missing), NMV

F60294; f, Ophiura irrorata, dorsal view (arm tips broken), NMV F60298; g, Ophiura meridionalis, dorsal

view of arm (arm tips broken), NMV F60297; h, Ophioplocus incipiens, dorsal view, NMV F60299;

i, Ophioleuce regulare, ventral view (arm missing from upper right), NMV F60314; j, Ophioleuce regulare,

dorsal view (arm missing from lower right), NMV F60314; k, Florometra austini, lateral view (several arms

broken, most cirri lost), NMV F60300.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 199

Figure 4. a-b, Ophiacantha vilis, NMV F60292: a, dorsal view of base of an arm and adjoining disc;

b, ventral view of arm base and adjoining jaws.

There are, however, differences in the reported

size, bathymetric distribution and reproductive

behaviour between the Macquarie Island and New

Zealand specimens, The Macquarie Island speci-

mens grow to 7 mm d.d., while the largest

reported New Zealand specimen is 4 mm d.d..

Several obvious growth changes occur in the

Macquarie Island specimens. Most notable is the

change in disc spines which are relatively shorter,

wider and with more thorns on large specimens.

Future research may find detectable differences

between Macquarie Island and larger New

Zealand specimens. The Macquarie Island form is

common in the upper subtidal zone (in fact it is

the only ophiuroid known above 20 m depth),

while the New Zealand form has only been col-

lected below 100 m. This species could prefer the

colder water which surrounds Macquarie Island.

Finally, the Macquarie Island specimens are

viviparous, while H.B, Fell (1958) states that the

New Zealand specimens are not. This again could

be due to the small size of the New Zealand

material. None of the present Macquarie Island

specimens under 5 mm d.d. show signs of

viviparity. Further evidence is needed to deter-

mine if the two populations should be separated

into distinct specific or subspecific taxa.

Another similar species, O. imago Lyman,

1878, has been recorded from Kerguelen, Marion

Island and New Zealand. Examination of the

holotype of that species (7 mm d.d.) revealed that

it differs from the Macquarie Island specimens in

having minute tentacle scales, widely separated

dorsal arm plates proximally, longer arm spines,

to 3 mm long, and conspicuously thorny distal

arm spines. Otherwise it is very similar to the

Macquarie Island specimens, particularly in the

shape of the disc spines, and it is also viviparous.

It agrees closely with Lyman’s (1882) description

and figures. Whether all the New Zealand speci-

mens referred to O. imago belong to that species

needs close examination. O. imago has not been

satisfactory separated from O. pentagona and

both share the same minute tentacle scale and arm

spine number. Very little data on the New

Zealand specimens has been published.

200 T. D. CHARA

Amphiuridae

Amphiura magellanica Ljungman

Plate 3d—e

Amphiura magellanica Ljungman, 1867: 320.—

Mortensen, 1924: 132, figs 14, 15a—Madsen, 1967:

129.—McKnight, 1984: 144 [limited synonymy].

Material examined. Macquarie Island, off Buckles Bay,

54°30'S, 158°58'E, 15 m, 19 Oct 1983, NMV

F82954(4); off Lusitania Bay, 54°42.5°S, 158°54.5’E,

69 m, 5 Dec 1930, BANZARE stn 83, SAM K1038(4),

ANARE 1986 Expedition, BT2, NMV F76112(1);

BT3, NMV F60294(2); BT4, NMV F76222(3); BTS,

NMV F60295(1); BT7, NMV F76111(7); BT8, NMV

F60296(2).

South America, Tierra del Fuego, Discovery stn

1321, identified by Mortensen (1936), BMNH

1936.12.30,531-540(10). Gough Island, Discovery stn

399, BMNH 1936.12,30.521-530(12). Auckland

Island, Tagua Bay, identified by H.B. Fell (1953),

BMNH 1952.11.18.31-32(6),

Comparative material examined. A. _ spinipes

Mortensen, 1924: New Zealand, Discovery stn 941,

BMNH 1936.12.30,541—550(S0).

Description of Macquarie Island material. Disc

2-6 mm d.d.. Disc scales small, 0.1—0.3 mm dia,

overlapping, flat. Radial shields, 0.2 times d.d., 5

times longer than wide, separated, divergent cen-

trally. Ventral disc scales smaller, persist until

oral shields. Oral shield rounded, usually wider

than long. Adorals separated radially and slightly

interradially. 2 large block-like infradental papil-

lae; distal oral papilla large oval flat, 2 times

higher than wide, larger than infradental papillae.

Dorsal arm plates rounded, slightly wider than

long proximally, triangular or fan-shaped distally,

contiguous. Ventral arm plates rectangular,

slightly longer than wide, contiguous, 6-7 arm

spines proximally, 4 distally, lowermost spine

enlarged, 1.6 mm, up to 3 arm segments in length,

slightly curved, second lowest 1 segment in

length, uppermost spines smaller, subequal, two-

thirds segment in length. Large oval tentacle

scale, half to two-thirds arm segment long.

Reproduction. Viviparous, juvenile arms emerg-

ing from some bursae of the Macquarie Island

specimens.

Colour. Dorsal disc surface grey, arms and

ventral surface tan (in alcohol).

Habitat. Maequarie Island specimens collected by

divers have been within Macrocystis holdfasts

(NMV F82954). Mortensen (1924) collected New

Zealand specimens from under stones, amongst

algal turf, on sand and sandy mud.

Distribution. Southern South America, Falkland

Islands Gough Island (0-140 m); Macquarie

Island (15-450 m); New Zealand (6-20 m);

Auckland and Campbell Islands (0-90 m);

southern Australia (220-310 m).

Remarks. These specimens are typical of this

species and closely match Mortensen’s (1924)

description and figures. They are also identical to

New Zealand and Atlantic specimens I have

examined. A. spinipes from New Zealand is mor-

phologically similar, differing only in its more

delicate appearance. Mortensen (1924) further

distinguished the two species by their repro-

duction, A. spinipes not being viviparous or

hermaphroditic.

A, magellanica has been assumed to have a cir-

cumglobal subantarctic distribution (Madsen,

1967). Mortensen (1924) suggested epiplanktonic

dispersal by drifting Macrocystis holdfasts. How-

ever, a close examination of its reported range

indicates that it has a disjunct mostly cool-

temperate distribution with two separate popu-

lations, the first around the southern tip of South

America and across to Gough Island in the South

Atlantic and the second in the Tasman region,

including New Zealand, Macquarie Island and

possibly southern Australia, The time required to

drift on the West Wind Drift current between

New Zealand and South America is over two

years making epiplanktonic dispersal unlikely

(O'Hara, 1998a). Another potential explanation

for this distribution is that it reflects vicariant

events associated with the break up of the ancient

continent of Gondwana (O’Hara, 1998a). How-

ever, it is also unlikely that two widely separate

populations would retain species integrity over

such a long time period and further research may

find that the two populations can be distinguished

on molecular or developmental grounds

Ophiactidae

Ophiactis hirta Lyman

Ophiactis hirta Lyman, 1879: 39, pl. 13 figs

365-367.—Lyman, 1882: 118-119, pl. 20 figs 4-6.—

Mortensen, 1924: 126, figs 12a-c—H.B. Fell, 1958:

26— MeKnight, 1984: 144-5,

Material examined. Northern Macquarie Ridge,

52°31'S, 160°31'E, 128 m, 23 Apr 1963, NZOI stn

D18, NIWA(26).

Description of Macquarie Island material. Disc

1—4 mm d.d., arms 5 times d.d. Arms 6, often 3

arms regenerating following fission. Disc round

or pentagonal, disc plates coarse, no obvious pri-

mary plates, Radial shields small, D-shaped,

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 201

divergent proximally. Few to many small pointed

spines on dorsal disc surface. Oral shields

diamond-shaped, as wide as long; adoral plates

contiguous interradially. One triradiate apical

papilla, 2-3 oral papillae. Dorsal arm plate

broadly contiguous in larger specimens, rounded

distally. Ventral arm plates contiguous, as wide as

long, pentagonal with straight distal edge and

tapered proximal end. 4 arms spines, upper spine

only marginally larger than other; lower spines

with swollen thorny tip. Tentacle scale to half the

length of ventral arm plate.

Reproduction. The Macquarie material includes

specimens asexually reproducing through fission.

Division appears restricted to small specimens

under 2 mm d.d.. Division fragments have a disk

half and three arms. The other half of the disk and

three more arms regrow along the division line. It

is possible that one specimen has divided twice,

having one large, two medium and three small

arms. Larger specimens usually have six equal

arms. One aberrant specimen has five equal arms,

but is otherwise identical, A common disc

anomaly is a single large radial shield rather than

two smaller plates on a regenerated disc.

Colour. White in alcohol.

Habitat. Australian specimens are known from

rock and sand substrates (O’Hara, unpublished

data).

Distribution. Northern Macquarie Ridge

(124-155 m); New Zealand and the Chatham

Rise (222-688 m); southeastern Australia

(160-770 m).

Remarks. These specimens are broadly similar to

specimens from southeastern Australia. They dif-

fer, however, in not having an obviously enlarged

upper arm spine, having thorny, swollen tips to

the lower arm spines, and slightly larger tentacle

scales. Fissiparity is rare in the Southern Ocean.

Of the fissiparous species listed by Emson and

Wilkie (1980), only the asteroid, Allostichaster

capensis (Perrier, 1875) is known from south of

40°S, occurring along the southern coast of South

America.

Ophiuridae

Ophiura (Ophiura) meridionalis (Lyman)

Plate 3g

Ophioglypha meridionalis Lyman, 1879: 56, pl. xvi

figs 447-449; Lyman, 1882: 40.

Ophiura meridionalis — Mortensen, 1936; 330-332,

figs 44a-c, 45 — Madsen, 1967: 131.

Material examined. Macquarie Island, off Lusitania

Bay, 54°42.5°S, 158°54,5'E, 69 m, 5 Dec 1930,

BANZARE stn 83, SAM K1097(4). ANARE 1986

Expedition, BT3, NMV F60297(2).

Description of Macquarie Island material. Disc

3.5-5.0 mm d.d., arms 3 times d.d.. Disc plates

thin, imbricating, primary plates and 2 interradial

plates large, other plates small, 0.4 mm dia., 5

from centre to margin. Radial shields small, one-

sixth d.d., contiguous. Arm comb papillae small

flat rounded, usually contiguous over arm. Ven-

trally 1, rarely 2, large plates on margin, 6-7

smaller plates. 1-2 pointed apical papillae, 3—4

smaller rounded oral papillae. Distal oral tentacle

pore with 3—5 inner, 4-5 outer tentacle scales.

Oral shields large, as wide as long, distal sides

rounded, proximally pointed. Adorals long and

bar-like, meeting interradially, separated radially

by large first ventral arm plate. Dorsal arm plates

fan-shaped, longer than wide, contiguous proxi-

mally, separated distally. Ventral arm plates twice

as wide as long, separated throughout. 3 short

subequal conical arm spines, 0.3 mm long, third

segment long. First tentacle pore with 3 scales, |

outer and 2 inner, second with 1-2 scales, inner

scale small and rim-like, other pores with | scale

incompletely covering pore.

Reproduction. This species is viviparous and a

hermaphrodite; male and female gonad separate;

eggs 0.3-0.4 mm dia, 6-8 young in a bursa

(Mortensen, 1936).

Colour. White in alcohol,

Distribution, Off eastern South America

(1098-1890 m); South Georgia, Falkland Islands

(60-249 m); Antarctica, off Enderby, Kemp and

Princess Elizabeth Lands (193-1266 m);

Macquarie Island (69-135 m).

Remarks. The present specimens closely match

Mortensen’s (1936) description and figures of this

species. O. meridionalis has an unusual distri-

bution. It has been found in the Antarctic and

Magellanic regions, but not at other subantarctic

locations besides Macquarie Island. The record of

O. meridionalis from off the Vestfold Hills, East-

ern Antarctica (Tucker and Burton, 1987, pl. 14,

fig. 30) is incorrect. The specimen (TM H1975) is

an Ophiomusium species.

Ophiura (Ophiuroglypha) irrorata Lyman

Plate 3f

Ophioglypha irrorata Lyman, 1878: 73, pl. 4 figs

106-108.—Lyman, 1882: 47-48, pl. 5 figs 7-9,

Ophiuroglypha irrorata—- Pawson, 1969:

figs 2e-f, 8-13 —MeKnight, 1984: 145,

52-54,

202 T. D. OHARA

Ophiura irrorata— Madsen, 1967: 130.

Ophiura (Ophiuroglypha) irrorata

Paterson, 1985; 123-124, figs 46, 47,

Material examined. Macquarie Island, ANARE 1986

Expedition, BT8, NMV F60298(6). Campbell Island,

58°07'S, 169°13'E, 526 m, NMV F52670(6).

irrorata.

Description of Macquarie Island material. Disc

6-9 mm d.d., arms (with broken tip) 3.5 times d.d.

Disc plates coarse, imbricating; primary plates

prominent; large circular plates to 1.1. mm dia.

exist midway to margin in each interradial and

radial axis; wide oval plates, 0.9 x 1.6 mm, are

present in each interradial margin; other disc

plates small, 0.3-0.6 mm dia. Radial shields an

irregular triangle or rectangle, contiguous dis-

tally, separated proximally by several disc plates,

to 1.8 mm long. Arm comb papillae short,

rounded, not contiguous dorsally over arm, dorsal

papillae longest. Genital slits extend from adoral

shields to dise margin, bordered by small papillae.

Oral shields roughly pentagonal, longer than wide

with acute proximal angle and rounded distal

edge. Adorals small, separated radially by first

ventral arm plate. 3-5 papillae on jaw apex, to 5

squarish papillae on each jaw side, smaller than

apical ones, distalmost papillae rarely widened.

Dorsal arm plates contiguous for two-thirds of

arm, fan-shaped with rounded distal edges, as

wide as long proximally, narrower distally. Ven-

tral arm plates contiguous for first 3-4 plates, first

2-3 plates squarish, fourth plate triangular, others

widely separated, 2-3 times as wide as long.

Second oral tentacle pore opens into jaw slit,

elongate with 5 inner and 8 outer scales. Pores

decrease rapidly in size over the next 6 plates, 4

inner and 5 outer scales on first arm pore, 2 and 3

on next 2 plates, reducing to 1 scale. Small sup-

plementary plate on outer distal corners of arm

segments 1-4. 3 small, peg-like arm spines, to 0.3

mm long, dorsal spine little separated from other

two. Middle spine modified into upturned hook

on distal arm segments,

Reproduction. Unknown.

Colour. Tan in alcohol.

Habitat. Mud (Lyman 1882): rock (Pawson,

1969; McKnight, 1984).

Distribution. Cosmopolitan

Macquarie Island (71-450 m).

(71-5870 m);

Remarks. These specimens are broadly similar to

those described by Pawson (1969) from New

Zealand, except that Pawson’s specimens differ

in having supplementary arm plates that persist

until the arm tip and low widened distalmost oral

papillae. O. irrorata is a very polymorphic

species (Paterson, 1985) and these differences do

not appear significant. Six specimens from off

Campbell Island were examined for comparison

(NMV F52670). These are smaller (to 4.5 mm

d.d.), but similar to the Macquarie specimens and

also lack distal supplementary arm plates after the

fifth arm segment. Pawson (1969) noted that his

specimens could be distinguished from other

O. irrorata material by the coarseness of the disc

plates and may represent a new species. In many

ways the Macquarie Island specimens are similar

to O. mundata (Koehler, 1906) (as described and

figured by Paterson, 1985) from the north

Atlantic. However, O. mundata has no supple-

mentary arm plates. Madsen (1967) noted that

O. irrorata is cosmopolitan in abyssal seas,

extending into the sublittoral in the Southern

Ocean.

Ophioplocus incipiens (Koehler)

Plate 3h

Ophioceres incipiens Koehler, 1922b: 48, pl. 84 figs

1-6, 13-14.—Mortensen, 1936: 307.--H.B. Fell, 1961:

69, pl. la fig. 3, pl. 2a fig. 1, pl. 3 fig. 2, pl. 9 fig. 2.—

Madsen, 1967: 137.—Pawson and Rowe, 1977: 352.—

Baker and Devaney, 1981, figs 3, 6-9.

Ophioplocus incipiens— Thomas, 1975: 239-240.

Material examined. ANARE 1986 Expedition, BT6,

NMV F60299(1).

Description of Macquarie Island material. Disc 9

mm d.d., arms > 35 mm. Dise scales small, 0.25

mm dia, imbricating, 15 scales from centre to

margin, primary plates evident. Radial shields

small, 0.8 mm long, oval, widely separated. Ven-

tral disc plates smaller, genital slit 1 arm segment

long. 4 oral papillae and 1 underlying oral ten-

tacle scale on jaw edge. Trapezoid oral shields,

adorals meet interradially, separated radially by

small first ventral arm plate. Dorsal arm plates

fragmented, proximally into 2 large outer frag-

ments separated by 2-3 smaller central fragments,

distally the 2 larger fragments joined at the distal

end and 2-3 smaller fragments are confined to

proximal end. Ventral arm plates pentagonal, con-

vex distally. 2, sometimes 3, stout flattened

pointed arm spines, lowermost longest, half seg-

ment long, 0.6 mm long. 2 tentacle scales, large

oval scale on lateral arm plate, smaller rim-like

scale on ventral arm plate.

Reproduction, Protandric

ep hermaphrodite,

viviparous (Mortensen, 1936).

Colour. Macquarie specimens (in alcohol) tan.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 203

H.B. Fell (1961) recorded live colour as blue-grey

or purple disc with cream or pinkish-yellow arms.

Habitat. Polyzoa (H.B. Fell, 1961)

Distribution. Antarctica, off Enderby, Mac-

Robertson, Princess Elizabeth, King George V

Lands, Ross Sea (110-603 m); South Georgia,

South Shetland, Clarence Islands (60-342 m);

Macquarie Island (25—29 m).

Remarks. The discovery of O. incipiens at

Macquarie Island is surprising as it has been pre-

viously reported only from Antarctic waters. A

closely related species is found in the subantarctic

islands of New Zealand. This was originally

referred to O. huttoni (Farquhar, 1899) but Baker

and Devaney (1981) restricted the known distri-

bution of O. huttoni to the northern island of New

Zealand, recognising O. marginata (H.B. Fell,

1953) as the southern and subantarctic New

Zealand species. Baker and Devaney (1981) dis-

tinguished O. marginata from O. incipiens by the

shorter arms, only twice d.d., and the relatively

simple pattern of dorsal arm plate fragmentation.

Ophioleuce regulare (Koehler)

Plate 3i-j

Ophiopyren regulare Koehler, 1901: 26, pl. VIII figs

52-54.

Ophiopyren _regularis— Madsen,

McKnight, 1984: 145.

Ophioleuce regulare— Madsen, 1983: 45-48, figs

7a-g [full synonymy].—O’ Hara, 1990: 293-294,

1967: 129,

Material examined. Macquarie Island, off Lusitania

Bay, 54°42.5'S, 158°54.5'E, 69 m, 5 Dec 1930, BAN-

ZARE stn 83, SAM(13). ANARE 1986 Expedition,

BT4, NMV F60314(1).

Description of Macquarie Island material.

1.5-5.0 mm d.d., arms delicate, slender, mostly

broken, approx. twice d.d., triangular in cross sec-

tion. Disc slightly convex dorsally, ventrally flat

or concave; plates large, 0.6 mm long, mostly

bordered by 1-4, usually 2, rows of spherical

granules; margin tapers to sharp edge, bears

several rows of elongate pointed granules. No

granules ventrally. Oral shields pentagonal, as

wide as long, distal edge overlain by ventral disc

plates. Adoral shields contiguous interradially,

separated radially by ventral arm plates. 1 elon-

gate pointed apical papilla, 5-7 oral papillae,

inner elongate, others low flat rounded, distal 1-2

papillae function as outer scales of second oral

tentacle pore, opposed by 1-2 inner scales on first

ventral arm plate. Dorsal arm plates with convex

distal edge, contiguous proximally, some slightly

carinate. Ventral arm plates rhombic or triangular,

contiguous until just outside disc margin. 2 small

pointed arm spines, Proximal tentacle pores elon-

gate with 1-3 inner and 1-2 outer scales, distal

pores round with 1 flat oval scale.

Reproduction. Sexes separate (Mortensen, 1936).

Colour. White in alcohol.

Habitat. Mortensen (1936) speculated that this

species lives attached to stones or other hard

substrates.

Distribution. Circumpolar Antarctica (100-

900 m); South Georgia (160 m); Macquarie

Island (65-438 m); southeastern Australia

(770-841 m).

Remarks. The present specimens closely match

the full description given by Madsen (1983).

Class Echinoidea

Key to Macquarie Island Echinoidea

l. Primary spines numerous, secondary spines similar but shorter and evenly

distributed over the test … en Pseudechinus novaezealandiae

— Primary spines few and very large, secondary spines much smaller,

distributed in wreathes around the primary spines and in rows up the

AMBUIACTAE pores eee et ee tete Adnan thy iat BPA win ON T end

2e Apical primary spines with prominent basal and terminal discs

AED, E TA reen oa A N EARTE Goniocidaris parasol?

— Apical primary spines with small terminal cup-like discs and basal spurs .....

wssn rn ERLE Goniocidaris umbraculum®

Notes on key:

4 Two cidaroid echinoids have also been reported from near Macquarie Island. Pawson (1968a:

13-15, fig. 1.1) and McKnight (1984: 145) recorded Goniocidaris umbraculum (Hutton,

1879a) from the northern Macquarie Ridge (NZOI stns D17 and D18, 52°31°S, 160°31/E,

124-128 m, 23 Apr 1963). F.J. Fell (1976) recorded Goniocidaris parasol H.B. Fell, 1958

204 T. D. O'HARA

from Eltanin Cruise 16, stn 1411 also on the northern Macquarie Ridge (51°00’S, 162°01/E.

333-371 m, 8 Feb 1965). G. parasol is distinguished from G. umbraculum by the presence of

basal and terminal discs on the apical primary spines. On the holotype of G. parasol (see H.B.

Fell, 1958: 32-34, pl. 3 fig. b, pl. 5 fig. b) these discs are very large and join together to form

a complete shielding system over the aboral surface. However, F.J. Fell (1976) found that not

all specimens have terminal discs and that basal discs do not develop until a horizontal dia-

meter of 20 mm is reached. As Pawson’s (1968a) specimens were small, < 19 mm diameter

(McKnight supplied no data for his specimens), it cannot be determined if one or two species

of Goniocidaris occur around Macquarie Island. F.J. Fell (1976) also records both species and

the similar species “Austrocidaris” pawsoni McKnight, 1974 and Ogmocidaris benhami

Mortensen, 1921 from the subantarctic islands of New Zealand. Both G. parasol and

G. umbraculum are known to brood their young (Barker, 1984).

Order Temnopleuroida

Temnopleuridae

Pseudechinus novaezealandiae (Mortensen)

Plate 4a—b

Notechinus novaezealandiae Mortensen, 1921: 153,

pl. 6 figs 7-10, pl. 7 figs 4, 5, 7-11 — Koehler, 1926:

36, pl. 54.

Pseudechinus novaezealandiae —- Mortensen, 1943:

237, figs 117-118, 120b, 121b, 126c— Pawson, 1968a:

15-16, fig. 1(2)— Bennett, 1971, pl. 56, fig. 5.—

Simpson, 1982: 50.—McKnight, 1984: 145.

Material examined. Macquarie Island, Garden Cove,

1957, NMV F60309(1); 14 Jul 1963, NMV F60310

(1); 5 May 1965, NMV F60311(1); opposite Gadget

Gully, 1 Jul 1962, NMV F60312(1). ANARE 1986

Expedition, BT2, MNV F60304(6); BT3, NMV

F60302(4); BT4, NMV F60305(3); BT6, NMV

F60303(1); BT10, NMV F60301(3). AM 1977-1978

Expedition, Gorilla Head Rock (9 m), MA-148(1);

Tottan Head, Goat Bay (14 m), MA-371(2),

MA-374(1), MA-375(3); Tern Rock Bay (10 m),

MA-37(1); Garden Cove (8-14 m), MA-14(3):

MA-87(4), MA-123(1), MA-124(3), MA-125(3),

MA-127(2), MA-128(11), MA-379(2); Sandy Bay (16

m), MA-241(6); Green Gorge (0.518 m),

MA-245(10), MA-246(1), MA-247(11), MA-248(1),

MA-251(11), MA-267(1), MA-281(1), MA-291(1).

MA-292(1), MA-294(4); Caroline Cove (13 m),

MA-306(3); Aerial Cove (5 m), MA-52(2), MA-107(1).

Eltanin Expedition, 54° 31.0’S, 159° 00,0°E, 110 m, 12

Feb 1965, cruise 34 stn 2215, USNM(1).

Description of Macquarie Island material. Hori-

zontal diameter to 51 mm, 36 mm high. Test

shape hemispherical, oral side flattened, circum-

ference round, slightly sunken perisome. Spines

short, to 7 mm, coarse and dense; not curved at

perisome. Primary ambulacral spines form regu-

lar series, their tubercules sometimes confluent.

Secondary spines, their tubercules half diameter

of primary spines, form longitudinal rows in

median area, and also present among pore pairs.

Interambulacral areas closely covered in spines,

some secondary tubercules almost as wide as

primary ones, forming 1—2 rows inside each pri-

mary series. Apical system small, one-sixth test

diameter, plates closely tuberculate. Ocular 1

insert, other oculars exsert. Periproct oval, anal

Opening posterior. Complete circle of larger

plates border periproct, smaller elongated plates

border anal opening. Perisome naked. 2 types of

globiferous pedicellariae, larger ones with short

robust valves, with 1—3 teeth on each side, smaller

ones with | tooth on each side. Tridentate pedi-

cellariae small inconspicuous, valves with narrow

serrated edges.

Reproduction. Separate sexes, female gonads

large, with numerous small eggs (0.08—0.1 mm),

The larval stage is probably planktonic

(Mortensen, 1921; Simpson, 1982).

Colour. Macquarie Island specimens (live) with

grey, green or brown test, dark green spines, with

white, grey or occasionally violet tips.

Habitat. On Macquarie Island it is found rarely by

shore collectors in deeper rock pools or as beach

drift. Divers (2-20 m) have found it mainly on

sponge-bryozoan mats, sheltered rock surfaces, or

sometimes amongst the holdfasts of Desmarestia

and Macrocystis. Common in dredged samples to

433 m. Pawson (1968a) recorded this species

from rock and sand substrata.

Distribution. South Island of New Zealand

(0-100 m); Auckland, Campbell, Bounty and

Antipodes Islands, Campbell Rise (0-306 m);

Macquarie Island (0.5-433 m).

Remarks. P. novaezealandiae is the most com-

mon echinoid from Macquarie Island as well

as southern New Zealand (Pawson, 1968a).

P. marionis (Mortensen, 1936) from Marion

Island, Kerguelen and Gough Island differs in

having slender white spines, a white or greenish

test, and much larger suranal plates than

P. novaezealandiae. P. magellanicus (Philippi,

1857) from the Magellanic region and Tristan da

Cunha has a red test and spines.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 205

Plate 4. a, Pseudechinus novaezealandiae, lateral view, NMV F60309; b, Pseudechinus novaezealandiae,

lateral view of test, NMV F60312; c, Psolus neozelanicus, dorsal view, NMV F60306; d, Taeniogyrus

dunedinensis, NMV F77775; e, Pseudopsolus macquariensis, lateral view, NMV F76189; f, Trachythyone

macphersonae, dorsolateral view, NMV F76113; g, Pseudocnus laevigatus, dorsolateral view, NMV F76118;

h, Trachythyone nelladana, holotype, dorsal view, NMV F83072; i, Trachythyone nelladana, holotype,

ventral view, NMV F83072.

206

T. D. O'HARA

Class Holothuroidea

Key to Macquarie Island Holothuroidea

1. Tube feet present

= No TUBE LEET raare eenn ares Sea tite Rune nee aD

10 dendritic (branched) tentacles; shallow water and shelf SPECIES noos 3

15-20 disc, quadrilobed or shield shaped tentacles; bathyl species ............. 8

3. Tube feet mostly absent from the dorsal surface (1-2 may be present

anteriorly),-ventral sole present sr stern ep aint aa hy ee le 4

— Tube feet common on dorsal Surface .......cesscsssessessessesssesssesseceereervecsersersesseens 6

4. Dorsal surface covered in thick leathery skin, few ossicles, pedicles in 3

rows on all ventral radii … …… … nn. Pseudopsolus macquariensis

= Dorsal surface covered by imbricating plates or scales. Midventral radius

without tube feet except at each end or rarely in middle … 5

5. Dorsal surface covered in large scales; up to 6.0 mm long, body flattened,

mouth and anus dorsal in position … nnn Psolus neozelanicus

— Dorsal surface covered in imbricating plates (0.3-0.5 mm long); mouth and

anus anterior/posterior in position … … … … Trachythyone nelladana sp. nov.

6. Tube feet scattered on dorsal surface, not restricted to radii …

n ideeen oun spediale ve Ocnus calcareus*

— Cup ossicles absent, plate ossicles “pine-cone” shaped, denticulate end con-

stricted into definite neck. 10 equal sized tentacles ........ Pseudocnus laevigatus

8. Body soft; 15 disc shaped tentacles, dorsal papillae apparently absent; ossi-

cles concave wheels of 2 types, largest with 10 or more spokes and ventral

rods finely spinous along their length … Laetmogone sp.°

— Body firm; 19-20 quadrilobed tentacles; pedicels restricted to double row on

posterior section of midventral radius; few scattered processes on expanded

dorsolateral margin of body; ossicles with 4 subequal elongate processes …

oc daten A AE EE ep enaar N ci Paelopatides ovalis®

— Body flaccid; 20 shield shaped tentacles, tube feet in 2—4 rows on ventral radii,

dorsal surface with 6 rows of papillae; ossicles cruciform plates with 3—4

bifurcated or perforated arms and central pillar ......... Synallactes challengeri®

9, Wheels ossicles present; skin smooth or wrinkled but not papillate … … … …

NN ey ee ot cs Taeniogyrus dunedinensis

— Wheels rare or absent; anterior body wall with papillae … … Taeniogyrus sp.

Notes on key:

® McKnight (1984: 145) recorded the New Zealand species Oenus calcareus ( Dendy, 1896)

(Cucumariidae: Dendrochirotida) from three NZOI stations off Macquarie Island (71-433 m).

I have re-examined holothurians from one of the stations (D10) and found only three species:

Trachythyone macphersonae, Pseudocnus laevigatus, and Taeniogyrus dunedinensis. As there

is no indication with the material whether it includes all McKnight’s specimens, further con-

firmation is required of the existence of an Ocnus species from Macquarie Island. Mortensen

(1925: 335) recorded O. brevidentis (Hutton, 1872) from Macquarie Island but suspected that

the locality label was an error.

A single damaged specimen of an undescribed Laetmogone species (Laetmoginidae: Elasipo-

dida) was collected south of Macquarie Island by the Eltanin expedition (56° 19.2’S, 158°

29'E, 833-842 m, 12 Feb 1965, cruise 16, stn 1422, identified by M. O'Loughlin, USNM

E27630). This species also occurs on the seamounts south of Tasmania (M. O’Loughlin, pers.

comm..) in depths of 1580-1700.

rotida) has been recently trawled near Macquarie Island (54° 46.2’S, 158° 42'E, 930-815 m,

17 Jan 1995, identified by M. O'Loughlin, NMV F80184). This species is otherwise known

from bathyl depths (911-1611 m) off the Andaman Islands in the Indian Ocean and east of

Cape York in the Coral Sea (M. O’Loughlin, pers. comm.).

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 207

d A single damaged specimen of S. challengeri (Théel, 1886) (Synallactidae: Aspidochirotida)

was collected off Macquarie Island by the Eltanin expedition (54° 31.0°S, 159° 00.0'E, 110 m,

12 Feb 1965, cruise 34, stn 2215, identified by M. O'Loughlin, USNM E47579). This species

also occurs off Crozet, Marion and Prince Edward Islands (237-600 m) (Massin, 1992) and off

Order Dendrochirotida

Psolidae

Psolus neozelanicus Mortensen

Plates 4c, 5a—b

Psolus neozelanicus Mortensen, 1925; 362-363, figs

44, 45 — Pawson, 1970: 28.

Psolus antarcticus—Pawson, 1968a: 19-21, fig.

2(1-4).—McKnight, 1984: 145—O’Hara, 1998a: 146.

[non Psolus antarcticus (Philippi, 1857)]

Material examined. Macquarie Island, ANARE 1986

Expedition, BT2, NMV F76117(1); BT3, NMV

F60306(9). Eltanin Expedition, 54° 30,0'S, 158°

59.0'E, 112-124 m, 15 Feb 1967, cruise 27 stn 1974,

USNM E33647(1).

New Zealand, 2 miles east of North Cape, 102 m,

2 Jan 1915, Mortensen, ZMUC(2 syntypes of P.

neozelanicus).

Description of Macquarie Island material, Up to

50 mm long, 40 mm wide, 10 mm high; body

flattened, limpet-like, mouth and anus dorsal,

mouth highest point, tapering on all sides to thin

sharp margin, Tentacles 10, ventral pair smaller,

dendritic. Anus and mouth each with 5 large tri-

angular valves. Dorsal surface covered in large

imbricating scales. Several rows of smaller scales

surround margin, mouth and anus, and lie

scattered between larger scales. No dorsal tube

feet. Conspicuous flattened ventral sole with 2

rings of tube feet, those on outer ring (on margin

of larger specimens) are small and sparse, those

on inner ring (several mm from margin) are larger

and form biserial or zigzag row; few tube feet at

each end of midventral radius on larger speci-

mens. Calcareous ring plates robust with concave

posterior edge and slender anterior process; 1

polian vesicle

Dorsal ossicles large multi-layered perforated

scales up to 6.0 mm long (body length 38 mm), to

2.5 mm long on smaller specimens (body length

12 mm), Scales with several small grain-like pro-

jections on the dorsal surface, 0.15 to 0.25 mm

wide, consisting of the same multi-layered frame-

work as the scale body. No cups. Ventral ossicles

scattered knobbed perforated buttons. Buttons

with 4 large central holes and 4-20 smaller

peripheral holes, up to 30 scattered small

knobs, jagged margins, up to 0.11 mm long

(pl. Sa-b). Tube feet each with typical ventral

ossicles, slightly concave, up to 0.25 mm wide,

with up to 33 perforations. Tentacles with per-

forated plates of 3 types: 1) triangular shaped to

narrow/bent plates with numerous perforations

and denticulate margin, typically 0.16 mm long;

2) flat irregular plates with 4-6 large perfor-

ations, typically 0.12 mm long; 3) smaller finer

plates, square, slightly concave, numerous small

perforations, typically 0.08 mm long (digit

endpieces).

Reproduction. Sexes separate, gonads consisting

of numerous long unbranched tubules, gonopore

dorsal, posterior to mouth; numerous eggs in

female gonad tubules, average size 0,5 mm dia.

Mortensen (1925) reported egg sizes of 0.2 mm in

his small specimens.

Colour. Macquarie Island material (in alcohol)

brown or pale dorsal surface, pale sole.

Habitat. Several specimens (NMV F60306) were

found attached to dead shells of the scallop

Chlamys patagonica delicatula (Hutton); also

recorded from rock (Mortensen, 1925).

Distribution. New Zealand (102 m); Macquarie

Island (91—415 m).

Remarks. The Macquarie Island Psolus material

appears conspecific with P. neozelanicus rather

than P. antarcticus as proposed by Pawson

(1968a). P. neozelanicus is only known from the

two small type specimens from North Cape, New

Zealand (length 7-10 mm). These specimens

have the same grain-like projections on the dorsal

scales as the Macquarie Island material. These

projections possibly function in a similar fashion

to cup ossicles. The types lack the small outer

tube feet, however this is also true of smaller

Macquarie Island specimens (< 13 mm body

length). The larger type specimen has lateral mar-

gins that are coiled ventrally giving the body an

atypical cylindrical shape, probably an artefact of

preservation (Mortensen, 1925). A similar body

form occurs on two small (9-13 mm length)

Macquarie Island specimens (NMV F60306).

P. antarcticus from the Magellanic region is

very similar except it appears to lack the small

grain-like projections on the dorsal scales;

instead its scales have been described as smooth

or very finely granulated (Théel 1886). Pawson

(1968a) distinguished P. neozelanicus from

P. antarcticus by the presence of several rings of

small dorsal scales surrounding the anus and

208 T. D. OHARA

Plate 5. Holothurian ossicles. a-b, Psolus neozelanicus, NMV F60306, ventral knobbed plates; c—d,

Pseudopsolus macquariensis, NMV F76198, dorsal ossicles, showing incipient secondary network in centre

of plates; e-k, Trachythyone nelladana, NMV F77770: e, dorsal plate; f-g, plates from sole; h, dorsal cups,

view from above; i, dorsal cups, view from below; j, dorsal cups, view from side; k, tentacle ossicles;

lo, Trachythyone macphersonae: |, dorsal plate from juvenile, NMV F77772; m, dorsal plate and cup

(viewed from below) from adult, NMV F77773; n, dorsal cup from juvenile (viewed from above),

NMV F77772; o, dorsal plate and cup (viewed from above) from adult, NMV F761 16; pq, Pseudocnus

laevigatus, NMV F76118, body wall plates; r-t, 7% aeniogyrus dunedinensis: r, sigmoid hook and wheels

from body wall, NMV F77775; s, tentacle ossicle with lateral process, NIWA; t, tentacle ossicles, NIWA;

u, Taeniogyrus sp., sigmoid hooks, AM J22904.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 209

greater number of perforations on the sole

ossicles. However, these features vary in the

Macquarie Island material. Both species are

distinguished from related Australasian and

Southern Ocean species by the lack of a contin-

uous row of midventral tube feet and the presence

of only five large anal and oral valves (Pawson,

1968).

Cucumariidae

Pseudopsolus macquariensis (Dendy)

Plate 4e, 5c-d

Psolus macquariensis Dendy, 1896: 41, pl. 7 figs

70-72.

Pseudopsolus macquariensis —Pawson, 1968b:

143-144, fig. 1 [full synonymy].—Pawson, 1970:

38.— Bennett, 1971, pl. 32.—Simpson, 1982: 45—48,

figs 7-9.

Material examined. Macquarie Island, Secluded Beach,

27 Nov 1989, NMV F76189(3); Garden Cove, 5 Dec

1960, NMV F45003(16); 12 Aug 1962, NMV

F76190(3); 17 Jan 1964, NMV F76200(10); 16 Jun

1967, NMV F76191(2); 14 Mar 1969, coll R.D. Simp-

son, AM 18177(12); 25 Nov 1989, NMV F76192(9); 26

Nov 1989, NMV F76193(6); Buckles Bay, 7 Dec 1986,

NMV F60307(3 juv); 8 Dec 1986, NMV F60308(2);

Nugget Point, 15 Dee 1962, NMV F76194(1); Hurd

Point, 29 Jan 1964, NMV F76195(1); Bauer Bay, 27

Jan 1967, NMV F76196(3); Hasselborough Bay, 23

Aug 1962, NMV F76197(8); Hasselborough Point, 24

Jan 1964, NMV F76199(2): Aerial Cove, 28 Nov 1989,

NMV F76198(5). AM 1977-1978 Expedition, Tern

Rock Bay (7 m), MA-35(1); Garden Cove (1-10 m),

MA-14(5), MA-356(33), MA-360(69); NE Buckles

Bay (intertidal), MA-93(100); Green Gorge (inter-

tidal-0.7 m), MA-255(1), MA-256(2), MA-258(2),

MA-259(2), MA-281(1), MA-282(5).

Description of Macquarie Island material. Up to

40 mm long and 16 mm wide (AM J8177), body

barrel-shaped, with a tapered anus (unless. fully

contracted), anus and mouth slightly dorsally

upturned. 10 equal tentacles, dendritic, each with

4 major bushes on pronounced stalk. Mouth in

centre of large oral disc. Introvert thick and mus-

cular. Dorsal body wall covered in thick smooth

leathery skin, dorsal radii sometimes visible. Con-

spicuous ventral sole, skin notably thinner than on

dorsum. Dorsal tube feet restricted to 1-2 small

ones on introvert. Ventrolateral radii with 1-3

rows of tube feet, outer row largest. Midventral

radii with 1—2 rows of pedicles, often tube feet in

each row placed alternately, in zigzag arrange-

ment. Pair of knotty protuberances are present

midventrally on specimens collected in late

winter-spring.

Dorsal ossicles perforated plates, varying in

density from scarce to abundant and overlapping,

typically 0.08-0.14 mm long, with 24 large

central perforations and smaller peripheral per-

forations, small knobs, sometimes coalescing into

secondary layers, margins denticulate (pl. 5c-d).

No tube foot or tentacle ossicles,

Reproduction. Macquarie Island material usually

with separate sexes, but with some specimens

showing successive hermaphroditism. Two com-

partmentalised brood sacs are present with an

opening to the exterior. Large eggs (to 1.8 mm

dia) develop in ovaries from December to June. In

May-June, eggs are transferred to the brood sacs,

possibly by an external process. Release of

juveniles occurs in late September—October,

apparently in a highly synchronised fashion

(Simpson, 1982).

Colour. Macquarie Island specimens (live) with

dorsal surface coloured light to dark brown, red-

dish or purplish; introvert, tentacles, anal cone

and sole sometimes paler. Brown spots some-

times present between tentacles,

Habitat. Lower intertidal pools and rock surfaces

(0-1 m), subject to wave action, sometimes

exposed at low tide; amongst Durvillaea hold-

fasts, sometimes on coralline algae or Codium.

Only two of the current 295 specimens were

collected from subtidal depths (7 and 14 m).

Distribution. Macquarie Island (0-14 m).

Remarks. This species is very common on rocky

substrates in the lower intertidal zone, The sole is

used to adhere strongly to the substrate and the

tentacles are extended into the surf. Pawson

(1968b) described internal anatomy in detail.

Simpson (1982) noted the pair of protuberances

which appear on the ventral body surface, near the

openings to the brood sacs, on specimens col-

lected during the juvenile release period in spring.

In this collection they are present in specimens

collected as early as August. Their function is

unknown.

P. macquariensis gruai has been described

from Kerguelen by Cherbonnier and Guille

(1975). Kerguelen specimens are similar to Mac-

quarie specimens in overall morphology and

habit. They appear to differ in having ossicles in

the tube feet and tentacles, and possibly in their

developmental biology. Cherbonnier and Guille

(1975) described the gonads as being very long

and fine, with relatively few eggs measuring

lum in diameter. A record of P. macquariensis

from Stewart Island south of New Zealand is

210 T. D. OHARA

probably a locality error (Mortensen, 1925;

Pawson, 1970).

Pseudocnus laevigatus (Verrill)

Plates 4g, Sp—q

Pentactella laevigatus Verrill, 1876: 68.

Cucumaria laevigata— Ludwig and Heding, 1935:

179-185, fig. 43.

Pseudocnus laevigatus.—Pawson, 1968b: 145, figs

2—11 [full synonymy].—Bennett, 1971, pl. 56 fig. 6a.—

Simpson, 1982: 48,

Pseudocnus leoninoides.—Pawson, 1968a: 22. [non

P. leoninoides (Mortensen, 1925)]

Material examined. Macquarie Island, Garden Cove, 26

Dec 1965, I. Bennett, det. Pawson, F45102(1); 26 Nov

1989, NMV F76119(1); off Buckles Bay, 54°30’S

158°58'E, 15 m, 19 Oct 1983, NMV F82953(8); Acrial

Cove, 28 Nov 1989, NMV F76118(1); 54°40'S,

159°01'E, 71 m, NZOI stn DIO, NIWA (3). AM

1977-1978 Expedition, Gorilla Head Rock (11-12 m),

MA-146(2); Tottan Head, Goat Bay (24-14 m),

MA-39(1), MA-41(1), MA-365(2), MA-371(9),

MA-374(11), MA-375(9), MA-376(16); Tern Rock

Bay (7 m), MA-35(6); Garden Cove (6-14 m),

MA-14(2), MA-91(4), MA-123(1) MA-124(8),

MA-125(23), MA-127(5), MA-128(18), MA-350(1),

MA-379(19); Sandy Bay (15-16 m), MA-237(2),

MA-241(57); Green Gorge (6-18 m), MA-245(7),

MA-247(8), MA-248(3), MA-250(4), MA-251(3),

MA-267(1), MA-275(3), MA-289(1), MA-291(1),

MA-292(13), MA-294(7); Caroline Cove (8-18 m),

MA-298(7), MA-300(2), MA-306(3), MA-307(2),

MA-311(2); Handspike Point (intertidal), MA-136(2);

Aerial Cove (0-6 m), MA-46(2), MA-47(14),

MA-50(8), _ _MA-54(2), MA-55(1), MA-83(12),

MA-85(65). MA-86(1), MA-98(1), MA-102(35),

MA-107(54), MA-108(100), MA-109(12), MA-110

(30), MA-382(10), MA-383(1), MA-386(155), MA-

388(4), MA-389(15); Anchor Rock (13-20 m), MA-

89(21), MA-95(6). ANARE 1986 Expedition, BT2,

NMV F76120(5); BT3, NMV F76121(14); BT4, NMV

F76122(3); BTS. NMV F76123(2). Eltanin Expedition,

54°24'S, 159°01'E, 10 Feb 1965, 79-93 m, cruise 16,

stn 1417, USNM E43134(5); 54°31.0’S, 159°00.0'E,

18 Jun 1968, 110 m, cruise 34, stn 2215, USNM(1).

Description of Macquarie Island material. Body

cylindrical, up to 35 mm long (preserved). Pre-

served material often highly contracted, with

body wall becoming thick and wrinkled or body

becoming twisted. 10 equal tentacles. Introvert

very long and thin when extended. Ventral and

dorsal body surface similar, no sole. Pedicels

restricted to radii, in 1-2 zigzag rows, extending

from anus to anterior end, including across intro-

vert. Body wall dense with small pine-cone

shaped ossicles (pl. 5p-q). Ossicles 0.08-0.15

mm long, 0.075-0.14 mm wide, with 4-5 large

central and smaller peripheral holes, covered in

large knobs, one end attenuated and denticulate,

Denticulate end orientated externally giving body

surface a rough texture. Some rare x-shaped plate

precursors, 0.03 mm long. Plates slightly smaller

ventrally. Introvert ossicles irregular perforated

plates, few knobs. Tube feet plates narrow and

bent, widened centrally and attenuated at each

end, large perforation in centre; end pieces

0.15-0.27 mm dia. Tentacle ossicles are 2 kinds

of perforated plates, irregular shapes, large plates

typically 0.2 mm long, occasionally knobbed,

smaller plates finer, 0.7 mm long.

Reproduction. Sexes separate. Gonads short

unbranched tubules. Females with few eggs in

each gonad; Macquarie Island specimens with

eggs in November to 0.7 mm dia; Kerguelen

specimens with eggs in January to 1.4 mm dia

(Pawson, 1968b). Eggs transferred to brooding

sacs in November. A pair of brooding sacs are

attached on either side of the ventral midline in

the posterior half of the animal. The sacs are

invaginations of the body wall opening to the

exterior through a vestibule with a slit-like aper-

ture (Pawson, 1968b). Brood juveniles in Decem-

ber measure 3 mm long, with a few tube feet on

the radii, visible tentacles and scattered ossicles in

the body wall. Simpson (1982) found up to 93

young in the internal brood pouches of a speci-

men 35 mm long from Macquarie Island (no date

supplied). Small juveniles are present in samples

from December to February, which may have

been free living or expelled from the brood sacs

during collection of adults,

Colour. Macquarie Island material (live) light

pink to orange-pink with distinct brown dots on

the tentacle trunks and at the bases of the pinnae.

Marion Island specimens white or orange

(Pawson, 1971).

Habitat. Macquarie Island: sublittoral, under

rocks or sheltering between encrusting sponges or

the green alga Codium, within Macrocystis hold-

fasts, and rarely on sand. Can be locally numerous

under sheltered rock overhangs. Marion Island

specimens recorded from rock, gravel, sand and

sand with mud (Branch et al., 1993).

Distribution. Marion Island (0.5—240 m); Crozet,

Kerguelen, Heard Islands (0.5-275 m); Mac-

quarie Island (0.5—135 m). Théel (1886) recorded

one small specimen from 1023 m south of Crozet

Island (as Cucumaria serrata var. marionensis).

Remarks. P. laevigatus is a polymorphic species

or species complex that has been reported widely

throughout the subantarctic. The shape of the

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 211

ossicles can vary from the typical “pine-conc”

shape, present at Macquarie Island, to elon-

gated plates found on animals near Heard Island,

to being very irregular at Kerguelen (M.

O'Loughlin, pers. comm.). Théel (1886)

attempted to separate regional varieties of this

species (as Cucumaria serrata) but these have

been abandonned by later researchers. However,

similar forms from South America and New

Zealand are known as P. perrieri (Ekman,

1927) and P, leoninoides (Mortensen, 1925)

respectively.

Pawson (1968a) recorded a single specimen of

P, leoninoides from NZOI station D18, 150 miles

NNE of Macquarie Island (52°31'S, 160°31/E,

128 m, 24 Mar 1963, NIWA). Pawson (1970)

suggest that further material may reveal that the

Macquarie specimen represents a different

species. M. O'Loughlin (pers. comm.) who has

examined this specimen believes its affinities lie

more with P. /aevigatus than P. leoninoides.

P. leoninoides is known from the rocky shallows

of Campbell, Auckland and the Snares Islands

and apparently differs from P. laevigatus in lack-

ing brood pouches (Mortensen, 1925). Material

referred to both P. laevigatus and P. leoninoides

requires revision.

Trachythyone macphersonae Pawson

Plates 4f, 51-0

Trachythyone macphersonae Pawson, 1962: 47, pl. 1

figs 1l-5.—Pawson, 1968b: 147-149, figs 12, 13.—

Bennett, 1971, pl. 56 fig. 6b (upper figure is probably

P. laevigatus)—Simpson, 1982: 48,

Material examined. Macquarie Island, Garden Cove, 26

Dee 1959, NMV F45001(holotype); Dec 1963, NMV

F76116(4); 12 Aug 1962, NMV F76115(4); 25 Nov

1989, NMV F76li4(1); 26 Nov 1989. NMV

F76113(3); north coast, under rock in pool, Dec 1965,

l. Bennett, ident. Pawson, NMV F45104(6); Buckles

Bay, 8 Dec 1986, NMV F77777(1); off Buckles Bay,

54°30°S, 158°58'E, 15 m, 19 Oct 1983, NMV

F82952(25); Aerial Cove, 28 Nov 1989, NMV

F76110(5); 54°40'S, 159°0 1'E, 71 m, NZOI stn D10,

NIWA(7). AM 1977-1978 Expedition, Tottan Head,

Goat Bay (9-14 m), MA-369(1), MA-371(2).

MA-376(2); Tern Rock Bay (1-10 m), MA-34(1),

MA-35(6); Garden Cove (3-14 m), MA-92(5),

MA-350(2), MA-354(18), MA-379(8); Sandy Bay

(15-16 m), MA-237(5), 241(45), 242(4); Green Gorge

(0.5-18 m), MA-245(4), MA-247(2), MA-248(7),

MA-251(8), MA-267(2), MA-281(6), MA-292(1),

MA-294(13); Caroline Cove (8-18 m), MA-300(4),

MA-306(8), MA-307(1), MA-311(2); Handspike Point

(intertidal), 136(4); Aerial Cove (0.5-6 m), MA-46(2),

MA-47(8), MA-50(3), MA-83(13), MA-85(2),

MA-98(1), MA-102(106), MA-108(26), MA-109(1),

MA-110(10), MA-382(14), MA-386(261), MA-387(4),

MA-388(18), MA-389(29): Anchor Rock (20 m),

MA-89(13). ANARE 1986 Expedition, BT2, NMV

F77773(6),. BT3, NMV F77771(1), BT4, NMV

F77772(3).

Description of material. Body (preserved) up to

65 mm long, 10 mm wide (AM stn MA-50). Body

cylindrical, 10 short dendritic tentacles, orien-

tated anteriorly, ventral pair reduced. Anus on

small posterior cone or taper, surrounded by 5

small valves and some small tube feet. Scattered

dorsal tube feet. Ventral tube feet slightly larger,

restricted to radii, in 2 rows midventrally, and 1-2

rows ventrolaterally, rarely with 3 rows. Ventral

skin thinner, distinct, no true sole. No tube feet on

introvert. Juveniles (3 mm d.d.) with 1-2 tube feet

on each ventral radius, few smaller tube feet

scattered dorsally.

Dorsal ossicles massed flat perforated plates,

overlain by cups. Small specimens have large

plates to 0.3 mm long (rarely to 0.37 mm) with

numerous knobs, up to 35 holes (pl. 51). Knobs

occasionally coalesce into incipient secondary

layer. Plates often overlap to form rigid crys-

talline body wall. Large specimens with smaller

plates, 0.06 to 0.23 mm long, with 2-13 holes,

few small low knobs (pl. 5m, 0). A few flat x-

shaped ossicles present, possibly plate precursors,

Cups oval-shaped with smooth cruciform base

and spined rim, spines directed up away from

cruciform piece (pl. Sm-o). Rim formed by

coalescing ends of developing cruciform piece,

typically to 0.05 mm long and 0.04 mm wide,

slightly wider than deep. Cruciform pieces with-

out rims or cups with incomplete rims sometimes

found. Cups more numerous in adults. Ventral

ossicles plates and cups, plates smaller, finer,

more irregular than dorsal plates, 0.15 to 0.25 mm

long. Introvert ossicles small thin plates with fine

knobs and large perforations, to 0.25 mm long

and 0.1 mm wide. Tube feet with endplates (0.3

mm dia) surrounded by a series of irregular nar-

row, bent or triangular perforated plates, typically

0.27 mm long 0.07 mm wide, sometimes with

ends or | edge strongly denticulate. Tentacles

with elongate curved knobbed perforated plates,

typically 0.2 mm long and 0.04 mm wide, and

smaller irregular perforated plates with large

holes and few knobs, typically 0.07 mm long and

0.05 mm wide.

Reproduction. Sexes separate. Gonads a cluster of

short tubules. Females with few relatively large

eggs, typically 0.2 to 0.35 mm dia (Simpson,

1982 recorded eggs up to 0.8 mm dia.), eggs

evident in specimens from all months sampled

212 T. D. OHARA

(July to February), no evidence of brooding in the

current specimens.

Colour. Body and introvert (live) pink or mauve;

anal cone and tube feet light orange; tentacles

dark orange with distinct brown dots at tentacle

bases.

Habitat. Sublittoral, under rocks or sheltering

amongst encrusting sponges, anemones or the

green alga Codium, common within Macrocystis

holdfasts. Locally numerous under sheltered rock

overhangs.

Distribution. Macquarie Island (0.5—135 m).

Remarks. Many species of Trachythyone

described from the Southern Ocean are ill-defined

and require revision. The ossicles of the various

species can change with age and some ossicle

types may be missing from old or poorly pre-

served specimens. 7. macphersonae is similar to

material of 7. parva deseribed by Panning (1964)

and Hernandez (1982) from off Chile and the

Falkland Islands (0-180 m) and may be con-

specific. Both species have knobbed plates and

cups, with large plates (pl. 51) and few cups in

juveniles and smaller plates (pl. Sm, o) overlain

by numerous cups in adults. Whether these speci-

mens are conspecific with the types of T. parva is

uncertain as Ludwig’s (1875) inadequate descrip-

tion of the ossicles in the type translates as “large

perforated plates” and “small x-shaped cups”.

Some specimens of T. macphersonae have a few

incomplete x-shaped cups consisting only of the

concave cruciform piece. Specimens from Ker-

guelen, originally recorded as T. parva by early

workers, were redescribed as 7. ekmani by

Ludwig and Heding (1935) on the basis that they

possessed cups instead of x-shaped plates. On the

other hand Cherbonnier and Guille (1975) sug-

gested that 7. ekmani is no more than a ecological

race of T. parva. Possibly T. parva, T. ekmani and

T. macphersonae represent a single widely-

dispersed subantarctic species. A revision will

require fresh material of several size classes

collected from throughout the subantarctic.

Trachythyone nelladana sp. nov.

Plates 4h—1, Se-k, Figure 5a

Material examined, Holotype, Macquarie Island,

ANARE 1986 Expedition, off Nugget Point, 54°33.4’S,

158°56.9'E, 108-135 m, 8 Dec 1986, stn BT3, NMV

F83072.

Paratypes, type locality and date, NMV F77770(10).

Description. Holotype 10 mm long, 4 mm wide, 3

mm high. Body dome shaped with convex dorsal

Figure 5. a, Trachythyone nelladana, NMV F77770,

radial and interradial plate of calcareous ring.

b, Taeniogyrus sp., AM J22904, tentacle ossicle

fragment.

surface and flattened sole. Mouth orientated ante-

riorly, 10 dendritic tentacles, ventral pair reduced,

Small thin introvert. Posteriorly orientated anus.

No dorsal tube feet, except pair of minute ones at

anterior margin. Sole present, surrounded by ring

of about 25 large tube feet, tube feet denser near

anterior and posterior ends, isolated midventral

tube foot occasionally present, skin of sole very

thin. No obyious anal or oral valves. Calcareous

ring with 5 radial and 5 interradial pieces, each

with incised anterior projection and posterior

notch; radials larger with smaller anterior inci-

sion.

Dorsal ossicles consist of several layers of

overlapping perforated plates overlain by cups.

Plates up to 0.55 mm, approximately 30 plates

between anus and introvert on largest specimen,

up to 50 perforations per plate in regular arrange-

ment (pl. 5e), no knobs, some larger plates with

ventral thickenings or arches. Cups oval, with

thick cruciform piece and spinose rim, typically

0.05 mm long, 0.04 mm wide, 0.04 mm deep, rim

often incomplete (pl. Sh—j). Ventral ossicles small

buttons with 2-8 large holes, margin with small

rounded knobs, up to 0.1 mm dia (pl. 5f-g). Tube

foot ossicles elongate perforated rods surrounding

end plate, end plate 0.3 mm dia. Tentacle ossicles

flat perforated plates, quadrangular, typically with

10 perforations and denticulate margin, to 0.12

mm; some plates slightly concave; other plates

elongate, 3 times as long as wide, with larger per-

forations (pl. 5k).

Paratypes 4—11 mm in length, similar morphol-

ogy and ossicles as holotype.

Colour, Tan/white, pale (preserved).

Reproduction. Female caeca in December with

long slender unbranched moniliform tubules (one

half the body length) containing scattered ovoid

eggs 0.15 to 0.25 mm long.

Habitat. Unknown.

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 213

Etymology, Latinised form of the name of the

former polar vessel Nella Dan from which these

specimens were collected.

Distribution. Macquarie Island (108—135 m).

Remarks. These specimens are intermediate

between the Psolidae and Cucumariidae having

the clearly defined sole of a psolid (thin skin,

mainly peripheral tube feet and small ventral

ossicles) and the small single-layered plates, well-

developed cups and terminal mouth/anus of a

cucumariid. Both families contain species that

lack dorsal tube feet. I am placing this species in

the Cucumariidae on the basis of the ossicles

present. However, the Cucumariidae is possibly

paraphyletic, the psolids representing an extreme

development of cucumariids with imbricating

dorsal plates.

I am reluctant to create a new genus for

these animals as the Cucumariidae is badly in

need of revision and generic limits are unclear

(O'Loughlin and O’ Hara, 1992). The ossicles. flat

imbricating plates and cups, are similar to those

found in Trachythyone species. It is clearly dis-

tinguished from all other Trachythvone species in

lacking dorsal pedicels. Species within the

Cucumariidae that lack dorsal tube feet

include Ocnus farguhari Mortensen, 1925 and

O. sacculus Pawson, 1983 from New Zealand.

These species differ from the 7. nelladana in

having multi-layered ossicles, knobbed buttons

and rows of tube feet on all three ventral radii.

Other cucumariid genera that lack dorsal tube feet

include Microchoerus Gutt, 1990 which has small

knobbed plates that diminish with age and

lacks cups; Neocnus Cherbonnier, 1962 and

Pseudopsolus Ludwig, 1898 which have few

body wall ossicles.

The new species can be differentiated from

species of Psolus by the terminal mouth and anus,

the flat single-layered dorsal plates, the well-

developed cup ossicles (absent in the sole), and

the flat reduced ventral plates. Several other

species have a similar body form, including

P. charcoti Vaney, 1906, P. murravi Théel, 1886,

and P. granulosus Vaney, 1906 from the Southern

Ocean. The body shape of P. granulosus is most

similar to the current species. However, this

species has multi-layered dorsal scales, knobbed

ventral plates and lacks true cups. Both

P. charcoti and P. murrayi are large cylindrical

animals with a small sole and a thick epidermis

that obscures the multilayered dorsal scales. Few

Psolus species have true cup ossicles, their

functional role often being replaced by multi-

layered grains (e.g., P. neozelanicus) or concave

spinous plates. Psolus ephipiffer Thomson,

1876 and P. paradubiosus Carriol and Féral,

1985 both from the Kerguelen region, have

shallow cup-shaped ossicles, but differ from the

current species in their body form, large dorsal

scales and the knobbed ventral plates. P. koehleri

Vaney, 1914 from the Antarctic Peninsula has

shallow cups in the sole, but has an upturned oral

cone, the mouth being surrounded by five brood

sacs,

Order Apodida

Chiridotidae

Taeniogyrus dunedinensis (Parker)

Plates 4d, Sr-t

Trochodota dunedinensis Parker, 1881: 418.

Dendy, 1896: 26-28, 49-50, pl. 3 figs 1-8—

Mortensen, 1925: 376-381, figs 59-61.—Pawson,

1968a; 24-25.—Pawson, 1970: 46-47 .—McKnight,

1984: 146.

Taeniogyrus dunedinensis. —Rowe, 1976: 204,

? Trochodota sp. Pawson, 1968b: 149.

Taeniogyrus sp. nov.—O’ Hara, 1998a: 146.

Material examined. Macquarie Island, shore, 1912,

H. Hamilton (AAE), det. Pawson (1968b) as Tro-

chodota sp., AM J4725(2); 54°40'S, 159°01’B, 71 m,

NZOL stn D10, det. Pawson (1968a), NIWA(4).

ANARE 1986 Expedition, BT4, NMV F77775(1).

New Zealand, Stewart Island, Paterson Inlet, 9 m, 12

Jan 1952, NMV F82702(4).

Description of Macquarie Island material. Up to

15 mm long. 10 uniform tentacles, approximately

8 digits per tentacle. Oral disc orientated

obliquely. Skin smooth, not papillate, one speci-

men (BT4) with a few white spots in the anterior

body wall formed from clusters of wheel ossicles.

Calcareous ring plates narrow, quadrangular, with

slight posterior notches.

Body wall ossicles wheels and sigmoid hooks.

Wheels slightly hexagonal, up to 0.09 mm long, 6

spokes, continuous teeth around inner rim, 4 teeth

across spokes, 5 teeth between spokes (pl. 5r). 1

specimen (BT4) with wheels grouped in anterior

clusters, other specimens with sparse scattered

wheels. Sigmoid hooks sparsely scattered, twice

as long as wheels, to 0.16 mm long, smooth. Ten-

tacle rods abundant (pl. St), to 0.12 mm, straight

to curved to bent, not branched laterally, a few

complex lateral knobs (pl. 5s), tips forked with 2

clusters of knobs,

Colour. Macquarie Island material (in alcohol)

light brown. Live Auckland Island material

reddish-brown with dark spots at the base of the

tentacles (Mortensen, 1925).

214 T. D. OHARA

Habitat. Auckland and Campbell Island speci-

mens found under stones at low tide (Mortensen,

1925) and in shell/sand (Pawson, 1968a).

Distribution. New Zealand (1-9 m); Auckland,

Campbell Islands (1-180 m); Macquarie Island

(65-433 m).

Remarks. The present specimens differ from typ-

ical T. dunedinensis material from New Zealand

in the relative size of the wheel and sigmoid hook

ossicles, the wheels being considerably smaller

than the hooks. However, the wheel ossicles are

known to vary in size from 0.053 to 0.16 mm dia.

in other material referred to T. dunedinensis

(Mortensen, 1925). Without a large number of

specimens it is impossible to determine whether

the Macquarie Island material is conspecific with

that from New Zealand.

Pawson (1968b) described two specimens of an

undetermined Trochodota species collected by

the ANARE expedition from the north coast of

Macquarie Island. Unfortunately the ossicles have

decayed, however, other characteristics suggest

that it also belongs to this species. The specimens

are 23 and 30 mm long, 1.5 mm wide, with ten

subequal tentacles, eight digits per tentacle, the

terminal digits are longest. The skin is not papil-

late and the mouth is orientated obliquely. The

anterior half of the body of one specimen is filled

with female gonads, the eggs are 0.3 mm dia.

Two other Taeniogyrus species occur in the

Southern Ocean. T. dendyi (Mortensen, 1925) is

known from northwest of the Auckland Islands

(Pawson, 1968b). It is large in size (to 180 mm

long), purple-white in colour, with 12-14 digits

per tentacle. The skin is covered in papillae con-

taining massed hooks, wheel ossicles are scarce,

and the tentacle rods are small (0.05 mm) with

2-5 branches at each end. T. contortus (Ludwig,

1875) recorded from Kerguelen, has 12 tentacles,

large sigmoid hooks, more than double the size of

the wheel ossicles, and wheels that are grouped

into distinctive papillae.

Taeniogyrus sp.

Plate 5u, Figure 5b

Material examined. Macquarie Island, AM 1977-1978

Expedition, Green Gorge (15 m), MA-289, AM

J22904(3).

Description of Macquarie Island material. Up to

9 mm long (strongly contracted), in poor condi-

tion, with tapered anal cone. Skin with numerous

longitudinal folds from contraction, probably

papillose. Mouth orientated anteriorly, not

oblique, 10 uniform tentacles, approximately 4

digits per tentacle, upper 2 largest. Body wall

ossicles sigmoid hooks. Sigmoid hooks scattered,

| per papillae, typically 0,07 to 0.11 mm long

(pl, Su). No wheels present. Tentacle ossicles

mostly eroded, remaining pieces are rod ends

with 3 terminal denticulations, entire ossicles

probably 0.05 mm long (fig. 5b).

Colour. Tan in alcohol.

Habitat. Coarse sand.

Distribution. Macquarie Island (15 m).

Remarks. Chiridotids with sigmoid hooks but

without wheel ossicles are generally referred to

the genus Scoliorhapis H.L. Clark, 1928. How-

ever, although these small, poorly preserved spec-

imens lack wheel ossicles they appear closer to

Taeniogyrus dendyi (Mortensen, 1925) from New

Zealand than the type species of Scoliorhapis,

S. theeli (Heding, 1928) from southern Australia.

Both species have groups of sigmoid hooks asso-

ciated with skin papillae that are present on the

anterior part of the animal. S. theeli clearly differs

from the Macquarie Island material in having

C-shaped unbranched tentacle ossicles, whereas

wheel ossicles are known to be scarce or even

absent in subantarctic specimens of 7. dendyi

(Mortensen, 1925, Pawson, 1968a). Better pre-

served material is required to finally determine

the identity of these specimens.

Discussion

Habitat. Detailed habitat information is only

available for Macquarie Island echinoderms col-

lected from rocky shores (0-2 m) or by divers

from shallow subtidal waters (0-25 m). The dis-

tribution of echinoderms across habitats at these

depths is shown in Table 3. No quantitative

survey has ever been conducted at subtidal depths

at Macquarie Island. Consequently abundance has

been estimated from museum collections.

1. Sublittoral fringe (0-2 m). The ecology and

zonation of the rocky shores at Macquarie Island

has been described by Kenny and Haysom

(1962), Bennett (1971) and Simpson (1976). The

sublittoral fringe is dominated by the large brown

alga Durvillaea antarctica, This habitat which is

affected by wave surge can range from 0-3 m on

the east coast to as deep as 15 m on the exposed

west coast (Ricker, 1984). Immediately seaward

of Durvillaea, the rocks are covered with foliose

red algae. A conspicuous but species-poor echin-

oderm fauna is present in the upper sublittoral,

occurring under rocks, in crevices and under the

large fronds of Durvillaea. Six echinoderm

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS

Table 3. Known habitat distribution of Macquarie Island echinoderms (0-20 m). No habitat

information is available for Taeniogyrus dunedinensis.

Sublittoral fringe(0-2 m)

Upper sublittoral (2-20 m)

Durvillaea

holdfasts

Species Red algal

zone

Macrocystis| Codium Rock

beds overhangs

Cycethra frigida

beds

P- H

Porania antarctica

Pteraster affinis

Henricia obesa

Smilasterias clarkailsa

Anasterias directa

Anasterias mawsoni

Ophiacantha vilis

Amphiura magellanica

Pseudechinus novaezealandiae

Pseudopsolus macquariensis

Pseudocnus laevigatus

Trachythyone macphersonae

Taeniogyrus sp

vv A] Ul vl | VT

R = rare (all collections with <= 2 specimens); P = present (3—19 specimens); N = locally numerous

(at least one collection with > 20 specimens)

species are generally present, the asteroids

Anasterias mawsoni, A. directa and Cycethra

frigida, and the holothurians Pseudopsolus

macquariensis, Trachythyone macphersonae and

Pseudocnus laevigatus. The first five of these

may be abundant. In particular P. macquariensis

may completely cover rocks at the low tide level.

P. laevigatus is not common in this zone,

2. Upper sublittoral (2-20 m). Habitats at these

depths have been described by Lowry et al.

(1978) and Ricker (1984). The habitats include 1)

exposed rocks and boulders dominated by a

canopy of the kelp Macrocystis pyrifera often

with an understorey of Desmarestia chordalis; 2)

sheltered rock caves and overhangs covered with

sponges, tunicates and hydroids; 3) protected

shallow coves and inlets often with mats of the

green algae Codium subantarcticum; and 4) areas

of coarse sand and gravel. The asteroids and

holothurians A. mawsoni, A. directa and C.

frigida, T. macphersonae and P. laevigatus are

common as in the sublittoral fringe, but P. mac-

quariensis is rarely found below | m. Several

shelf species are also present. The echinoid

Pseudechinus novaezealandiae and the ophiuroid

Ophiacantha vilis may be locally common. The

asteroids Henricia obesa, Smilasterias clarkailsa,

Porania antarctica, Pteraster affinis, the ophi-

uroid Amphiura magellanica and the holothurian

Taeniogyrus sp. also reach their upper limit in this

zone.

3. Deep water (< 20 m). The continental slope

surrounding Macquarie Island is covered by very

little sediment (Williamson, 1988). The slopes are

very steep, the east side plunges over 5000 m

from Macquarie Island to the Macquarie Trench

in a little over 25 km. The total submerged area of

the Macquarie Ridge occurring at depths of less

than 1000 m deep is only 3300 km? (Williams,

1988). Most of the Macquarie Island material

from deeper water has been trawled and lacks

detailed substratum information. The lack of sed-

iment indicates that many species would occur on

rock or amongst sessile invertebrates growing on

rock, although there are some species that are

known from elsewhere to prefer a soft sediment

substratum (e.g., Psilaster charcoti, Taeniogyrus

spp.).

The echinoderm fauna is similar at all shallow

water sites that haye been examined around the

island (Table 2) although relative abundance can

vary with exposure (Bennett, 1971; Ricker,

1984). There are some exceptions. Porania

antarctica and Pteraster affinis have been found

by divers only at Caroline Cove despite the low

level of sampling at the site. Taeniogyrus sp. was

collected only at Green Gorge off a sandy sub-

stratum. Relatively few soft sediment species

216 T. D. OHARA

have been found from 0-20 m. This may be due

to the lack of suitable substratum or the lack of

collecting with dredges and boats within this

depth range (G.C.B. Poore, pers. comm.).

The shallow subtidal environment is broadly

similar at other subantarctic Islands including

Marion Island (Beckley and Branch, 1992;

Branch et al., 1993), Heard Island (Smith and

Simpson, 1985) and Kerguelen (Guille, 1974;

Cherbonnier and Guille, 1975; McClintock,

1985), with the exception that the kelp Mac-

rocystis is apparently absent from Heard Island.

A similar echinoderm fauna is present at all loca-

tions. Anasterias rupicola, Pseudechinus mario-

nis and Pseudocnus laevi-gatus dominate the

echinoderm fauna at Marion Island, although the

asteroid Anteliaster scaber and the ophiuroid

Ophiurolepis intorta can be locally common.

Other shallow water species include Henricia

praestens, Pteraster affinis, Porania antarctica

and Smilasterias scalprifera. There has been no

subtidal SCUBA survey of Heard Island, but

beach washed specimens indicate that the aster-

oids Anasterias mawsoni, Odontaster meridion-

alis, Porania antarctica and Pteraster affinis are

common (O’ Hara, this report). At Kerguelen shal-

low rocky reefs support Anasterias rupicola and

A. perrieri, Henricia spinulifera, Porania antarc-

tica, Pteraster affinis, Cvcethra frigida, Pseudop-

solus macquariensis gruai, Pseudocnus laeviga-

tus and Trachythyone ekmani. Kerguelen differs

from Macquarie Island in having wide sandy bays

which support a significant soft sediment fauna.

Systematics and origin. One of the difficulties

identifying echinoderms from Macquarie Island

is the polymorphism exhibited by many species

(or species complexes) that are widespread

throughout the Southern Ocean. There is consid-

erable morphological variation both between and

within isolated populations.

widespread polymorphic species include the

asteroids Odontaster penicillatus and Cycethra

verrucosa which can vary in form from subpen-

tagonal to stellate with a range of abactinal spina-

tion. In other cases a number of discrete regional

or sibling species are currently recognised, such

as the numerous Anasterias and Henricia species.

The current and past geography of the Southern

Ocean is likely to have promoted regional varia-

tion and polymorphism. The break up of the

ancient continent of Gondwana would have iso-

lated populations of formerly widespread species,

which could then have evolved independently

Examples of

(Palacoaustral species). Examples include Psolus

neozelanicus and P. antarcticus, very similar

species that are now widely separated in the Tas-

man region and off South America respectively.

Overlaying these vicariant patterns are species

(Neoaustral species) that have subsequently dis-

persed eastward across the Southern Ocean by

epiplanktonic rafting on the West Wind Drift

(H.B. Fell, 1962). Infrequent colonisation of iso-

lated islands would promote regional variation

and speciation through founder effects and

genetic drift (Palumbi, 1994). Polymorphism has

possibly been accentuated by a complex process

of colonisation, genetic drift and recolonisation

by parental stock. Dispersal by epiplanktonic raft-

ing is restricted to shallow water rocky reef

species living amongst kelp holdfasts, the only

known transportation vector (O'Hara, 1998a),

Species rafting to Macquarie Island from the west

probably include Anasterias spp., Cvcethra

frigida, Trachythvone macphersonae, Pseudoc-

nus laevigatus and Pseudopsolus macquariensis,

all of which have been found within kelp hold-

fasts on Macquarie Island.

The problem for a taxonomist is whether these

morphological variants are separate species, sub-

species or races. Key (1981) has provided some

relevant definitions. Species are populations that

are reproductively isolated from all others, where

reproductive isolation requires that either no

hybrids are found in the field or that such hybrids

are infertile. A race is any population within a

species that it is convenient to recognise on the

basis of characteristic attributes. A subspecies is a

special case of a geographical race which has

been given a formal trinomial name under the

terms of the International Code of Zoological

Nomenclature.

The trend in echinoderm systematics over this

century has been to synonymise regional forms

into polymorphic widespread species. Early

researchers, with relatively few specimens at their

disposal, described numerous species differenti-

ated by small morphological differences. As more

comprehensive collections have become avail-

able, the distinction between these nominal

species became blurred and many were syn-

onymised or reduced to subspecies (e.g., Fisher,

1940; A.M. Clark, 1962). Later monographs have

tended to abandon the use of subspecies as it

becomes clear that the morphological variants are

not confined to distinct regions (e.g., A.M. Clark

and Downey, 1992). However sibling species

are known to be common in marine systems

SYSTEMATICS AND BIOLOGY OF MACQUARIE ISLAND ECHINODERMS 217

(Knowlton, 1993), and recent research indicates

that morphologically similar echinoderm species

can be distinguished using reproductive or

molecular data (e.g., Bosch, 1989; Hart et al.,

1997),

The approach in this work is to follow Key’s

(1981) recommendations not to designate new

subspecific names and retain existing species

names for regional variants until we have evi-

dence to the contrary. Thus I have refrained from

establishing new subspecies for Macquarie Island

variants of Henricia obesa and Odontaster peni-

cillatus. On the other hand I have retained the

nominal species Anasterias directa, Smilasterias

clarkailsa and Trachythyone macphersonae even

though they are very close to 4. antarctica,

5. scalprifera and T. parva respectively. The final

determination of whether these are distinct

specific or infraspecific taxa will have to await

developmental or molecular research as tradi-

tional external morphological characters appear

insufficient to resolve the problem. Amongst the

numerous pentamerous Anasterias species, only

A. studeri and A, suteri can be adequately distin-

guished from A. antarctica on morphological

grounds. Other species complexes requiring

investigation are Henricia pagenstecheri/

H., simplex/H. lukinsi and H. obesa/H. aucklan-

diae. On a global scale Pteraster affinis is very

close to P. militaris from the Arctic, and Cera-

master patagonicus is similar to C, grenadensis

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

This report retains four species as Macquarie

Island endemics: Anasterias directa, Odonto-

henricia anarea, Trachythvone macphersonae,

and Trachythyone nelladana, In addition other

poorly known species listed from Macquarie

Island, such as Ophiomyxa sp. and Taeniogyrus

sp. may also be endemic. However, in no case is

the endemic status certain. A. directa and

T. macphersonae are morphological variants of

widespread Southern Ocean species. The poorly

known species are likely to be regional variants of

New Zealand or deep water species. Only Odon-

tohenricia anarea and Trachythyone nelladana

are completely distinct from their cogeners. Both

are known from shelf habitats (69-135 m) and as

such are likely to have migrated to the recently

emergent Macquarie Island via the North or South

Macquarie Ridge from New Zealand or eastern

Antarctica respectively (O’Hara, 1998a). As such

we can expect to collect these species in the future

from nearby shelf or ridge localities. The lack of

apparent speciation on Macquarie Island is a

possible indication of the conservative evolution

of echinoderms (H.B, Fell, 1962) or the contin-

uing input of genetic material from neighbouring

localities via epiplanktonic or larval dispersal

(O'Hara, 1998a).

Acknowledgments

I am most grateful to Dr C.C. Lu (now at the

National Chung Hsing University, Taiwan) and

Ms S. Boyd for their support and ready assistance

with the resources of Museum Victoria; to the

Australian Antarctic Division for supporting my

field trip to Macquarie Island in 1989 and provid-

ing the opportunity for Tonia Cochrane (NMV) to

participate in the 1986 Benthic survey; to all the

ANARE personnel who assisted me with my field

studies; to Mark O'Loughlin for his assistance

with the holothurian laboratory work and identifi-

cation of clasipods and aspidochirotes; to

K. Nolan (NMV) for diagrams; to D. Paul

(University of Melbourne) and C. Rawley (NMV)

for the photography; to Dr D. Eibye-Jacobsen

(ZMUC) and A. Morgan (BMNH) for hosting

short visits to their institutions in April 1997, to

Dr D. Pawson (USNM) for his help with

holothurian systematics and information about

the USS Eltanin echinoderms; to Dr P. Berents

(AM) and W. Zeidler (SAM) for providing

research facilities and loan of comparative speci-

mens; to Paul Anderson (NIWA), G. Paterson

(BMNH), C. Ahearn (USNM), C. Nielsen

(ZMUC) and M. Jensen (ZMUC) for the loan

of comparative material; to Dr F.W.E. Rowe and

an anonymous reviewer for reviewing the

manuscript. This paper is an outcome of

ASAC Antarctic Project 38: The biology and

classification of Macquarie Island Echinoderms.

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Memoirs of Museum Victoria 57(2): 225-236 (1999)

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA

(ACARI: HYDRACHNIDIA: ARRENURIDAE)

HARRY SMIT

Emmastraat 43-a, 1814 DM Alkmaar, The Netherlands

(smit-h@pzh.nl)

Abstract

Smit, H., 1999, New species of the water mite genus Arrenurus from eastern Australia (Acari:

Hydrachnidia: Arrenuridae). Memoirs of Museum Victoria 57: 225-236.

Five new species of the water mite genus Arrenurus from Queensland, Victoria and

Tasmania are described, viz. A. acutipetiolatus sp. nov., A. maria sp. nov., A. hybridus sp.

nov., A. queenslandicus sp. nov and A. perplexus sp. nov. The name Arrenurus mantonensis

Smit is preoccupied, and therefore the new name Arrenurus bifurcatus nom. nov. is proposed.

Arrenurus madaraszi Daday is recorded for the first time from Australia. Some measurements,

additional characters and new records are given for already known species.

Introduction

Of the cosmopolitan water mite genus Arrenurus

35 species and subspecies are known from

Australia (Smit, 1997), a relatively low number

compared to the 150 species from Europe (Smit,

1996). A key of all Australian species was pro-

vided by Smit (1997). In this paper five new

species from Victoria, Tasmania and Queensland

are described, and one new name is given to a

previously described species. Additionally,

one species new to the fauna of Australia is

reported and some additional characters and

measurements are given for already known

species.

All material has been collected by the author.

Victorian and Tasmanian holotypes and para-

types have been deposited in Museum Victoria,

Melbourne (NMV), and in the Tasmanian

Museum and Art Gallery, Hobart (TM) respect-

ively. Holotypes and paratypes from Queensland

have been deposited in the Queensland Museum,

Brisbane (QM). Other paratypes and all non-type

material have been deposited in the Zoological

Museum of the University of Amsterdam

(ZMAN).

The following abbreviations have been used

(see figs 1, 2 and 6): Al and A2, pre- and post

antennal glandularia; C24, coxoglandularia 2-4;

D1 44, dorsoglandularia 1-4; L1-4, lateroglandu-

laria 1—4; V2, ventroglandularia 2; PI-PV, palp

segments 1—5; IV-leg-4—6, fourth—sixth segments

of fourth leg; NHRS, Swedish Museum of

Natural History. For the description of the

glandularia, Jin and Wiles (1996) and Wiles

(1997) are followed. All measurements are in um,

measurements of leg and palp segments are of

the dorsal margins, Measurements of paratypes in

the description of new species are given in

brackets. Scale lines are 200 um for most figures

and 50 um for figures of the palp.

Arrenurus (Arrenurus) acutipetiolatus sp. nov.

Figures 1-6

Material examined. Holotype. Male, Victoria, swamp

at junction of Victoria Valley Road and Bundol Road,

SW of Grampians National Park, 30 Sep 1997 (NMV).

Paratypes: Victoria: 8 males, 22 females, same data

as holotype (NMV, TM, ZMAN).

Tasmania: 14 females, Reservoir of Darlington,

Maria Island National Park, Tasmania, 18 Oct 1997

(ZMAN).

Other material. New South Wales: 1 female, Mt Vic-

toria, 24 Oct 1936, leg. F. Linder (NHRS, slide 3414,

"A. fissipetiolatus").

Diagnosis. Petiole without ligulate process, nar-

rowed posteriorly, with pointed extension.

Female with L4 and V2 on small humps.

Description. Male: Body 1499 (1464-1584) in

length and 1222 (1128-1248) in width. Body

brownish. Anterior body margin concave. Body

with well developed pygal lobes. Dorsal shield

601 in width, dorsal furrow incomplete. D1 on

small humps, D3 on large humps. Setae associ-

ated with D4 on long tubercles. Genital plates

226 H. SMIT

Figures 1—4. Arrenurus acutipetiolatus. 1, holotype male, dorsal view. 2, ventral view. 3, lateral view. 4, palp.

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA 227

Figures 5, 6. Arrenurus acutipetiolatus. 5, paratype female, ventral view. 6, dorsal view.

straight, medially widened, extending to lateral

body margin. Petiole posteriorly narrowed, with a

pointed extension; ligulate process absent. Hya-

line membrane well developed, with more or less

pointed lateral angles, posteriorly concave. Setae

associated with petiole extending beyond pos-

terior margin of petiole. Lengths of PI-PV: 60,

130, 108, 149, 88; PII with 4 setae on medial side,

of which 2 in anteroventral corner. Antagonistic

bristle of PIV 106 long. Lengths of I-leg-4—6:

301, 262, 252. Lengths of IV-leg-4—6: 407, 204,

262; IV-leg-4 with a short spur. Second, third and

fourth legs with numerous swimming setae.

Female: Body 1656 (1608-1992) in length and

1416 (1344-1764) in width. Anterior margin of

body concave. Body truncated posteriorly, pos-

terolateral corners present. Dorsal shield 1056

(1032-1344) in length and 563 (446-582) in

width. Dorsal shield slender, usually widest in

anterior part, slightly tapering posteriorly, poste-

rior margin straight. Body shape and shape of

dorsal shield variable, less truncated specimens

can be found, as well as specimens in which the

dorsal shield is contracted in middle or with par-

allel margins. L4 on small humps. Distance of

fourth coxal plates larger than width of 1 geni-

tal valve. Medial margin of fourth coxal plates

larger than medial length of third coxal plates.

Gonopore 116 long. Gonopore without chitinized

patches. Genital plates wide, slightly bowed. V2

on small humps. Lengths of PI-PV: 48, 132, 118,

144, 89. Palp as in male, antagonistic bristle of

PIV 103 long. Lengths of I-leg-4—6: 281, 233,

242. Lengths of IV-leg-4-6: 349, 301, 267.

Second, third and fourth legs with numerous

swimming setae.

Etymology. The name refers to the pointed

petiole.

Remarks. The new species is close to A. fissipeti-

olatus Lundblad. Both have an almost similar

palp (long antagonistic bristle of PIV, 2-3 setae in

anteroventral corner of PII). Males are easily dis-

tinguished by the shape of the petiole, but distin-

guishing the females is more difficult. Both have

a rather slender dorsal shield, which character

they share with A. balladoniensis Halik and A.

ensifer Smit. The last species is the smallest, 1416

in length. A. balladoniensis is the largest, measur-

ing 1848-2232 in length (Smit, 1997). A. bal-

ladoniensis can be told apart from fissipetiolatus

and acutipetiolatus by the shape of the genital

plate, which is much narrower than that of the last

two species. A. acutipetiolatus differs from A. fis-

sipetiolatus by having L4 on tubercles and a pos-

teriorly more truncated body. Another useful

228 H. SMIT

character by which to distinguish the two species

is the shape of the dorsal shield, which is more

slender in acutipetiolatus. The ratio length/width

of the dorsal shield is 1.73-2.04 in acutipetiolatus

(usually >1.80) and 1.50-1.67 in fissipetiolatus.

Lundblad (1947) described the female of A.

fissipetiolatus, but he had some doubts if the

assignment to this species was correct, because

the male and the female came from different loca-

tions. The female described by Lundblad matches

the description of 4. acutipetiolatus (L4 on

humps, body truncated, ratio dorsal shield 1.89),

and should therefore be assigned to that species.

Arrenurus (Arrenurus) hybridus sp. nov.

Figures 7—10

Arrenurus sp. — Harvey, 1998: 106, fig. 32C.

Material examined. Holotype. Male, Victoria, pond,

Hospice Plain, Mt Buffalo National Park, 10 Oct 1997

(NMV).

Figures 7-10. Arrenurus hybridus, holotype male, 12, dorsal view, 13, ventral view. 14, palp. 15, paratype

female, ventral view.

Figure 11. Arrenurus fissipetiolatus Lundblad, holotype male, detail of petiole.

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA 229

Paratypes. Victoria: | male (ZMAN), 1 male (NMV),

same data as holotype; 1 male (not sclerotized, ZMAN),

1 female (NMV), small ponds, Kowan Plain, + 1400 m

above sea level, Mt Buffalo National Park, 10 Oct 1997.

Diagnosis. Body with distinct cauda. Petiole pre-

sent, spatulate. Hyaline membrane and ligulate

process absent.

Description. Male: Body 951 (912-951) long and

543 (504-553) wide. Anterior body margin

rounded. Body posteriorly gradually tapering into

a distinct cauda. Cauda with a shallow concavity.

Dorsal furrow not closed posteriorly. Posterior

margin almost straight, with small indentations.

First coxal plates extending beyond anterior body

margin. Second and third coxal plates lying very

close. Gonopore 72 long. Genital field 197 long,

not extending to lateral body margin, slightly

undulating. Petiole spatulate, hyaline membrane

and ligulate process absent. Lengths of PI-PV: 38,

82, 65, 91, 31; PII with 7 (8?) setae on medial

side. Lengths of I-leg-4—6; 136, 136, 146. Lengths

of IV-leg-4-6: 194, 184, 155. IV-leg-4 without

spur, but IV-leg-5 with dorsodistal extension.

Second, third and fourth legs with numerous

swimming setae.

Female: Body 1106 long and 825 wide, egg-

shaped, without posterolateral corners. Anterior

body margin rounded. Dorsal shield 970 long and

679 wide, dorsal furrow closed. Medial lengths of

third and fourth coxal plates of equal length,

fourth coxal plates almost without posteromedial

corner. Gonopore large, 165 in length; gonopore

with indistinct sclerotized patches. Genital plates

bowed, laterally somewhat rectangular. Lengths

of PI-PV: 46, 98, 82, 108, 48; PII with 8 setae on

medial side. Second, third and fourth legs with

numerous swimming setae.

Etymology. The name refers to the somewhat

intermediate position of the new species between

the subgenera Arrenurus and Megaluracarus.

Remarks. No other Indo-Australian Arrenurus

species has a distinct cauda with a spatulate peti-

ole. The female is characterized by the combina-

tion of the absence of posterolateral corners, the

absence of posteromedial corners of the fourth

coxal plates and the medial margins of the third

and fourth coxal plates being of equal length.

Arrenurus (Arrenurus) bifurcatus nov. nom.

Remarks. When describing A, (Arrenurus) manto-

nensis Smit, 1997, T was not aware of the exis-

tence of A. (Megaluracarus) mantonensis

George, 1903. This species was synonymized by

Viets (1956) with A. buccinator (Müller). There-

fore, the species described by me is a junior

homonym of A. mantonensis George and thus

requires a new name for which I propose Arrenu-

rus bifurcatus nom. nov. The new name refers to

the bifurcated setae of the petiole.

Arrenurus (Arrenurus) fissipetiolatus Lundblad

Figure 11

Lundblad,

1986, figs

Arrenurus

1947: 73,

1632-1637,

(Arrenurus) _fissipetiolatus

figs 46A—D. — Cook,

Material examined. Holotype. Male, Victoria, Mary-

borough, 24 Sept 1926, leg. E.J. Semmens (NHRS,

slide 3413).

Other material. Victoria: 1 female, swamp at junction

of Victoria Valley Road and Bundol Road, SW of

Grampians National Park, 30 Sep 1997.

Tasmania: 1 male, 14 females, Blackmans Lagoon,

Waterhouse Protected Area, 21 Oct 1997; 4 females,

Little Waterhouse Lake, Waterhouse Protected Area, 21

Oct 1997,

Description. Male: Body 1488 (1337-1483)

in length and 1272 (1064-1138) in width (in

brackets the measurements of Lundblad, 1947

and Cook, 1986).

Female: Body 1680-1944 (Cook, 1986; 1581)

in length and 1392-1656 (Cook, 1986: 1292) in

width. Dorsal shield 1104-1320 in length and

281-364 in width; usually widest in middle. L1

shifted dorsally towards dorsal shield and there-

fore visible in dorsal view.

Remarks. Contrary to Lundblad's description, the

male does not have a long spur. The holotype

(Figure 16) and my own specimen have a notch in

the posterior margin of the petiole. However,

Cook (1986) could not observe this in all his

specimens. The genital plate of most of my

female specimens fit well with the description of

Cook (1986). However, occasionally some speci-

mens have a narrower genital plate, while others

have a genital plate with a slightly undulating

posterior margin, The species has been reported

from Victoria and Tasmania. However, the num-

ber of specimens collected so far is very limited.

Therefore, some additional characters and

measurements are given. The female described by

Lundblad (1947) belonged to another species (see

under A. acutipetiolatus).

Arrenurus (Megaluracarus) maria sp. nov,

Figures 12-16

Material examined. Holotype. Male, Tasmania, reser-

voir of Darlington, Maria Island National Park, 18 Oct

1997 (TM).

230 H. SMIT

Figures 12-16. Arrenurus maria. 7, holotype male, dorsal view. 8, ventral view. 9, lateral view. 10, palp. 11,

paratype female, ventral view.

Paratypes. Tasmania: | female, same data as holo-

type (TM); 1 female, Apsley River, at crossing with

Tasmanian Highway, 19 Oct 1997 (ZMAN).

Diagnosis. Male with a long cauda, posterior

margin of cauda with a small hyaline area and

4 rounded hyaline extensions, 1 peg-like

rudimentary petiole.

Description. Male: Body 1465 long and 708 wide.

Anterior and posterior body part brownish, mid-

dle part bluish, cauda purple; legs bluish. Anterior

body margin almost straight, body truncated ante-

riorly. In the middle between the eyes an area

without body pores. Postocularia setae and setae

associated with dorsoglandularia 2 and 4 very

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA 231

long. Cauda much longer than wide, width of

cauda 359. D4 on small humps. Posterolateral

corners of cauda rounded. Cauda posteriorly with

a distinct hyaline area and 4 rounded, hyaline

extensions. Petiole rudimentary, peg-like. Genital

field 252 in width. Genital plates short and

rounded, Lengths of PI-PV: 24, 94, 62, 116, 53.

PII with a large patch of setae lying on a bulge.

Lengths of I-leg-4—6: 206, 243, 180. Lengths of

IV-leg-4-6: 272, 184, 243; IV-leg-4 with a short

spur. Second, third and fourth legs with numerous

swimming setae.

Female: Body 1416 (1320) long and 1077

(1009) wide. Dorsal shield 897 long and 902

wide; dorsal furrow complete. Body brownish,

posterior and anterior body parts purple; legs

bluish. Between eyes an area without body pores.

Body egg-shaped, without posterolateral or with

indistinct posterolateral corners. Medial margin

of third coxal plates larger than medial margin of

fourth coxal plates. Medial distance of fourth

coxal plates slightly smaller than width of 1 gono-

pore valve. Gonopore 194 long. Genital plates

short and wide, sloping posteriorly, extending

beyond posterior margin of gonopore. Lengths of

PI-V: 36, 94, 67, 122, 50; palp as in male. Lengths

of I-leg-4—6: 165, 204, 146. Lengths of IV-leg-

4-6; 233, 272, 276. Second, third and fourth legs

with numerous swimming setae.

Etymology. Named after the island where the type

specimen was collected. Noun in apposition.

Remarks. The male of the new species is close to

anumber of Australian species with a long cauda,

Le. A. otodus Cook, A. gilvus Smit and A. vander-

palae Smit. The new species differs from

A. gilvus and A. vanderpalae by its large size and

the presence of only one peg-like rudimentary

petiole (two in gilvus and vanderpalae), and from

A. otodus by the lack of the pointed postero-

lateral corners of the cauda. Moreover, A. maria

is larger than A. otodus, has a larger hump on

which D4 are located, and its rounded hyaline

extensions are not found in 4. otodus. The female

can be distinguished from all other species by

having the medial margin of the third coxal

plates larger than the medial margin of the fourth

coxal plates.

Arrenurus (Micruracarus) forpicatoides

Lundblad

Arrenurus (Micruracarus) forpicatoides Lundblad,

1941: 160. — Lundblad, 1947: 75, figs 47A-D. —

Uchida and Imamura, 1951: 353, figs 18a-d. — Smit,

1992: 109.

Material examined. Victoria: 10 males, 21 females,

swamp at junction of Victoria Valley Road and Bundol

Road, southwest of Grampians National Park, 30 Sep

1997; 3 females, Lake Catani, Mt Buffalo National

Park, 10 Oct 1997; 1 male, unnamed creek 4,5 km east

of Shipwreck Creek, Croajingolong National Park, 23

Oct 1997.

Tasmania: | male, 2 females, old river branch of Coal

River, north of Richmond, 17 Oct 1997; 1 female,

swamp 12 km south of Gladstone, along road B82, 20

Oct 1997; 2 males, 10 females, Big Waterhouse Lake,

Waterhouse Protected Area, 21 Oct 1997; 2 females,

Little Waterhouse Lake, Waterhouse Protected Area, 21

Oct 1997.

Description. Male: Body 689-786 in length and

543-640 in width. Petiole occasionally reaching

posterior body margin. Caudal lobes in some

specimens rounded.

Female: Body 776-936 in length and 650-786

in width. Dorsal shield complete.

Remarks. The species was previously reported

from Victoria, South Australia and Queensland.

The specimens from Queensland reported by Smit

(1992) do not belong to this species but to a new

species described below. The record from China

(Uchida and Imamura, 1951) needs confirmation

as only females have been collected. Females of

the subgenus Micruracarus with two pairs of

rounded chitinized patches on the gonopore are

difficult to identify. Moreover, the genital field of

the specimens illustrated by Uchida and Imamura

(1951) is laterally narrowed, a character not found

in the Australian specimens.

Arrenurus (Micruracarus) queenslandicus

sp. nov.

Figures 17-20

Arrenurus (Micruracarus) forpicatoides. —

1992: 109.

Smit,

Material examined. Holotype. Male, Queensland, pond

north of Normanton, 14 Aug 1989 (QM).

Paratypes. Queensland: 3 males, 5 females

(QM), 3 males, 5 females (NMV), 3 males, 5

females (ZMAN), all same data as holotype; 3

males, 1 female (ZMAN), pond near Crocodile

Road, Bowling Green Bay National Park, 31 Jul

1989; 3 males, 1 female (ZMAN), pond

Townsville Common National Park, | Aug 1989;

1 male, Freshwater Lagoon, Horseshoe Bay,

Magnetic Island, 3 Aug 1989.

Other material. Queensland: 2 females, pond north of

Longreach, 15 Aug 1989.

Diagnosis. Cauda of male indistinctly set off from

anterior part of body. Petiole tong-shaped,

hyaline, fused with body by small hyaline area.

Dorsal shield of female incomplete.

232 H. SMIT

Figures 17 20. Arrenurus queenslandieus. 17, holotype male, dorsal view. 18, ventral view. 19, palp. 20,

paratype female, ventral view.

Description. Male: Body 708 (660-795) in

length and 611 (553-679) in width. Body yellow-

ish brown to greenish. Anterior margin of body

straight to slightly concave. Dorsal shield com-

plete, 423 (359-456) in length and 417 (373-441)

in width. Genital plates long and narrow, extend-

ing onto dorsum, Cauda indistinctly set off from

anterior part of body. Cauda with median cleft.

Petiole hyaline, tong-shaped, fused with body by

a small hyaline area. Lengths of PI-PV: 31, 61,

40, 84, 50; PIL with 4 setae on medial side, of

which 2 close to ventral margin. Lengths of I-leg-

4-6: 110, 113, 132. Lengths of IV-leg-4-6: 144,

134, 142; IV-leg-4 without a spur. Second, third

and fourth legs with numerous swimming setae.

Female: Body egg-shaped, 951 (936—1048) in

length and 786 (771-844) in width. Anterior body

margin straight or slightly concave. Posterolateral

corners of body almost absent. However, occa-

sionally posterolateral corners more pronounced,

and in these specimens body truncated pos-

teriorly, Dorsal shield incomplete, 524 (495-572)

in width. Medial distance of fourth coxal plates

longer than width of | genital valve. Medial

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA 233

margin of fourth coxal plates longer than medial

margin of third coxal plates. Gonopore 98 long.

Genital valves with large chitinous patches, ante-

rior and posterior patches connected by chitinous

strip. Genital plates straight to slightly bowed,

tapering laterally. Lengths of PI-PV: 36, 70, 48,

89, 43; palp as in male. Lengths of I-leg-4-6: 130,

134, 125. Lengths of [V-leg-4 6: 155, 155, 136.

Second, third and fourth legs with numerous

swimming setac.

Etymology. The name refers to the state of

Queensland in which the species has been found,

Remarks. 1 erroneously identified the species as

A. forpicatoides (Smit, 1992). However, in the

last species the cauda is distinctly set off from the

body, the body is reddish brown and the tong-

shaped part of the petiole is narrower. A. anbang-

bang Smit has a different shaped petiole which is

not fused with the cauda. The female of the new

species closely resembles A. forpicatoides but is

larger, has a different colour and an incomplete

dorsal shield.

Arrenurus (Micruracarus) perplexus sp. nov.

Figures 21-24

Material examined. Holotype. Male, Victoria, pond

near Buckland River, at crossing with Buckland Valley

Road (west of Bright), 11 Oct 1997 (NMV).

Paratype. Victoria: 1 female, same data as holotype

(NMV).

Diagnosis. Cauda of male distinctly set off from

body, lateral margin of cauda convex, body bluc,

petiole hyaline, tong-shaped. Body of female

tapering posteriorly.

Description. Male: Body 635 long and 519 wide.

Dorsal shield complete, 320 long and 320 wide.

Body blue. Cauda distinctly set off from anterior

part of body, lateral margins of cauda convex,

caudal lobes rounded. Cauda posteriorly with a

median cleft. D1 on small humps. Petiole hyaline,

tong-shaped, fused with body by a small hyaline

area. Genital plates narrow and long, extending

onto dorsum. Lengths of PI-PV: 28, 50, 38, 72,

41; PII with 4 setae on medial side. Lengths of I-

leg-4-6: 98, 94, 113. Lengths of IV-leg-4-6: 132,

125, 108; IV-leg-4 without spur. Second, third

and fourth legs with numerous swimming setae.

Female: Body 757 long and 630 wide. Dorsal

shield complete, 582 long and 432 wide; dorsal

shield somewhat tapering posteriorly. Body taper-

ing posteriorly, slightly truncated. Anterior mar-

gin of body straight. Medial distance of fourth

coxal plates slightly larger than 1 genital valve.

Medial margin of fourth coxal plates larger than

medial margin of third coxal plates. Gonopore 97

long, each valve with 2 rounded chitinous

patches. Genital plates straight, slightly sloping

posteriorly and laterally somewhat enlarged.

Lengths of PPV: 31, 55, 41, 86, 43. Palp as in

male, but medial side of PIL with 3 setae. Lengths

of I-leg-4-6: 121, 125, 120. Lengths of IV-leg-

4-6: 170, 156, 130. Second, third and fourth legs

with numerous swimming setae.

Etymology. The name refers to the complicated

structure of the petiole.

Remarks. The new species is close to A. forpica-

toides, but differs in size, body colour and shape

of the cauda. The hyaline petiole of the two

species are nearly identical, although the tong-

shaped part of the petiole is narrower in the new

species. In A. queenslandicus the cauda is not

set off from the body, but the petioles of the

two species are very similar. The female of

A. perplexus can be distinguished from A. forpi-

catoides by the body colour, shape of the body

and the broader genital plates, and from

A. queenslandicus by the complete dorsal shield.

The species of Micruracarus with a hyaline

petiole form a complex group within the genus

Arrenurus. In my material (from Hasties Swamp,

Atherton Tablelands, Queensland) another

species might be present, of which the only male

is somewhat intermediate between A, queens-

landicus and A. perplexus. There are differences

in shape of the cauda and the body between this

male and the males of the two aforementioned

species. Unfortunately, all three females from this

location are different, and I refrain from describ-

ing it as a new species until more material is

available.

Arrenurus (Micruracarus) madaraszi Daday

Arrenurus Madarászi Daday, 1898: 99, figs 49a i.

Arrenurus (Micruracarus) forpicatoides. Smit,

1992; 109 (part).

Material examined. Queensland: | male, pond in

Townsville Common National Park, Queensland, | Aug

1989 (ZMAN).

Remarks. Initially, the specimen has been identi-

fied erroneously as A. forpicatoides to which it is

closely related. A. madaraszi is a very widespread

species, known from Japan, China, Burma, India,

Sri Lanka throughout Indonesia. This is the first

record for Australia.

234 H. SMIT

Figures 21-24. Arrenurus perplexus. 21, holotype male, dorsal view. 22, ventral view. 23, palp. 24, paratype

female, ventral view.

Figure 25. Arrenurus haswelli Cook, ventral view female.

Figure 26. Arrenurus novaehollandiae Lundblad, ventral view male.

NEW SPECIES OF THE WATER MITE GENUS ARRENURUS FROM EASTERN AUSTRALIA 235

Arrenurus (Truncaturus) haswelli Cook

Figure 25

Arrenurus (Truncaturus) haswelli Cook, 1986: 309,

figs 1653-1658, 1660.

Material examined. Holotype. Male, Tasmania, pond

on southwest side of Great Lake, 18 Mar 1981 (NMV,

slide K705). Paratype, female, same data as holotype

(NMV, slide K706).

Other material, Tasmania: 7 males, 13 females, wet-

land south of Derwent River, 10 km east of New Nor-

folk, 17 Oct 1997 (ZMAN).

Description. Male: Body brownish.

757-834 in length and 485-533 in width.

Female: Body yellowish-brown to reddish-

brown.

Males

Remarks. Males in the series collected by me are

smaller than the males of the type series, which

measured 897-927 in length and 593-616 in

width (Cook, 1986). Otherwise, males from this

study fit well in the description of Cook. The

females in my collection differ in a number of

characters from females of the type series. Geni-

tal plates are much narrower as illustrated by

Cook (1986). Moreover, the medial length of the

fourth coxal plates is of equal length or smaller

than the medial length of the third coxal plates

(Figure 25). In the paratype female the medial

lengths are more or less of equal size. The females

from this study measure 960—1140 in length and

757-883 in width. As in the males, the females of

this study are smaller than the type series,

So far, the species was only known from few

specimens from the type locality, which is situ-

ated in Tasmania as well, The variation in genital

plates is not unusual in female Arrenurus species

(Smit, 1995). Because of this, and the fact that the

males in my collection fit very well in the descrip-

tion of Cook (1986), all females are assigned to

A, haswelli,

Arrenurus (Truncaturus) novachollandiae

Lundblad

Figure 26

Arrenurus (Truncaturus) novaehollandiae Lundblad,

1947: 79, figs 50A-D.

Material examined. Tasmania: 2 males, 7 females,

reservoir of Darlington, Maria Island National Park, 18

Oct 1997 (ZMAN).

Description. Male: Body brownish. Body

771—786 in length and 640-645 in width, Genital

plates wide, slightly bowed.

Female: Body colour as in male. Body 849-980

in length and 713-805 in width.

Remarks. Tasmanian specimens differ in colour

and size from the type series but apart from this fit

well with the description of Lundblad (1947).

Specimens of the type-series are yellow-green

and smaller. The species has only been reported

from Victoria.

Acknowledgements

l am indebted to the Department of Natural

Resources and Environment (Melbourne) and the

Parks and Wildlife Service (Hobart) for permis-

sion to collect water mites in the national parks of

Victoria and Tasmania respectively, to Dr T. Kro-

nestedt for the loan of type material of Arrenurus

fissipetiolatus and A. australicus and to Dr K.

Walker for the loan of type material of 4.

haswelli. Johannes Postma (Ann Arbor) corrected

the English.

References

Cook, D.R., 1986, Water mites from Australia.

Memoirs of the American Entomological Institute

40: 1-568,

Daday, E. von, 1898. Mikroskopische Süsswasserthiere

aus Ceylon. Természetrajzi Füzetek, Anhangshefl

21: 1-123.

George, C.F.. 1903. New British

Naturalist 28: 215-216.

Harvey, M.S., 1998. The Australian water mites. A

guide to families and genera. Monographs on

Invertebrate Taxonomy 4. CSIRO Publishing:

Collingwood, 150 pp.

Jin Daochao and Wiles R., 1996. New species of

Arrenurus Dugès (Acari: Hydrachnidia: Arrenuri-

dae) from China and first records of watermites

from Laos. Acarologia 37: 317-344.

Lundblad, O., 1941. Neue Wassermilben aus Amerika,

water mites.

Afrika, Asien und Australien. Zoologischer

Anzeiger 133: 155-160.

Lundblad, O., 1947. Zur Kenntnis Australischer

Wassermilben. Arkiv för Zoologi 40A: 1-82.

Smit, H., 1992. Water mites from New South Wales

and Queensland, Australia (Acari, Hydrachnellae).

Tijdschrift voor Entomologie 135: 91— 112.

Smit, H., 1995. New records of water mites from

Morocco, with the description of one new sub-

species (Acari, Hydrachnellae). Aquatic Insects 17:

17-24.

Smit, H., 1996. A revision of enigmatic species within

European members of the genus Arrenurus Dugès

(Acari, Hydrachnellae), Annales de Limnologie 32

(3): 137-146.

Smit, H., 1997. Australian water mites of the genus

Arrenurus, with the description of 12 new species,

from northern and western Australia (Acari:

Hydrachnellae: Arrenuridae). Records of the

Western Australian Museum 18: 233-261.

236 H. SMIT

Uchida, T. and Imamura, T., 1951. Some water-mites

from China. Journal of the Faculty of Science,

Hokkaido University, Sapporo, Serie VL, Zoology

10: 324-358.

Viets, K., 1956. Die Milbes des Süßwassers und des

Meeres. Zweiter und dritter Teil. Katalog und

Nomenclator. Gustav Fisher: Jena.

Wiles, P.R., 1997. The homology of glands and glandu-

laria in the water mites (Acari: Hydrachnidia).

Journal of Natural History 31: 1237-1251.

Memoirs of Museum Victoria 57(2): 237-239 (1999)

A SECOND SPECIES OF ATRIPLECTIDES MOSELY FROM AUSTRALIA

(TRICHOPTERA: ATRIPLECTIDIDAE)

ARTURS NEBOISS

Entomology, Museum Victoria, 71 Victoria Crescent, Abbotsford, Victoria 3067, Australia

Abstract

Neboiss, A., 1999. A second species of Atriplectides Mosely from Australia (Trichoptera:

Atriplectididae). Memoirs of Museum Victoria 57: 237-239.

Atriplectides ikmaleus sp. nov. is described from the North Queensland Wet Tropics Her-

itage Area. Its adult and larval features are compared with those of several recently described

neotropical atriplectididines.

Introduction

Investigation of the North Queensland Wet Trop-

ics Heritage Area (Walker et al., 1993; 1995)

revealed many undescribed species of Tri-

choptera which were given code-numbers pend-

ing formal species description. This material

included a small number of specimens of

Atriplectides dubius Mosely from several local-

ities, as well as an undescribed Afriplectides

species (code-number PT-2010).

Atriplectides dubius, hitherto the only

Australian atriplectidid species known, is widely

distributed and has been collected from numerous

localities. It was described from Tasmania and

subsequently found to be common throughout

eastern Victoria and New South Wales. It has

been recorded from Queensland as far north as

the Mareeba district north of Cairns. It has been

taken on Kangaroo Island and in the Adelaide

Hills, South Australia, and from southwestern

Western Australia. The single Western Australian

specimen (Seldom Seen Brook, Jarradale, S.

Bunn, 4.xi.1983, NMV collections) is marginally

smaller than the average specimens from eastern

Australia but shows no structural differences. The

new species differs in wing and genitalic features;

it overlaps the northern distribution of A. dubius.

Specimens were prepared for examination by

clearing the abdomens in cold KOH solution. The

dissected and figured specimens are identified by

the author’s notebook number with the prefix

‘PT-’. Material is deposited in the Australian

National Insect Collection, Canberra (ANIC) and

Museum Victoria, Melbourne (NMV).

Atriplectides ikmaleus sp. nov.

Figures 1—4

Material examined. Holotype male, North Queensland,

Upper Mulgrave river via Gordonvale, 17°10'S,

145°53’E (Wet Tropics Investigation area 5),

29-30.iv.1970, S.R. Curtis, ANIC (genitalic prep. PT-

2010, illustrated).

Paratype male, North Queensland, Yuccabine Creek,

Kirrama State Forest, 18°12’S 145°54E (Wet

Tropics Investigation area 9), xii.1985, R. Pearson &

L. Benson, NMV 117263 (genitalic prep. PT-1588).

Diagnosis. Forewing length 15.2-15.4 mm,

slightly larger than A. dubius. Forewing fork 1

with footstalk short; cross-vein r-m at about dis-

tal third of discoidal cell; a small cross-vein

between M1+2 and M3+4 about halfway between

arculus and wing margin; male genitalia with

single-segmented inferior appendages.

Description. Male. Wings (Fig. 4) fuscous with-

out mottling (both specimens preserved in alco-

hol and faded, paratype male slightly teneral).

Forewing fork 1 with short footstalk, fork 2 long,

broadly sessile; an unusual cross-vein between

M1+2 and M3+4 halfway between arculus and

wing margin; hind wing Sc joins RI shortly

before wing margin, and at this point small cross-

vein connects to R2+3: discoidal cell elongate tri-

angular. Antennae slender, scape short, distinctly

bulbous; segment 2 very short, slightly thicker

than segment 3, subsequent segments long and

slender, Maxillary palpi 5-segmented, first three

segments more robust than distal segments,

elongate; segments 4 and 5 distinctly thinner.

238 ARTURS NEBOISS

EL Sn,

4 ER

Figures 1—4. Atriplectides ikmaleus sp. nov. 1-3, male

venation.

Figures 5, 6. Atriplectides sp. (presumably ikmaleus). 5, head and thorax, dorsal view; 6, larval case, dorsal

view.

genitalia, dorsal, ventral and lateral views; 4, wing

A SECOND SPECIES OF CADDIS FLY (ATRIPLECTIDIDAE) 239

Genitalia (Figs 1-3). Abdominal segment IX

short, lateral margins slightly lobose; segment X

with a deep V-shaped cleavage separating rather

slender, triangular, distally tapered lobes. Supe-

rior appendages slightly shorter than segment X,

stout, rounded distally. Inferior appendages

single-segmented, slightly compressed later-

ally, divergent distally, shorter than segment

X. Phallus short, truncate, apically a distinct lip.

Female. Unknown.

Etymology. Latinised from ikmaleos, Greek —

damp, wet — in reference to the wet tropical envi-

ronment of the North Queensland World Heritage

Area

Remarks. Atriplectides ikmaleus adults are known

from only the two North Queensland localities,

150 km apart. The new species is distinguished

from A. dubius by being larger (forewing

15.2-15.4 versus 12-14 mm), wing venation and

the single-segmented inferior appendages of the

male genetalia.

Two larval specimens are assigned to this

species, collected from the same general arca as

the paratype (Yuccabine Creek, Kirrama State

Forest, x.1983, R. Pearson & L. Benson, NMV).

They are figured and keyed out as “Species AVI

in the ‘Preliminary keys ..... Atriplectididae, etc.’

by John Dean (in litt.). The sand-grain case (Fig.

6) is broad, dorsoventrally flattened with larger

ballast particles attached along the lateral mar-

gins. The larva (Fig. 5) is distinguished from that

of A. dubius (Neboiss, 1978) by the presence of

five sclerites on the anterior half of the pronotum;

and the hind tarsal claw less than half the length

of the tarsus. The number and position of the gills

resemble the arrangement in an unidentified

Brazilian atriplectidid larva described and figured

by Holzenthal (1997) as does also the curiously

elongate arrangement of the pronotum, strength-

ened by additional sclerites. A similar prothoracic

arrangement was described by Marlier (1978: 43)

in an ondontocerid larva, Hughscottiella auri-

capilla Ulmer, from the Seychelles. This larva

differs, however, in having strongly enlarged

midlegs and its species was transferred to

Atriplectididae by Neboiss (1978: 67).

The Atriplectididae were recently recorded

from the Neotropics for the first time by Holzen-

thal (1997) who erected a new genus, Neoa-

triplectides, for a species from Peru, Bolivia and

Ecuador (N. froehlichi Holzenthal). Neoatriplec-

tides is characterised by the single-segmented

inferior appendages of the male, a feature also

seen in the new Australian species. However, the

wing venation of the Australian species differs

from that of the Neotropical genus.

Acknowledgements

My sincere thanks are extended to Mr John Dean,

Melbourne, for providing the informative details

of the larvae and giving permission to use his

illustrations, and to Dr Alice Wells, Canberra, for

continuous encouragement, discussions and help

in the preparation of this manuscript.

References

Holzenthal, R.W., 1997. The caddisfly (Trichoptera)

family Atriplectididae in the Neotropics. Pp.

157-165 in Holzenthal, R.W. and Flint, O.S. Jr

(eds) Proceedings of the Sth International Sym-

posium on Trichoptera. Ohio Biological Survey :

Colombus, Ohio, USA.

Marlier, G., 1978. Les larves et nymphes des Trichop-

tères des Seychelles. Pp. 31 54 in Crichton, M.I.

(ed) Proceedings of the 2nd International Sym-

posium on Trichoptera. Dr W. Junk Publishers:

The Hague.

Neboiss, A., 1978. Atriplectididae, a new caddisfly

family (Trichoptera: Atriplectididae). Pp. 67-73 in

Crichton, LM. (ed.) Proceedings of the 2nd Inter-

national Symposium on Trichoptera. Dr W. Junk

Publishers: The Hague.

Walker, K., Neboiss, A., Dean, J. and Cartwright, D.,

1993. A Preliminary Investigation of the Caddis-

flies (Trichoptera: Insecta) of the Queensland Wet

Tropics World Heritage Area. Museum of

Victoria, Department of Entomology: Melbourne.

177 pp.

Walker, K., Neboiss, A., Dean, J. & Cartwright, D.,

1995. A preliminary investigation of the caddis-

flies (Trichoptera: Insecta) of the Queensland

Wet Tropics. Australian Entomologist 22: 19-31

[Summary of the 1993 report].

Memoirs of Museum Victoria 57(2): 241-250 (1999)

A NEW GENUS OF SUBCORTICAL COCCOIDS (HEMIPTERA: COCCOIDEA:

ERIOCOCCIDAE) ON EUCALYPTUS

P. J. GULLAN

Division of Botany and Zoology, The Australian National University

Canberra, A.C.T. 0200, Australia

(Penny.Gullan@anu.edu.au)

Abstract

Gullan, P.J., 1999. A new genus of subcortical coccoids (Hemiptera: Coccoidea: Eriococcidae)

on Eucalyptus. Memoirs of Museum Victoria 57: 241-250.

A new genus of Eriococcidae (Hemiptera: Coccoidea), Subcorticoccus gen. nov., is

described for three new species of scale insects collected under eucalypt bark in southeastern

Australia. S. beardsleyi sp. nov. occurs on Eucalyptus macrorhyncha near Melbourne,

Victoria, whereas both S. huonamnis sp. nov. and S. murrindindi sp. nov. feed on E. regnans

in Tasmania and Victoria, respectively. The adult females of all three species and the

first-instar nymph of S. beardsleyi are described and illustrated. Subcorticoccus appears to be

morphologically most similar to the Australian genus Phacelococcus Miller.

Introduction

The Eriococcidae are a speciose family of scale

insects with major radiations in North and South

America (e.g., Miller and Gonzalez, 1975; Miller

and McKenzie, 1967; Miller and Miller, 1992),

New Zealand (Hoy, 1962) and Australia (e.g.,

Froggatt, 1921; Hoy, 1963; Gullan, 1984). Except

for the gall-inducing taxa, Australia’s eriococcids

have been poorly studied since the pioneering

work of Froggatt (1921). It is thus not surprising

that current taxonomic and cladistic studies of the

Australian Eriococcidae are revealing a number

of new taxa. In particular, one undescribed taxon

exhibits unique morphology which appears most

similar to that of Phacelococcus Miller. This

taxon is represented by three undescribed species

all of which were collected under Eucalyptus

bark in southeastern Australia.

The adult females of these three new species

differ from those of the speciose and cosmopoli-

tan genus ZEriococcus Targioni-Tozzetti and

related genera in lacking anal lobes, enlarged

dorsal setae, differentiated marginal setae and

microtubular ducts, and in possessing a ventral,

noncellular anal ring. They resemble Phacelococ-

cus (Miller, 1970; Gullan and Strong, 1997) in

possessing clusters of quinquelocular pores on

the ventral abdomen and in having very small

legs but differ in lacking anal lobes and

241

microtubular ducts, and in having ventral frontal

lobes, a mediolongitudinal band of microtrichia

on the anterior dorsum and a single pair of anal

ring setae on a simple anal ring. Phacelococcus

and the undescribed species also share the habit

of living under the bark of their eucalypt hosts.

All of these species have been collected only

rarely probably because of their cryptic habit and

yet they may form an important dietary com-

ponent for a number of other arboreal animals

including mammals (Gullan and Strong, 1997)

and arthropods. Since no observations are avail-

able on live specimens of the new species it is not

known whether nymphs and adult females pro-

duce honeydew. The biology of these eriococcids

warrants further study.

This paper erects a new genus, Subcorticoccus,

for these three new species collected under euca-

lypt bark in southeastern Australia. The adult

females of S. beardsleyi sp. nov., S. huonamnis

sp. nov. and S. murrindindi sp. nov. and the first-

instar nymph of S. beardsleyi are described

and illustrated. The terminology and the slide-

mounting and illustrative techniques employed

are the same as those in Gullan and Strong (1997)

except that the antennal sensilla are named

according to Koteja (1980) and Le Rü et al.

(1995). Thus trichoid sensilla are equivalent to

the antennal hair-like setae mentioned in most

other coccoid descriptions and the different types

242

of pegs have been variously called ‘fleshy setae’

or ‘antennal bristles’ by most previous authors:

the antennal basiconic and coeloconic sensilla

(which can be difficult to distinguish) and the

campaniform sensillum usually are not mentioned

in descriptions of coccoids.

Material is deposited in The Australian

National Insect Collection, CSIRO Entomology,

Canberra ACT, 2601, Australia (ANIC); The

Natural History Museum, London SW7 5BD, UK

(BMNII); Bernice P. Bishop Museum, Honolulu,

Hawai, USA (BPBM); and Museum Victoria,

Melbourne, Victoria, Australia (NMV).

Subcorticoccus gen. nov.

Type species. Subcorticoccus murrindindi sp.

nov.

Description. Adult female with abdomen tapering

to rounded apex; derm membranous with rugu-

lose microsculpturing, especially obvious margin-

ally, and distinctive, mediolongitudinal band of

microtrichia on dorsum of head and anterior

thorax and much shorter band or indistinct cluster

of microtrichia ventrally between antennae; pair

of eyespots on body margin; antennae 6-7 (rarely

5) segmented, segments II to apical one subequal

in length; 0-4 trichoid sensilla per antennal seg-

ment, lengths and distribution on segments vari-

able but apical segment always with 2 4 sensilla,

20-35 um long, and segment IV (or V if 7 seg-

ments) always lacking trichoid sensilla; pegs

5 25 um long on antennal segments IV-VI

(V VIL if 7 segments), distributed as follows: 1

on IV (or V), 1 on V (or VI), usually 3 on VI (or

VID; usually 2 either basiconic or coeloconic sen-

silla, 5-12 um long, only on apical segment (Vl or

VII); pair of oval, slightly raised, frontal lobes

with rugulose surface, posteromedial to antennae:

labium conical, width equal or greater than

length, segmentation not apparent, segments pos-

sibly fused; legs reduced, less than 160 um long;

digitules of tarsi and claws capitate; tarsal claws

with or without small denticle near apex; anal

lobes completely lacking, their position indicated

P. J. GULLAN

by pair of apical seta, one on each side of abdom-

inal apex; anal ring ventral, simple, noncellular,

usually with one flagellate anal ring seta (7-18

um long) laterally on each side of ring and pair of

flagellate suranal seta (8-20 um long) just outside

ring; dorsal setae short (4-16 um long) and flag-

ellate, in segmental rows, enlarged setae absent;

ventral setae flagellate, longest (13-33 um) near

vulva; remainder similar in length to dorsal setae;

differentiated marginal setae absent; slender

macrotubular ducts, 10-16 jum long, 1-2 um

wide, present on dorsum and venter, scattered in

bands across segments and in clusters on body

margin, each duct with delicate inner filament

(= ductule) with terminal knob barely distinguish-

able; microtubular ducts absent; multilocular

pores mostly quinquelocular, very occasionally

trilocular especially near spiracles, 3-6 um in

diameter, distributed in bands on ventral posterior

abdomen and sometimes also scattered on

margins of body, small clusters at opening of

spiracles; bilocular pores absent,

Etvmologv. The genus name is descriptive of the

under-hark habit of the species (sub, meaning

under, Latin; corticis, meaning bark, Latin).

Comments, Adult females of Subcorticoccus can

be distinguished from those of other Australian

genera of Eriococcidae by the combination of a

tapered abdomen, a mediolongitudinal band of

microtrichia on the head and anterior thorax, a

pair of oval frontal lobes, very small legs relative

to the size of the body, a ventral and noncellular

anal ring, all body setae flagellate and mostly less

than 15 um long, very slender macrotubular

ducts distributed over both dorsum and venter,

ventral bands of clustered quinquelocular pores

on the posterior abdomen and sometimes

scattered quinquelocular pores on the body

margin, and the absence of anal lobes and

microtubular ducts.

First-instar nymphs and a single prepupal male

are known only for S. beardsleyi sp. nov. The

male nymph is too poorly preserved to describe

adequately.

Key to adult females of Subcorticoccus

l. Antennae usually 7 (rarely 6) segmented; legs of typical form but reduced in

size; quinquelocular pores densely scattered around margins of entire body...

vee arin S. murrindindi sp. nov.

Antennae usually 6 (rarely 5) segmented; legs highly reduced so that com-

bined femur, tibia, tarsus and claw resembles an elongate cone; quinqueloc-

ular pores either absent from margins of body or sparsely scattered on

abdominal margins … …………… ….

NEW SUBCORTICAL COCCOIDS ON EUCALYPTUS 243

Je Quinquelocular pores sometimes present on margins of body, at least on

abdomen, in dense bands ventrally on last 4 abdominal segments (V_ VIII)

plus a few pores on LV wo...

wter en D ANTARI S, huonammnis sp. nov.

— Quinquelocular pores absent from margins of abdomen, in sparse bands ven-

trally on last 4 abdominal segments (V VHI) only ....... 8. beardslevi sp. nov.

Subcorticoccus beardsleyi sp. nov.

Figures | 2

Type material. Holotype: adult female (1.6 mm long,

largest of 3 females on slide), Victoria, near Heathcote,

15 Mar 1972, ex Eucalyptus macrorhyncha, under twig

bark, J.W. Beardsley (ANIC)

Paratypes: 17 adult females (11 slides), | prepupal

male (on slide with 2 adult females) and 4 first-instar

nymphs (each on slide with | or 2 adult females), same

data as holotype (2 slides in ANIC, 8 slides in BPBM,

l slide in NMV T-17318).

Description of adult female (measurements based

on 10 slide-mounted specimens) (Fig. 1). Body

1.1-1.7 mm long, 0.6-1.0 mm wide; segmenta-

tion distinct only on posterior half of abdomen.

Eyespots 10-15 um wide. Antennae (Fig. la)

43-70 um long, with 6 (rarely 5) segments.

Frontal lobes irregularly oval, each 30 80 um

long, 25-50 um wide. Labium 50-70 um long,

60-70 um wide across base. Clypeolabral

shield 120-170 um long, 105-140 pm maximum

width. Spiracles (Fig. le): mesothoracic 35 56

um long, 14-25 um wide; metathoracie 39-60 um

long, 14-27 um wide. Legs (Figs 1b,d) 45 65 pm

long, with segments highly reduced: tarsal

digitules 8-20 um long; claw digitules 8 15 um

long; claw denticle not discernible. Apical

setae 20-40 um long; anal ring (Fig. 1g) 14-20

um in diameter with | pair of anal ring setae

8-10 um long; suranal setae (Fig. 1g) 8-10 um

long.

Dorsum with mediolongitudinal

microtrichia (Fig. Lh) on head and anterior thorax

140-260 um long, widest (25-33 um) posteriorly;

flagellate setae, 4-10 jum long, sparsely dis-

tributed across all segments, longest on posterior

abdominal segments; macrotubular ducts (Fig.

le) 10-16 (mostly 13) um long, 1-2 pm wide,

scattered across all segments; quinquelocular

pores absent.

Venter usually with a few microtrichia in indis-

tinct cluster between antennae; flagellate setae

3-12 um long, sparsely distributed across all seg-

ments, a pair of longer setae (13-18 um) near

vulva; macrotubular ducts (Fig. le) 10-14

(mostly 13) um long, 1-2 pm wide, scattered

across all segments; quinquelocular pores about 5

um in diameter (Fig. 1f) in sparse bands on

abdominal segments V to VIII, absent from

band of

margins of body, a small cluster of 3 8 pores

(mostly quinquelocular, rarely trilocular) (Fig.

le), each 3-4 pm in diameter, at opening of each

spiracle and a few in each spiracular furrow.

Description of first-instar nymph (measurements

based on 4 slide-mounted specimens) (Fig. 2).

Body 0.32-0.41 mm long, 0.11 0.14 mm wide;

segmentation indistinct; microtrichia absent. Lyc-

spots 7-8 um wide. Antennae 75 80 um long,

with 6 segments; trichoid sensilla 8 30 um long,

distributed as follows: 3 on 1, 2 on H, 2 on HI, 0

on IV, 2 on V, 3 on VI; pegs 4-13 um long, dis-

tributed as follows: Lon IV, | on V,3 on VL 3 4

basiconie or coeloconie sensilla, 5 6 ym long, on

VI; a single campaniform sensillum on apical part

of I. Labium conical, segmentation not apparent,

35-43 um long, 30 34 um wide across base.

Clypeolabral shield 75 90 um long, 42 50 um

maximum width. Spiracles about 10 um long, 5-7

um wide. Legs of typical form; tarsal digitules

capitate (Fig. 2b), 15-25 um long; claw digitules

capitate, 10-15 um long; each claw with small

denticle near apex. Anal lobes absent;

apical setae 52 60 um long: anal tube, about 10

um long, with simple, ventral anal opening about

3 um in diameter without setae; suranal setae

flagellate, about 6-7 um long.

Dorsum with short cone-like setae, 1-2 um

high and 1-2 um wide with a base 2.0 3.5 um in

diameter (Fig. 2e), distributed in a transverse row

of 6 setae per abdominal segment, segmentally

arranged on thorax, scattered on head, with setae

of adjacent body segments lining up to form 3

pairs of longitudinal rows: | medial, | submedial

and | submarginal, with submarginal setae largest

especially on abdomen; macrotubular ducts,

microtubular ducts and quinquelocular pores

absent,

Venter with flagellate setae 3-7 pm long, 3

pairs on head near antennae, | seta near base of

each fore leg, and abdominal setae distributed in

transverse rows of 6 setae on each segment with

setae of adjacent segments lining up to form 3

pairs of longitudinal rows; macrotubular ducts

and microtubular ducts absent; a single trilocular

pore (Fig. 2a), 3 um in diameter, adjacent to each

spiracle.

Etymology. This species is named in honour of

Professor Jack Beardsley who collected all known

P. J. GULLAN

244

Figure 1. Adult female of Subcorticoccus beardsleyi sp. nov. Enlargements show: a, antenna; b, fore leg:

c, metathoracic spiracle and associated pores; d, hind leg; e, macrotubular duct; f, abdominal quinquelocular

pore; g, anal ring with suranal setae lateral to ring (NB. all of these structures are ventral); h, microtrichia

from dorsal band on head and anterior thorax.

NEW SUBCORTICAL COCCOIDS ON EUCALYPTUS 245

Figure 2. First-instar nymph of Subcorticoccus beardsleyi sp. nov. Enlargements show: a, trilocular pore;

b, midleg claw and tarsal apex; c, dorsal seta.

246 P. J. GULLAN

specimens of this species and other

Australian eriococcids.

many

Subcorticoccus huonamnis sp. nov.

Figure 3

Type material. Holotype: adult female (3.4 mm long),

Tasmania, Huon River near Judbury, 24 Oct 1978, ex

Eucalyptus regnans, under bark, D.J. Williams (ANIC),

Paratypes: 28 adult females (9 slides), same data as

holotype (2 slides in ANIC, 7 slides in BMNH).

Description of adult female (measurements based

on 10 slide-mounted specimens). Body 1.2-3.9

mm long, 1.1-2.7 mm wide; segmentation indis-

tinct except on posterior abdomen. Eyespots

25 35 um wide. Antennae (Fig. 3a) 60-95 um

long, with 6 (rarely 5) segments. Frontal lobes

each 70-130 um long, 40-90 um wide. Labium

60-85 mm long, 70-100 mm wide across base.

Clypeolabral shield 140-230 um long, 125-200

um maximum width. Spiracles (Fig. 3c):

mesothoracie 42 58 um long, 20 30 um wide;

metathoracic 45 70 um long, 23 30 um wide.

Legs (Figs 3b,d) 70 120 um long, with segments

reduced and coxa mostly membranous; tarsal dig-

itules 20-32 um long: claw digitules 15-25 um

long; claw with small denticle discernible near

apex on some specimens. Apical setae 30-50 um

long: anal ring (Fig. 32) 23-29 mm in diameter,

with 0-1 pair of anal ring setae, 10-18 um long;

suranal setae (Fig. 3g) 10-12 um long.

Dorsum with mediolongitudinal

microtrichia (Fig. 3h) on head and anterior thorax

280-600 um long, widest (40-100 um) in poster-

ior two-thirds; flagellate setae, 5-10 um long,

sparsely distributed across all segments, longest

on posterior abdominal segments; macrotubular

ducts (Fig. 3e) 10-15 um long, 1.5-2.0 um wide,

scattered across all segments; quinquelocular

pores absent.

Venter with microtrichia in short, mediolongi-

tudinal cluster 60-100 um long, 25-38 um wide,

between antennae; flagellate setae 5-12 um long,

sparsely distributed across all segments, one pair

of longer setae (15-23 um) near vulva; macro-

tubular ducts (Fig. 3e) 10-15 um long, 1.5-2.0

um wide, scattered across all segments; quinque-

locular pores 4-5 um in diameter sometimes pre-

sent on margins of body, at least on abdomen,

larger pores about 5-6 um in diameter (Fig. 3f) in

dense bands on abdominal segments V to VIII, a

few scattered or clustered on IV, plus a loose

cluster of 13-25 pores (Fig. 3c), each 3-5 um in

diameter, at opening of each spiracle and a few in

each spiracular furrow.

band of

Etymology. After the type locality on the Huon

River, Tasmania, and amnis, meaning river,

Latin.

Subcorticoccus murrindindi sp. nov.

Figure 4

Type material. Holotype: adult female (3.5 mm long),

Victoria, c. 10.5 km NE of Toolangi, near Murrindindi

River, off Murrindindi Road, 31 Oct 1978, ex Euca-

lyptus regnans, under bark, P.J. Gullan and A. Smith

(ANIC).

Paratypes: 8 adult females, same data as holotype

(4 slides in ANIC, 2 slides in BPBM, 2 slides in

BMNH).

Description of adult female (measurements based

on 7 slide-mounted specimens). Body 3.4-3.9

mm long, 2.1-2.4 mm wide; segmentation dis-

tinct, at least on abdomen. Eyespots 27-35 um

wide. Antennae (Fig. 4a) 85-125 um long, with 7

(rarely 6) segments. Frontal lobes oval, each

65-155 um long, 50-65 um wide. Labium 70-85

mm long, 100-110 um wide across base. Clypeo-

labral shield 150-170 um long, 140-160 um max-

imum width. Spiracles (Fig. 4c): meso-

thoracic 55-72 ptm long, 30-35 um wide;

metathoracic 60-75 um long, 30-35 um wide.

Legs (Figs 4b, d) of typical form but small,

112-160 um long, tibia and tarsus of each leg

fused, tibia + tarsus about equal in length to femur

of each leg; tarsal digitules 20-35 um long; claw

digitules 13-20 um long; claw sometimes with a

barely discernible denticle. Apical setae 35-40

um long: anal ring (Fig. 4g) 27-33 um in

diameter, with | pair of anal ring setae 7-13 um

long: suranal setae (Fig. 4g) 12-20 pm long.

Dorsum with mediolongitudinal band of

microtrichia (Fig. 4h) on head and anterior thorax,

600-800 um long, widest (100-130 um) for pos-

terior half to two-thirds; flagellate setae, 5-16 um

long, sparsely distributed across all segments,

longest on posterior abdominal segments; macro-

tubular ducts (Fig. 4e) 12-15 um long, 1.0-1.5

mm wide, scattered across all segments; quinque-

locular pores 4-5 um in diameter on margins of

body (Fig. 4c).

Venter with microtrichia in indistinct, medio-

longitudinal cluster about 50 um long, 25-30 um

wide, between antennae; flagellate setae 7-22 um

long, sparsely distributed across all segments, a

pair of longer setae (30-33 um) near vulva;

macrotubular ducts (Fig. 4e) 12-15 um long,

1.0-1.5 um wide, scattered across all segments:

quinquelocular pores 4-5 um in diameter (Fig.

4c) densely scattered on margins of body, larger

pores about 5—6 um in diameter (Fig. 4f) in dense

NEW SUBCORTICAL COCCOIDS ON EUCALYPTUS

Figure 3. Adult female of Subcorticoccus huonamnis sp. nov. Enlargements as for caption of Fig. 1 except

that, in c, the enlarged quinquelocular pore represents both pores near the spiracle and on the body margin.

248 P. J. GULLAN

Figure 4. Adult female of Subcorticoccus murrindindi sp. nov. Enlargements as for caption of Fig. 1 except

that, in c, the enlarged quinquelocular pore represents both pores near the spiracle and on the body margin.

NEW SUBCORTICAL COCCOIDS ON EUCALYPTUS 249

bands on abdominal segments IV to VIII, a loose

cluster of 15-25 pores, each 4-5 pm in diameter,

at opening of each spiracle and in an irregular row

in each spiracular furrow.

Etymology. For the type locality near Toolangi,

Victoria, a noun in apposition.

Comments. These eriococcids were collected

from under pieces of decorticating bark on their

host trees. A single adult female from Euca-

lyptus acmenioides in Brisbane (A.R. Brimble-

combe No. SC2207, 18.iv.1948, ANIC via H.M.

Brookes) is similar to the specimens from

Murrindindi Road but is only one third the body

size and differs in the shape of the claws, which

are distinctly hooked with a small denticle near

the apex, and the shape of the frontal lobes. This

female probably represents a fourth species, but

its description must await the collection of further

specimens.

Discussion

In addition to Subcorticoccus and Phacelococcus

several other Australian eriococcid genera also

have species with stationary females that live in

or under eucalypt bark. The coccids are either

tightly fitted into crevices or in blister galls as in

Floracoccus Beardsley, Ourococcus Fuller and

Sphaerococcopsis Cockerell (Fuller, 1899;

Beardsley, 1974a, b), are under bark on twigs as

in a few species of Lachnodius Maskell (J.W.

Beardsley, pers. comm.), in resinous secretion as

in Olliffia Fuller (P.J. Gullan, unpubl. data), or in

bark crevices under felted tests as in a few

species of Eriococcus (Froggatt, 1921). Outside

Australia, some eriococcid genera, including

Capulinia Signoret, Cryptococcus Douglas,

Ovaticoccus Kloet and Xerococcus Ferris, have

one or more species that either live under bark or

in bark crevices (D.R. Miller, pers. comm.). Some

of these taxa, particularly species of Ovaticoecus

(Miller and McKenzie, 1967), display some

similarity to Subcorticoccus, for example in pos-

sessing a reduced anal ring and small legs. The

morphological reductions that accompany many

bark-dwelling eriococcids make it difficult to

estimate their phylogenetic relationships using

cuticular features; it is hoped that their relation-

ships may be more accurately estimated by future

cladistic analysis of molecular data.

Acknowledgments

Gordon Nishida and Jon Martin kindly arranged

the loan of specimens from the Bishop Museum

and the Natural History Museum, respectively,

The financial support of the Australian Biological

Resources Study (ABRS) is gratefully acknowl-

edged. Thanks to Peter Cranston and Dug Miller

for helpful comments on the manuscript and to

Robert Hoare for advice on the Latin used to form

the new names. The Zoology Department of the

University of Western Australia generously pro-

vided laboratory space while this paper was writ-

ten, and Alan Muir at the Division of Botany and

Zoology kindly made the stencil that was used to

draw the tiny circles on the figures.

References

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from Australian Eucalyptus (Homoptera), Pro-

ceedings of the Hawaiian Entomological Society

21: 325-328.

Beardsley, J.W. Jr, 1974b. A review of the genus

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two new species (Homoptera: Coccoidea).

Proceedings of the Hawatian Entomological

Society 21: 329-342,

Froggatt, W.W., 1921. A descriptive catalogue of

the scale insects (“Coccidae”) of Australia. Part LL

Science Bulletin, Department of Agriculture, New

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Memoirs of Museum Victoria 57(2): 251-259 (1999)

A NEW SPECIES OF OGYRIS ANGAS (LEPIDOPTERA: LYCAENIDAE)

FROM SOUTHERN ARID AUSTRALIA

R. P. FIELD

Museum Victoria, PO Box 606E, Melbourne, Vic. 3001, Australia

(rficld@mov.vic.gov.au)

Abstract

Field, R.P., 1999, A new species of Ogyris Angas (Lepidoptera: Lycaenidae) from southern

arid Australia. Memoirs of Museum Victoria 57; 251-259.

Ogyris subterrestris sp. nov. is described with the nominal subspecies O. s. sublerresiris ssp.

nov. from northwestern Victoria and the subspecies Ogyris subterrestris petrina ssp. nov. from

near Kalgoorlie, Western Australia. Adults, male and female genitalia, and first instar larvae

are figured and characters to distinguish adults and larvae of Ogyris subterrestris trom O. idmo

Hewitson are discussed.

Introduction

The genus Ogyris Angas, is an Australasian

genus in the tribe Ogyrini (Eliot, 1973; Edwards,

1996). There are 15 described species in the

genus, 12 occurring in Australia and three in

Papua New Guinea. Of the Australian species, the

larvae of ten are known to feed on mistletoes

(Loranthaceae), and one feeds on the root para-

sitic plants in the genera Choretrum and

Leptomeria (Santalaceae). The remaining species

is Ogyris idmo (Hewitson), the largest and one of

the rarest species in the genus. The life history

of this species is unknown but the larvae are pre-

sumed to be predatory on ants (Field, 1997).

Many species of Ogyris have an obligatory asso-

ciated with ants whereas the others have a facul-

tative relationship. For species in which the life

history is known, these associations are presumed

to be mutualistic. By day the larvae shelter with

the ants in crevices, under bark or underground,

emerging at night with ants in attendance, to feed.

The larvae will usually pupate in these same or

sheltered sites nearby.

Populations of O. idmo show differing wing

colours and patterns, and morphologically (wing

shape, antennal segments) it is a very

variable species. It has been collected from

western Victoria, southern South Australia, and

southwestern Western Australia, from Cape Arid

National Park to near Geraldton. Few specimens

have been recorded outside Western Australia

since 1950 but recently the species has been

locally common near Perth, and at Cape Arid

National Park (Field, 1990, 1992) and it has

been rediscovered in South Australia (Hunt

et al., 1998). Specimens from Mildura (Victoria),

Broken Hill (New South Wales) and near

Kalgoorlie (Western Australia), initially incor-

porated within the O. idmo complex, are here

described as a new species of Ogvris with two

new subspecies.

Material examined is stored in Museum

Victoria, Melbourne (NMV), South Australian

Museum, Adelaide (SAM), Australian National

Insect Collection, CSIRO, Canberra (ANIC),

Australian Museum (AM), Natural History

Museum London (BMNH) and the private collec-

tions of the author (RPFC), R. Hay (RHC), M.

Moore (MMC), and B.H. Vardy (BH VC).

Key to females of Ogyris idmo (Hewitson) group

L Upperside of forewing with 2 black bars at proximal end of postmedian

cream patch and with subapical region to distal end of postmedian cream

patch dark brown.

haft. O. subterrestris sp. nov. … 2

— Upperside of forewing with black patch at proximal end of postmedian

cream patch and with subapical region to distal end of postmedian cream

patch black, ans errans oaavorreneneneren

EERE ee O. idmo … 3

R.P. FIELD

Upperside of forewing with postmedian cream patch regular, oval, often

faint and small; upperside of hindwing with central blue patch extending

anteriorly to M, and reaching base…

EN teneerste tects free A: O. subterrestris subterrestris ssp. nov. (figs 1, 2)

Upperside of forewing with postmedian cream patch irregular, distinctly

divided by M, forming a small faint cream patch distally between M, and

CuA); upperside of hindwing with central blue patch not extending

anteriorly to M} and not reaching base nnen eenen

ETE O. subterrestris petrina ssp. nov. (figs 5, 6)

Upperside of wings brown with broad basal area bright blue or bluish-purple

and extending to postmedian cream patch on the forewing..… 4

Upperside of wings brown with broad basal area purple but not extending to

postmedian cream patch on the forewing…………… O. idmo idmo (figs 9, 10)

Upperside of wings with broad basal area bluish-purple…

EEEN EN O. idmo halmaturia (figs 13, 14)

Upperside of wings with basal area bright blue nnn.

Mr Bee VDATA desetero A G O. idmo (Mt Ragged form) (figs 17, 18)

Key to males of Ogyris idmo (Hewitson) group

Forewing with termen at right angles to inner margin; upperside of wings

with fine black line at termen … O. subterrestris sp. nov. … 2

Forewing with termen obtuse to inner margin; upperside of wings with nar-

row black band to termen nnee O. idmo … 3

Upperside of wings predominantly bluish-purple with strong bronze sheen to

brown marginal areas; underside of hindwing with well defined markings,

the grey patches contrasting the black-edged brown patches.

EE AN ND O. subterrestris subterrestris ssp. nov. (figs 3, 4)

Upperside of wings predominantly brownish-purple without strong bronze

sheen to brown marginal areas; underside of hindwing with poorly defined

brown markings and without contrasting grey patches.......

OLEATE EDER NCN DENS O. subterrestris petrina ssp. nov. (figs 7, 8)

Forewing termen CONVEX... O. idmo halmaturia (figs 15, 16)

Forewing termen straight. inte nare eustaatrheevan cacansatdnsr ain tas cen benauwde 4

Upperside of forewings brownish-purple… … … … O. idmo idmo (figs 11, 12)

Upperside of forewings dark purple. O. idmo (Mt Ragged form) (figs 19, 20)

Ogyris subterrestris sp. nov.

Types. See types of nominal subspecies.

Diagnosis. Females with upperside of forewing

with 2 black bars at proximal end of postmedian

cream patch and with subapical region distal to

cream patch dark brown, males with termen at

right angles to inner margin and upperside of

forewing with fine black line at termen and with

faint to strong bronze sheen to brown areas: first

instar larvae with a pair of prominent mesotho-

racic dorsal spines and a pair of long, posteriorly

curved dorsal spines on each of abdominal

segments 6 and 7.

Remarks. The syntypic series of O. idmo and the

holotype of O. orontas (Hewitson) (male), a

junior subjective synonym of O. idmo, are held in

the Natural History Museum, London. Colour

images of these specimens have been examined.

Although a female specimen of O. idmo in the

Natural History Museum has the word “type” on

a label, Hewitson (1862) described the species

from two (female) specimens. The syntype (male)

of O. halmaturia (Tepper) is in the South Aus-

tralian Museum and has been examined. Tepper

(1890) described the species from three speci-

mens, two small specimens being the males and a

larger specimen, which he believed to be the

female. However, the latter was a male and the

smaller specimens were male O. otanes (C. Felder

and R, Felder). Waterhouse (1903a) synonymised

part of halmaturia (presumably referring to the

large male) with O. idmo and part with O. otanes.

Waterhouse (1903b) later synonymised halma-

turia with ofanes and made no mention of hal-

maturia under idmo. Tepper’s female (the large

male) is the first specimen described and thus

halmaturia can be synonymised with idmo but

not also with ofanes. This specimen is thus

A NEW SPECIES OF BUTTERFLY OF THE GENUS OGYRIS 253

designated the lectotype of O. halmaturia and is

so labelled. O. waterhouseri (Bethune-Baker)

was described, but not figured, from Victoria

(Bethune-Baker, 1905) and subsequently figured,

| synonymised and treated as a subspecies of O.

idmo by Waterhouse and Lyell (1914). O. water-

houseri has generally ar treated as a junior sub-

jective synonym of O. i. halmaturia (Common

and Waterhouse, 1981) but both McCubbin

(1971) and D’Abrera (1971) retained water-

houseri as a distinct subspecies. This paper recog-

nises waterhouseri as a junior subjective syn-

onym of halmaturia. The type

O. waterhouseri could not be located but presum-

ably consisted of one female and at least two

males (based on size ranges given in the descrip-

tion). The original description, which highlighted

the “strongly arched” termen, Waterhouse and

Lyell’s figuring of the species along with locality

data, clearly separates, O. waterhouseri from

O. subterrestris sp. noy.

Etymology. Sub- and terrestris (Latin), under-

ground, referring to the larval and pupal stages

thought to be completely subterranean.

Ogyris subterrestris subterrestris ssp. nov.

Figures 1—4

Types. Holotype: female, Pink Lakes, Murray-Sunset

NP, 15 km N Lima, Victoria, 35°03.45'S, 142°43.13'E,

20.x.1996, R.P. Field (NMV T-17264).

Paratypes (all Victoria): | male, same data as holo-

type (NMV T-17265); 1 female, 150 m NW of Ring

Road at 1 km NNE of junction with Grub Tk, Pink

Lakes, Murray-Sunset NP, F. Noelker, 18.11.1996

(NMV T-17267); 11 males, 2 females, same data as

holotype; | male, Pink Lakes, Murray-Sunset NP, 15

km N of Lima, 12.iv.1996, R.P. Field (RPFC); 1

female, Mildura 16.x.1972, B. H. Vardy, | male, Mil-

dura, 26.x.1972, B. H. Vardy (ANIC); 1 male, 2

females, Mildura, 26.x.1972, B. H. Vardy (BHVC).

Other material examined. Victoria: | female, Lake

Waltah, xi. 1918, F.R. Spry, G.A. Waterhouse Collec-

tion (AM KL 20211).

New South Wales:

(NMV LEP 6272).

South Australia: 1 female, Koonibba Mission, nr

Ceduna; 1 male, Loxton, 1986, Hudson (SAM); 1 male,

7.5 km NW Ramco, 13,x.1993, 34°07'38"S,

139°53'22"E, P.J. Peile; 3 females, 7.5 km NW Ramco,

34°07'38"S, 139°53'22"E, 12.11.1994, R.P. Field;

1 male, 1 female, 7.5 km NW Ramco, 13.11.1994, R.P.

Field: 1 female 7.5 km NW Ramco, 34°07'38"S,

139°53'22"E, 14.11.1994, R.P. Field; 4 males,

3 females, 7.5 km NW Ramco, 34°07'38"S,

139°53'22"E, 30.x.1994, R.P. Field (all RPFC);

] female, 10.v.1992, Qualco; 1 female, 19.1.1993,

1 male, Broken Hill, xii.1912

series of

Qualco; 4 males, | female, 25.11.1993, Qualco; | female

26.11.1993, Qualco; 2 males, 19.iii.1993, Qualco;

| female, 20.i11.1993, Qualco; 1 female, 20.iv.1993,

Qualco; 1 female, 28.ix.1993, Qualco; | male,

2.x.1993, Qualco (all MMC),

Description. Female. (figs 1, 2). Antennal length

(of holotype) 7.1 mm, flagellum 37 segments,

brown and bronze with segmental bands narrowly

banded black with lateral white scales; club short,

apically broad, rounded, tipped orange. Head, pal-

pus, thorax and abdomen dorsally brown with

white scales, ventrally white with brown scales;

legs speckled brown and white; ventral surface of

head, thorax, base of abdomen, all of coxae with

long white hair scales; dorsal surface of thorax

with long bronze hair scales. All tibiae of equal

length, first and third femora equal length of tib-

iae, mid femur much longer than tibiae. Forewing

length (of holotype) 21,0 mm, apex weakly acute,

rounded, termen slightly convex; above central

area from base to subterminal area and from sub-

costal area to inner margin royal blue, remaining

areas and veins browny bronze except for narrow

black bar proximal to discocellulars, wide black

bar distal to discocellulars between M, and M,

proximally edging a postmedian cream patch

between M, and M, which extends faintly

towards M, and C uA,. Hindwing termen crenated

convex; above central area from base to subtermi-

nal area and from M, to CuA, royal blue, remain-

ing areas, veins and bands at discocellulars

browny bronze, long tan hair scales throughout

cell and from base to termen at CuA, and to inner

margin. Cilia of both wings white, browny bronze

at veins. Beneath forewing base colour grey, dark

brown cell, extending to basal fifth of area

between M, and Ms, basal third of M, to CuA;,

basal quarter of CuA, to CuA,, anid circular

brown patch between "action of cubitus and

CuA, and 1A + 2A narrowly edged near LA+2A

with ‘white and iridescent blue scales; cell with 2

iridescent light blue bands 1 median band from

proximal end of Rs to base of CuA,, the other

subbasal parallel to radius with bend towards

cubitus distally; 2 fine white inner subbasal lines

between radius and cubitus; prominent post-

median cream band extending from M, to nearly

CuA, slightly stepped towards termen between

M, and CuA,; broad subterminal brown band

from costa to CuA,, twice as wide at costa than at

CuA, edged dark brown; prominent white scales

in subcostal, subapical and apical areas. Hind-

wing base colour grey with irregular brown or

white patches edged dark brown; 2 subbasal dark

brown lines, between costa and Sc +R, subparal-

lel to basal Sc + R, then bending back to costa

A NEW SPECIES OF BUTTERFLY OF THE GENUS OGYRIS

subparallel to mid Sc + R} and between Se + R,

and cubitus displaced towards termen between Sc

+ R, and radial sector; submedian lines dark

brown from Sc + R, to cubitus (2), radial sector to

mid cell, cubitus (nearly) to anal vein; median

lines from Se + R1 to Rs (2), Rs to M}, M} to M,

(2) 1 either side of discocellulars, M; to" CuA,,

CuA, to CuA;, 1A+2A to anal vein; postmedian

dark brown lines between Rs and M}, M} and M,

(2), dark brown edged v markings between M

and CuA, and CuA, and CuA,; brown areas basal

fifth of M, to CuA, extending proximally into

cell, basal quarter of CuA, to CuA, extending

proximally into cell and spat distal to junction of

CuA, and CuA, between CuA, and IA + 2A

edged dark brown; costa and termen edged dark

brown,

Genitalia (Fig, 21): Apophyses anteriores long,

slender; papillae anales broadly acute, setose;

ostium bursae broad, weakly sclerotised; ductus

bursae not sclerotised, membranous, broad, mod-

erately long, expanding to rounded, membranous

corpus bursae.

Male. (figs 3, 4). Antennal length 6.6-8.3 mm

(mean 7,6 mm, n=20), flagellum 34-41 segments

(mean 38, n=20). Colour of head, palpi, antennae,

thorax, abdomen and legs similar to female.

Forewing length 19,5-23.5 (mean 21.7 mm,

n=20), apex acute, termen straight or slightly con-

cave, above similar to female but without post-

median cream patch and darker purple blue;

remaining areas, veins and narrow band at disco-

cellulars bronze. Hindwing above similar to

female but darker purple blue central area and

bronze submarginal areas; beneath similar t

female (holotype) but postmedian cream band of

forewing more grey and median iridescent light

blue band in cell often broader, subbasal irides-

cent band often more extensive forming a cross

and an additional narrow basal iridescent band in

cell; subterminal brown band narrower than in

female but extending past CuA, often to CuA,,

Genitalia (fig. 22): Vinculum-tegumen ring

oval, saccus small, blunt; uncus lobes narrow,

bases close dorsally, setae numerous, brachia

long, U-shaped, curved laterally inwards, broad

basally tapering to a blunt point; valva broadly tri-

angular, numerous long setae on posterior margin

concentrated near dorsal and ventral regions;

juxta prominent, v-shaped; aedeagus long,

straight, basal fifth swollen, zone broad; post-

zonal sheath length longer than half the pre-zonal

sheath, broad, displaced ventrally near base with

cornuti well developed near zone, apically round.

Variation. Female, size of the cream spot on the

forewing varies from almost absent to a small

patch; flagellum 36-44 segments (mean 39,

n=20), length 6.8-8.3 mm (mean 7.7 mm, n=20),

wing length 20-25.5 mm (mean 22.7 mm, n=20).

Distribution. In Victoria the subspecies occurs in

the Murray Sunset National Park, near Linga and

was recorded from Mildura, in October 1972 (B.

Vardy collection), A single specimen was

recorded from Broken Hill, New South Wales, in

December 1912 (Museum Victoria). In South

Australia it occurs locally in a small remnant

roadside strip of mallee vegetation, near Ramco,

(Moore, 1999) and has also been recorded near

Loxton and near Ceduna.

Ogyris subterrestris petrina ssp. nov.

Figures 5—8

Types. Holotype: female, Western Australia: female,

Lake Douglas, 12 km SW of Kalgoorlie. 12.xi. 1989,

R.P. Field (NMV T-17268),

Paratypes: 1 male, Lake Douglas, 12 km SW of Kal-

goorlie, 4.x. 1991, R.P. Field (NMV T-17269). 1 male,

1 female, Lake Douglas, 12 km SW of Kalgoorlie,

L.ii.1982, A.J. Graham 12 males, 3 females, Lake Dou-

glas, 12 km SW of Kalgoorlie, 16,.x,1986, A.J. Graham;

4 females, Lake Douglas, 12 km SW of Kalgoorlie,

17.x,1986, A.J. Graham; 1 male, Lake Douglas, Kalgo-

orlie, 22.x1.1991, L. R. Ring, (all ANIC); 1 female,

Lake Douglas, 12 km SW of Kalgoorlie, 5.x.1987, R.P.

Field; 3 males, 5 females, Lake Douglas, 12 km SW of

Kalgoorlie, 6.x.1987, R.P. Field; 2 males, Lake Dou-

glas, 12 km SW of Kalgoorlie, 7.x.1987, R.P. Field; 1

male, | female, Lake Douglas, 12 km SW of Kalgoor-

lie, 9.xii. 1989, R.P. Field; 2 males, Lake Douglas, 12

km SW of Kalgoorlie, 10.xii. 1989, R.P. Field; 3 males,

Lake Douglas, 12 km SW of Kalgoorlie, 11.xii.1989,

R.P. Field; 2 males, 4 females, Lake Douglas, 12 km

Figures 1-4, Ogyris subterrestris subterrestris ssp. nov. 1, 2, holotype female upperside and underside; 3, 4,

paratype male upperside and underside.

Figures 5-8, Ogyris subterrestris petrina ssp. nov. 5, 6, holotype female upperside and underside; 7, 8,

paratype male upperside and underside.

Figures 9-12, Ogyris idmo idmo. 9, 10, female upperside and underside; 11, 12, male upperside and under-

side.

Figures 13-16, Ogyris idmo, Halmaturia. 13, 14, female upperside and underside; 15, 16, male upperside and

underside.

Figures 17-20, Ogyris idmo (Mt Ragged form). 17, 18, female upperside and underside; 19,

side and underside. Scale lines 1 mm.

20, male upper-

SW of Kalgoorlie, 12.xii.1989, R.P, Field; 1 female,

Lake Douglas, 12 km SW of Kalgoorlie, 3.x.1991, R.P.

Field; 2 males, 2 females, Lake Douglas, 12 km SW of

Kalgoorlie, 4.x.1991, R.P. Field, (all RPFC); 1 female,

13.x11, 1986, R. H. (WAM 96/177); 2 males, 1 female,

14.x11.1986, R.H. (WAM 96/175, 96/176, 96/178); 1

female, Lake Douglas, 30.xi.1985, R.H.; 1 male, Kal-

goorlie, 14.xii.1986, R.H; 1 male, Lake Douglas,

28.ii. 1988, R.H.; 1 female, Lake Douglas, 3.x.1991, R,

HL; 4 males, Lake Douglas, 4.x.1991, R.H,; 1 female,

Lake Douglas, 5.x.1991, R. H.; 2 males, 2 females,

Lake Douglas, 6.x.199], R.H.; 1 male, Lake Douglas,

8.x.1991, R, H.; 1 male, Kalgoorlie, 9.x.1991, R.H. (all

RHC),

Other material examined. Western Australia: 1

female, S. W. Australia, Kalgoorlie district, W. Subi-

aco, 23.x.1911, W.J. Brooks (BMNH).

Description. Female. (figs 5, 6). Antennal length

(of holotype) 7.9 mm, flagellum 41 segments,

brown and bronze with segmental bands narrowly

banded black with lateral white scales; club short,

apically broad, rounded, tipped orange. Head,

palpus, thorax and abdomen dorsally brown with

white scales, ventrally white with brown scales:

legs speckled brown and white; ventral surface of

head, thorax, base of abdomen, all of coxae with

long white hair scales; dorsal surface of thorax

with long bronze hair scales. All tibiae of equal

length, first and third femora equal length of

tibiae, mid femur much longer than tibiae,

Forewing length (of holotype) 21.0 mm, apex

weakly acute, rounded, termen slightly convex;

above central area from base to median area in

cell and subterminal area at CuA, and from sub-

costal area to inner margin purple blue: discocel-

lulars and veins browny bronze, remaining areas,

brown with scattered browny bronze scales

except for narrow brown black bar proximal to

discocellulars and a narrow brown black bar

distal to discocellulars between M, and M, prox-

imally edging a large postmedian cream patch

between MI and M3 which extends faintly

towards CuAl. Hindwing termen crenated con-

vex; above central area from base to subterminal

area and from M, to CuA, purple blue, discocel-

lulars and veins browny bronze, remaining areas,

brown with scattered browny bronze scales, long

tan hair scales throughout cell and from base to

termen at CuA, and to inner margin. Cilia of both

wings white, brown at veins. Beneath forewing

base colour grey with dark brown cell, extending

to basal sixth of area between M, and Ms, basal

quarter of M, to CuA,, basal third of CuA, to

CuA,, cell with median iridescent light blue patch

and 2 iridescent light blue bands 1 sub-medial and

| subbasal, 2 fine white inner subbasal lines

between radius and cubitus; prominent post-

median cream band extending from M, to nearly

R.P. FIELD

CuA, slightly stepped towards termen between

M, and CuA,; broad subterminal brown band

from costa to midway between CuA, and CuA,,

twice as wide at costa as at CuA, edged dark

brown; prominent white scales in subcostal, sub-

apical and subterminal areas. Hindwing base

colour grey and flecked with white scales with

irregular brown lines; 2 subbasal dark brown

lines, between the costa and Sc + R} and back to

costa subparallel to basal Sc + R} and between

radial sector and cubitus, subbasal brown ring

between anal vein and inner margin; submedian

dark brown lines between Se + R, and radial

sector (2), radial sector and cubitus (2), cubitus to

I A+2A; median dark brown lines from Sc + R, to

Rs (2), M‚ to M, (2) | either side of discocellu-

lars, LA+2A to anal vein median dark brown rings

(3), in cell half the width of the cell at the base of

CuA,, between M, and CuA, and between CuA,

and CuA,, median brown spot distal to junction

of CuA, and CuA, between CuA, and 1A + 2A;

postmedian dark brown lines Rs to M (2), M, to

M, (2), M, to CuA CuA, to CuA,, CuA, to

1A+2A; termen edged dark brown.

Male. (figs 7, 8). Antennal length 7.3-8.5 mm

(mean 8.1, mm, n=19), flagellum 37-43 segments

(mean 40, n=20) colour of head, palpi, antennae,

thorax, abdomen and legs similar to female.

Forewing length 21.5-24.5 mm (mean 23.4 mm,

n=19), apex acute, termen straight or slightly con-

cave, above similar to female but without post-

median cream patch and central area browny

purple. Hindwing above similar to female but

central area browny purple and browny bronze

submarginal areas; beneath similar to female but

postmedian cream band of forewing more grey

and iridescent light blue bands in cell often

broader; subterminal brown band narrower than

in female but extending past CuA, often to CuA,.

Etymology. Petra and -ina (Latin), small rock,

upon which females will sometimes oviposit; also

the name of my wife.

Variation. Female, flagellum 39-43 segments

(mean 41, n=20), length 6.8-8.6 mm (mean 8.3

mm, n=18), wing length 22-25 mm (mean 22.7,

n=18).

Distribution. Western Australia, Ogyris subter-

restris petrina is known only from a few square

kilometres to the north east of Lake Douglas, near

Kalgoorlie

Biology

The life history of O. subterrestris is largely

unknown, It is associated with the sugar ant

Camponotus terebrans (Lowne), the same species

A NEW SPECIES OF BUTTERFLY OF THE GENUS OGYRIS 257

that is associated with all populations of O. idmo

and O. otanes (C. and R, Felder) (McArthur et al.,

1997), O. subterrestris has been recorded every

month from September to May with peak flight

activity in mid-spring and late summer and is

probably bivoltine at all locations. Near Waikerie

and at Pink Lakes males fly low in open grassland

and on nearby ridges whereas females generally

fly close to the trees where the ant nests occur. At

Kalgoorlie, the butterflies are sparse but have def-

inite flight paths and often exhibit hill-topping

and individuals, particularly males, fly to the tops

of small rises and settle on the ground, Females

are also often found on these rises.

The eggs are laid close to the ground at or in

the entrance to the ants’ nest usually on the bark

of trees or occasionally on small stones. At

most locations the trees are mallee eucalypts,

Eucalyptus concinna Maiden & Blakely near

Kalgoorlie (Field, 1992), E. oleosa F. Muell.,

E. foecunda Schauer, and E. pileata Blakely but

also exotic garden eucalypts and Mvoporum

platycarpum R. Br. near Waikerie, South Aus-

tralia (Moore, 1999) and E. largiflorens F. Muell.

at Mildura, Victoria. At the type location only one

egg has been found (on a mallee eucalypt)

although ant nests are abundant, occurring at the

base of many plants, not only eucalypts.

At Lake Douglas in 1991, ant nests at the base

of 50 randomly chosen E. concinna trees were

examined. Egg clusters of O. subterrestris were

found on 18 trees with 26 clusters present, four of

which were unhatched. Egg shells seem to remain

attached to the bark for several years. The cluster

size averaged 7.8 eggs (range 2-20) with 11% of

the eggs parasitised. In 1989 an encyrtid wasp

(Ooenevrtus sp.) was reared from one cluster of

eggs (Field, 1990). Captive females readily

oviposit on stones and bark taken from the

entrance of ant nests, producing clusters of 40 or

more eggs if left undisturbed. Nearly 90% of the

eggs laid in the field were in the northern to west-

ern quarter of the tree. However, the entrances to

the ant nest seemed to be well distributed around

the base of the tree and averaged 7.2 holes/tree

(range 1—18). Female butterflies that were caged

over ant nests readily oviposited, but only

between 1145 h and 1300 h, when the sun was

shining on the northwestern sector of the base of

the tree. During this period the ants, which are

predominantly nocturnal, exhibit little activity

above ground. If disturbed during the day the ants

will leave the nest and attack intruders. It is likely

that the butterflies oviposit during periods when

the sun is shining on the base of the tree and the

ants have retreated further underground. This

enables the females to back into the nest entrances

to oviposit undisturbed. Near Waikerie, hundreds

of old eggs occur on some trees with few eggs

showing evidence of parasitism.

Newly hatched larvae are carried in the

mandibles of ants or walk into the ant nest where

larval growth and pupation occurs. The first instar

larvae of O. subterrestris (Fig. 23) has morpho-

logical differences from O. idmo idmo (Fig. 24).

O. subterrestris has two pairs of dorsal posteri-

orly curving abdominal spines, a pair on segment

6 and on segment 7, A smaller pair of dorsal

spines also occurs on the mesothorax, These

spines do not occur on second instar larvae. There

are no spines on first instar O. idmo, but numer-

ous long hairs occur on the lateral margin of the

abdomen and thorax and there are numerous

clubbed secondary setae on the thorax and

abdomen, As with O. idmo, the larval food is

unknown. The larvae and pupae are thought to be

totally subterranean and probably have an obliga-

tory association with C. ferebrans. Adult pinned

specimens of O. idmo and O, subterrestris often

develop greasy wings, a feature commonly occur-

ring in lycaenids that are known to have predatory

larvae (Sands, 1980).

Discussion

A female specimen of O. subterrestris subter-

restris in the AM and labelled Lake Waltah, a

nonexistent location, and bearing the name F. R.

Spry, is most likely incorrectly labelled but may

refer to Lake Hattah, a location between the two

known Victorian locations of the species. Spry’s

diaries (held in the NMY) do not indicate that he

travelled in northwestern Victoria during

November 1918, hence he probably labelled this

specimen with misinterpreted data from the

collector.

Braby et al. (1997) proposed a common name

for this species of Mallee Bronze Azure. How-

ever, there are records of O. subterrestris subter-

restris from mallee and non-mallee areas (Broken

Hill, NSW, Mildura, Vic and near Ceduna, SA)

and O. idmo, both the western and eastern popu-

lations, also occurs in mallee vegetation as well as

heathland. A more appropriate common name

would therefore be Arid Bronze Azure, reflecting

the more arid climatic regions in which the

species is found in comparison to O. idmo (Field,

1997).

Most colonies of O. subterrestris occur in dis-

turbed areas. The colony at Lake Douglas occurs

within a public recreation area that can be sub-

jected to significant human interference with

numerous vehicle tracks crossing the main breed-

ing areas. In some years, since its first discovery

258 R.P. FIELD

Figures 21, 22. Genitalia of Ogyris subterrestris subterrestris sp.

nov.: 21, female; 22, male.

Figures 23, 24. First instar larvae of Ogyris: 23, O. subterrestris sp. nov; 24, O. idmo idmo. Scale line 1

mm.

A NEW SPECIES OF BUTTERFLY OF TI IE GENUS OGYRIS 259

in the early 1980s, the butterflies have been com-

mon, but few specimens have been seen since

1991 (Field, 1997). Near Waikerie, citrus

orchards and vineyards surround the breeding

area. These crops are regularly sprayed with pes-

ticide during flight periods of the butterfly, Adults

fly commonly in the adjoining farmland as well as

in the breeding area and a nearby ridge. The only

known extant location of O. subterrestris in

Victoria is the type location in the Murray Sunset

National Park. This relatively undisturbed loca-

tion is the most secure of all populations of the

species (Field, 1997),

The Ogyris idmo complex covers colour and

pattern variants, morphological and/or behav-

iouraly distinct populations of O. idmo and

O. subterrestris. Nowhere are specimens

common. Both O. idmo idmo and O. idmo halma-

turia are univoltine with peak flight activity in

mid to late November. The Mt Ragged form of

idmo has a longer flight activity period, flying in

early/mid October with specimen activity still

occurring in late December. Specimens from near

Geraldton more closely resemble the Mt Ragged

form of O. idmo than typical idmo and fly in

September, Flight activity during the day also dif-

fers amongst the complex. Female idmo tend to

be active only in the late morning, whereas males

are mainly active in the afternoon. However, at

Mt Ragged, males and females are active

throughout the day. Hunt et al. (1998) reported

male halmaturia having a peak flight activity in

the late morning. Both sexes of O, subterrestris

are active throughout the day although during the

heat of mid afternoon, specimens may shelter in

trees. The complex may yet reveal more species

than O. idmo and O. subterrestris. Larval charac-

ters have proven useful in separating species in

the complex. Further studies on the behaviour and

immature stages of the various idmo populations

may clarify the colour and morphological differ-

ences that occur in the adult populations.

Acknowledgements

I thank Drs Don Sands (CSIRO) and Ken Walker

(Museum Victoria) for comments and advice on

early drafts of the manuscript, Mr Ted Edwards

(CSIRO) for comments and access to images of

type specimens, Mr Phillip Ackery (Natural His-

tory Museum, London) for specimen loans and

label data on specimens in the Natural History

Museum, Mr Bob Hay for field assistance in

Western Australia and Ms Kate Nolan for the line

drawings of the larvae. Research on the Ogvris

idmo complex was partially funded by a research

grant from the Australian Entomological Society

in 1991,

References

Bethune-Baker, F\L.S., 1905. A monograph of the

genus Ogyris. Transactions of the Entomological

Society of London 3: 269-292.

Braby, M.F., Atkins, A.F., Dunn, K.L., Woodger, T.A.

and Quick, W.N.B., 1997. A provisional list of

common names for Australian butterflies. Aus-

tralian Journal of Entomology 36: 197-212.

Common, I. F. B. and Waterhouse, D. F., 1981. Butter-

Eliot, J. N., 1973. The higher classification of the

Lycaenidae (Lepidoptera): a tentative arrange-

ment, Bulletin of the British Museum (Natural His-

tory), Entomology 28: 373 505.

Field, R.P., 1990. Range extensions and the biology of

some Western Australia butterflies. Victorian

Entomologist 20(4): 76-82.

Field, R.P., 1992. Research grant report. Myrmecia

26(4): 12-17.

Field, R.P., 1997. The Ogyris idmo Hewitson (Lepi-

doptera: Lycaenidae) complex as flagship species

for conservation in southern Australia. Memoirs of

the Museum of Victoria 56(2): 389-392,

Hewitson, W.C., 1862, Specimen of a catologue of

Lycaenidae in the British Museum. British

Museum (Natural History): London. iii+15 pp. 8

pls.

Hunt, L., Moore, M. and Moore, D., 1998. Rediscovery

of Ogyris idmo halmaturia (Tepper, 1890) Victo-

rian Entomologist 28(6): 113-116.

McArthur Al, Adams, M. and Shattuck, S.O., 1997, A

morphological and molecular review of Campono-

tus terebrans (Lowne) (Hymenoptera: Formici-

dae). Australian Journal of Zoology 45(6):

579-598,

McCubbin, C.W., 1971, Australian butterflies. Nelson:

Melbourne. xxxiit+206 pp.

Moore, M., 1999. Some field notes on the as yet

unnamed Ogyris species (formerly included in the

species Ogyris idmo halmaturia) from Waikerie.

Victorian Entomologist 29(1); 12-18.

Sands, D.P.A., 1980. A new genus, Acrodipsas, for a

group of Lycaenidae (Lepidoptera) previously

referred to Pseudodipsas C. & R. Felder, with

descriptions of two new species from northern

Queensland. Journal of the Australian Entomolog-

ical Society 18: 251-265.

Tepper, J.G.0., 1890. Common native insects of South

Australia, A popular guide to South Australian

entomology. Part IT Lepidoptera, or butterflies and

moths, Adelaide. iv+65 pp.

260 R.P. FIELD

Waterhouse, G.A., 1903a. Catalogue of the Rhopalo-

cera of Australia. Memoirs of the New South Wales

Naturalists’ Club 1: 1-49.

Waterhouse, G.A., 1903b. Notes on Australian

Rhopalocera: Lycaenidae. Part IH. — Revisional.

Proceedings of the Linnean Society of New South

Wales 1903: 132 275.

Waterhouse, G.A. and Lyell, G., 1914. The Butterflies

of Australia. A monograph of the Australian

Rhopalocera. Angus and Robertson: Sydney.

vit+239 pp.

Memoirs of Museum Victoria 57(2): 261 (1999)

REPLACEMENT NAMES FOR FIVE SPECIES OF AUSTRALIAN LASIOGLOSSUM

(CHILALICTUS) (HYMENOPTERA: HALICTIDAE).

K. L. WALKER

Museum Victoria, PO Box 666E, Melbourne, Vic. 3001, Australia

(kwalker@mov.vic.gov.au)

Abstract

Walker, K.L., 1999. Replacement names for five species of Australian Lasioglossum

(Chilalictus) (Hymenoptera: Halictidae). Memoirs of Museum Victoria 57: 261,

Replacement names for one primary homonym and four secondary homonyms. within

Lasioglossum (Chilalictus) are proposed. The primary homonym change is: L. clariventre

(Friese, 1924) = L. impunctatum nom. nov. The secondary homonym changes are: L. bidens

Walker, 1995 = L. bidentatulum nom. nov., 1

… megacephalum Walker, 1995 = L, platychilum

nom. nov., L. occidens Walker, 1995 = L. occiduum nom. nov., L. soror Walker, 1995 -

L. sororculum nom. nov., and L. striatum Walker, 1995 = L. seriatum nom. nov.

Walker (1995) erected 73 new species of

Lasioglossum Curtis within the endemic

Australian subgenus Chilalictus Michener.

Unfortunately, four of these names created sec-

ondary homonyms and one existing primary

homonym was not identified and corrected. |

propose new names for these homonyms.

Halictus clariventris Friese, 1924 is a

primary homonym of Halictus clariventris Dalla

Torres, 1896 and is replaced with Lasioglossum

impunctatum nom. nov.

Lasioglossum bidens Walker, 1995 is a

secondary homonym of Halictus bidens

Cameron, 1904 and is replaced with Lasio-

glossum bidentatulum nom. nov.

Lasioglossum megacephalum Walker, 1995 is

a secondary homonym of Halictus megacephalus

Schenck, 1869 and is replaced with Lasioglossum

platychilum nom. nov.

Lasioglossum occidens Walker, 1995 is a sec-

ondary homonym of Halictus occidens Smith,

1873 and is replaced with Lasioglossum

occiduum nom. nov.

Lasioglossum soror Walker, 1995 is a sec-

ondary homonym of Halictus soror Saunders,

190land is replaced with Lasioglossum soror-

culum nom. nov.

Lasioglossum striatum Walker, 1995 is a sec-

ondary homonym of Halictus striatus Schenk,

261

1868 and is replaced with Lasioglossum seriatum

nom. nov.

I sincerely thank Pastor Andreas Werner

Ebmer for alerting me to these homonyms.

References

Cameron, P., 1905. On the Hymenoptera of the Albany

Museum, Grahamstown, South Africa. Records of

the Albany Museum 1: 185-265.

Dalla Torres, C.G. de, 1896. Catalogus hymenopterum

Vol. 10. Leipzig.

Friese, H., 1924. Ueber die Bienen

Konowia 3: 216-249,

Saunders, E., 1901. Balearic insects-Hymenoptera

Aculeata collected in Majorca and Minorca

(March and April, 1900) by E.B. Poulton, Oldfield

Thomas, and R.I. Pocock, with description of new

species. Entomologists’ Monthly Magazine 37:

208-211.

Schenck, A., 1869. Beschreibung der nassauischen

Bienen, Zweiter Nacktrag. Jahrbucher des

Nassauischen Vereins fur Naturkunde 21-22:

1-114. [Preprint, dated 1868, published 1869]

Smith, F., 1873. Descriptions of aculeate Hymenopter

a of Japan collected by Mr George Lewis

at Nagasaki and Hiogo. Transactions of the

Entomological Society of London 11: 181-206.

Walker, K.L., 1995. Revision of the Australian native

bee subgenus Lasioglossum (Chilalictus)

(Hymenoptera: Halictidae) Memoirs of the

Museum of Victoria 55(1-2); 1-423,

Australiens.

Memoirs of Museum Victoria 57(2): 263-266 (1999)

DESIGNATION OF TYPE SPECIES FOR THE GENERA OF AUSTRALIAN PAROPSINE

BEETLES (COLEOPTERA: CHRYSOMELIDAE)

PETER G KELLY

16 Roberts Street, East Brunswick, Victoria 3057, Australia

(pgkelly@melbpc.org.au)

C.A.M. REID

CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia

(Chris.Reid@ento.csiro.au)

Abstract

Kelly, P.G. and Reid, C.A.M., 1999. Designation of type species for the genera of Australian

paropsine beetles (Coleoptera: Chrysomelidae). Memoirs of Museum Victoria 57: 263-266.

Type species are listed for all Australian genera in the subtribe Paropsina, including neces-

sary first designations for Chrvsophtharta Weise, Faex Weise, Notoclea Marsham, Pyrgoides

gen. nov., Sterromela Weise, Trachymela Weise and Trochalodes Weise. The synonymy of

Notoclea with Paropsis Olivier, and Pyrgoides with Acacicola Lea are confirmed. A new

species synonym is recorded: Paropsis hamadrvas Stal = Acacicola tristis Lea.

Introduction

The paropsine chrysomelids are among the most

diverse and taxonomically intricate group of

beetles associated with natural vegetation in

Australian. A few paropsines have in recent years

become serious pests in eucalypt plantations in

Australia, New Zealand and South Africa. Many

of the 450 described species have been allocated

uncritically to genus and the genera themselves

have not always been delineated clearly. This

paper is a start towards resolving these taxonomic

problems.

The genus Puropsis was erected by Olivier

(1807) for 14 Australian chrysomelid species.

Marsham (1808) erected the genus Notoclea with

20 species which were incorporated into Paropsis

by Olivier (1825). Subsequently, Boisduval

(1835), Erichson (1842), Germar (1848), Bohe-

man (1858), Stal (1860), Clark (1864, 1866),

Baly (1864), Chapuis (1877) and Blackburn

(1890-1901) added large numbers of species to

the genus, so that by 1916, 406 species were

listed from Australia (Weise, 1916). Since 1916,

a number of species have been described by

Weise (1817, 1923), Lea (1924), Gressitt (1963)

and Selman (1983), and the current total is about

450 species.

This large number of rather similar species,

many with quite inadequate descriptions, has

263

made identification by later workers very

difficult. Several attempts have been made to

split Paropsis into smaller groups. Motschulsky

(1960) split three genera from Paropsis, namely

Dicranosterna, Niliosoma, and Paropsisterna.

Blackburn, in a series of papers (1897a-1901)

arranged Paropsis into five “groups” and a num-

ber of “subgroups” while still retaining the genus

Paropsis for all species. Weise (1901-1915) split

Paropsis into nine genera: Sterromela Weise,

Trochalodes Weise, and Dicranosterna Mot-

schulsky in the tribe Dicranosternini Weise and

Paropsis Olivier, Paropsisterna Motschulsky,

Chrysophtharta Weise, Trachymela Weise, Faex

Weise, Procris Weise, Philhydronopa Weise and

Pyrgo Weise in the tribe Paropsinini Weise. Most

of these genera are poorly diagnosed with no

accompanying illustrations. Some are defined

only by the species that Weise placed in them and

most lack designated type species.

Daccordi (1994) presented a checklist of world

chrysomeline genera and made several new syn-

onymic designations in the paropsine genera, thus

at present 15 Australian genera are considered

valid.

For this paper type specimens were examined

for most designated type species and where none

was available specimens named by Blackburn or

Lea were used. The validity of these genera is not

discussed here.

204 PETER G. KELLY AND C.A.M. REID

Australian generic names in the Paropsina

(sensu Daccordi, 1994), with designated type

species

The name Coccinelloides Latreille is not listed

below. Selman (1963) following Weise (1916)

claimed that this is an available generic-rank

name based on a species, “australasiae”, without

authorship. He then decided that Chrysomela

australasiae Fabricius was the intended spec-

ies and declared this to be the type species.

We believe that the phrase “coccinelloides

australasiae” of Latreille (1807:58) was never

meant to be generic but was merely descriptive

and this is clearly how Latreille uses the term

‘coecinelloides’ in a later work (1825: 10).

Acacicola Lea, 1902: 392

Type species. Acacicola tristis Lea, 1903 by

monotypy.

Chondromela Weise, 1915: 435

Type species. Chondromela mjoebergi Weise,

1915 by monotypy.

Chrysophtharta Weise, 1901: 165

lype species. Weise (1901) did not nominate a

type species but listed six species. The first of

these, Paropsis nobilitata Erichson, 1842, is here

designated type species.

Dicranosterna Motschulsky, 1860: 193

Type species. Paropsis picea Olivier, 1807 by

original designation.

Faex Weise 1901: 165

Type species. Weise (1901) did not designate a

type species but listed three species names. The

first, Paropsis notatipennis Chapuis, 1877 is here

designated type species.

Niliosoma Motschulsky, 1860: 194

Type species. Paropsis testacea Olivier, 1807 by

original designation. This species is unrecognis-

able and the type is apparently lost. Niliosoma is

therefore a nomen dubium and unusable, but it

any case tt has always been ignored by Australian

workers.

Notoclea Marsham, | 808: 283

Type species. Marsham described 20 species in

this genus but failed to designate a type. Notoclea

obsoleta Marsham, 1808 is hereby designated

type species. Notoelea obsoleta is a junior

synonym and homonym of Paropsis obsoleta

Olivier, 1808 therefore synonymy of the two

genera is confirmed (Daccordi, 1994).

Novacastria Selman, 1983a: 179

Type species. Novacastria nothofagi Selman,

1983 by original designation and monotypy.

Paropsides Motschulsky, 1860: 192

Type species. Paropsis — duodecimpustulata

Gebler, 1825 by original designation.

Paropsis Olivier, 1807: 596

Type species. Paropsis obsoleta Olivier, 1807 by

subsequent designation (Selman, 1963).

Paropsisterna Motschulsky, 1860: 192

Type species. Notoclea sexpustulata Marsham,

1808 by original designation.

Philhydronopa Weise, 1901: 166

Type species. Paropsis aeneipennis Chapuis,

1877 by original designation.

Procrisina Aslam, 1968: 129

Procris Weise, 1901: 165, nec Fabricius, 1807

(junior homonym).

Clorindina Jolivet, 1957: 170,

lerich, 1930 (junior homonym).

Clorindiella Jolivet, 1971: 382 (unecessary

nom. nov).

nee Khoda-

Type species. Paropsis pictipennis Boheman,

1858 by original designation (Weise, 1901).

Pyrgoides gen. nov.

Pvrgo Weise 1901: 166, nee Defrance, 1824

(junior homonym).

Pyrgoides Aslam, 1968: 129 (nomen nudum,

no type species)

Type species. The name Pvrogoides was proposed

by Aslam (1968) as a replacement name for the

preoccupied Pyrgo Weise. Published after 1930

Aslam’s name is unavailable as no type species

was designated (ICZN Article 13(b)). We make it

available here. Weise (1901) did not designate a

type species of Pyrgo but listed a number of

species to be included in his genus. Paropsis

hamadryas Stal, 1860, one of these, is hereby des-

ignated type species. This species is a senior syn-

onym of Acacicola tristis Lea (syn. nov.; types

examined), thereby confirming synonymy of

Pyrgoides with Acacicola Lea (Daccordi, 1994).

TYPE SPECIES OF GENERA OF AUSTRALIAN PAROPSINE BERTI ES 265

Rhaebosterna Weise, 1917: 124

Type species. Rhaebosterna sciola Weise, 1917

by monotypy.

Sterromela Weise, 1915: 436

Type species. Weise (1915) placed four species of

Paropsis in Sterromela without designating a type

species. The first named of these, Paropsis sub-

costata Chapuis, 1877 is here designated type

species.

Trochalodes Weise, 1901: 167

Type species. No type species was designated

by Weise (1901) but three species were named,

two erroneously. The first and only correctly

named species, Paropsis circe Stil, 1860 is here

designated the type species.

Trachymela Weise, 1908: 8

Type species. No type species was deisgnated by

Weise but he listed three species names and, by

inference, all species in Blackburn's Group HI of

Paropsis. One of the listed names, Puropsis

sloanei Blackburn 1897, is here designated type

species.

Xanthogramma Weise, 1923: 63

Type species. Xanthogramma pellucida Weise,

1923 by monotypy.

Acknowledgements

We are grateful to Dr T. New (LaTrobe Uni-

versity, Melbourne), Dr K. Walker (Museum

Victoria, Melbourne) and Dr E. Matthews (South

Australian Museum, Adelaide) for access to

collections and helpful discussion. This work

was completed with assistance to PGK from the

Setence and Industry Endowment Board.

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Memoirs of Museum Victoria 57(2): 267-285 (1999)

STORTHYNGURELLA, NEW GENUS OF MUNNOPSIDAE (CRUSTACEA: ISOPODA),

WITH DESCRIPTIONS OF THREE NEW SPECIES FROM DEEP-SEA BASINS

OF THE SOUTHERN HEMISPHERE

M. V. MALYUTINA

Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences, 17 Palchevsky St,

Vladivostok 690041, Russia

(inmarbio@mali.primorye.ru)

Abstract

Malyutina, M.V., 1999. Storthyngurella, new genus of Munnopsidae (Crustacea: Isopoda),

with descriptions of three new species from deep-sea basins of the Southern Hemisphere.

Memoirs of Museum Victoria 57: 267-285.

A new genus Storthyngurella and three new species are described from deep-sea basins and

trenches of the Southern Hemisphere. Storthyngurella is characterised by: long dorsal and lat-

eral spines on the body; a terminal spine on the pleotelson; a long spine on the basal article of

both antennae; narrow posterior pereopods 5-7; elongated distal lobes of male pleopod 1

extending far beyond the distal margins of pleopods 2; a long stylet of male pleopod 2, which

is more than two-thirds as long as the protopod; and elongated uropods with tubular protopod

and rami. The new genus is most similar to Storthyngura and Microprotus.

Introduction

A specialised deep-sea family Munnopsidae Sars,

1869 is characterised by natatory posterior legs

with paddle-like carpi and propodi bearing

plumose setae, fused compact natasome and

streamline body outline. A small group of mun-

nopsid genera is distingushed by spinose body,

fused but non-compact natasome, and relatively

narrow pereopods 5—7, which in some cases are

lacking natatory setae. This group comprises

Acanthocope Beddard, 1885, Storthyngura

Vanhöffen, 1914, and Microprotus Richardson,

1910. The most numerous (more than 40 species)

and widespread genus Storthyngura shows vari-

able external morphology. Its species are distin-

guished by different number and length of body

spines, shape of pleotelson, and morphology of

appendages. Among the representatives of the

genus large active swimmers, which swim back-

wards, are characterised by relatively compact

muscular body, paddle-like pereopods 5-7, and

short body processes directed forward; also a few

elegant, brittle species have thin, long spines and

narrow, often walking posterior legs. George and

Menzies (1968a) tried to divide the genus into

five groups and 14 subgroups. Although they

wrote that their analysis was based on 158

267

characters, description of these subdivisions was

based mainly on pleotelson morphology. Never-

theless, the groups were distinguished by other

morphological characters such as armament of

the antennae, morphology of the posterior pairs

of pereopods and male pleopods. The artificiality

of such a classification was noted by subsequent

authors (Birstein, 1969; Wilson et al., 1989).

Working with Storthvngura specimens collected

by RV Akademik Kurchatov (cruises 11 and 43)

and RV Dmitrii Mendeleev (cruises 16 and 43)

from deep-sea trenches and basins of the South-

ern Hemisphere, I was able to compare 12 species

of this genus. The new genus Storthyngurella

is erected for three new species described in

this paper and five previously known species of

Storthyngura.

I am indebted to colleagues from the Labora-

tory of Benthos in Shirshov’s Institute of Ocean-

ology, Moscow, for the privilege of being able to

examine the Storthyngura collections. The ter-

minology and measurements follow Wilson and

Hessler (1980). When measuring the body pro-

portions the body spines were omitted. The

type material is deposited in the Zoological

Museum of Moscow University, Moscow, Russia

(ZMMU) and Museum Victoria, Melbourne,

Australia (NMV).

268 M. V. MALYUTINA

Suborder Asellota

Munnopsidae Sars, 1869

Storthyngurella, gen. nov.

Type species. Storthyngurella hirsuta sp. nov.

Diagnosis. Body slender, with long spines: dorsal

on cephalon, pereonites and pleotelson, lateral on

coxae of pereopods 1-4, pereonites 4-7 and pleo-

telson, and terminal on pleotelson. Posterior body

part (natasome) longer than anterior part, non-

compact, with distinct waist (narrowed anterior

part of pereonite 5), pleotelson longer than broad,

with long preanal ventral process. Antenna | sex-

ually dimorphic: in males 1.5-2 times as long as

in females, basal article of antenna 1 with long

medial spine, basal article of antenna 2 with long

lateral spine. Mandible normal, with cuticular

posterolateral projection articulated with

cephalon in elongate notch, proximal portion of

mandibular body up to condyle and palp abbrevi-

ated, Pereopods 5-7 thin, either with tubular

carpus and propodus lacking natatory setae or

with flattened articles provided with short

plumose setae, either on dorsal margin or in both.

Dactyli of pereopods 2-7 with acute dorsal claw

provided with inner acute projection. Male pleo-

pod | clongated without distinct waist midlength,

with long distal lobes making angle to each other

and extending in situ far beyond distal margins of

pleopods 2. Male pleopod 2 endopod inserted in

midlength of protopod medial margins, stylet

length more than two-thirds of protopod length,

exopod with long distal hook. Pleopod 3 endopod

with 3-4, exopod with numerous plumose setae.

Uropod elongate, with tubular protopod and rami

subequal in length.

Additional description. Body widest at pereonite

4, long dorsal and lateral spines directed perpen-

dicularly to body axis and (or) backward (only in

S. digitata body spines are short and of unusual

bulbous shape). Cephalon without rostrum,

broader than long, with pair (2 pairs in S. digitata)

of long dorsal spines, laterally projecting cheeks

at mandibular articulation point; frons sloping

with frontal arch semicircular in frontal veiw,

clypeus significantly broader than labrum, Pere-

onites 1—4 slightly broadening and lengthening

from 1 to 4, pereonites 2-4 with 3 dorsal spines

(in S. wolffi sp.nov. and S. benti only with |

spine), anterolateral margins of pereonites 1-3

rounded, of pereonite 4 with long spines, coxae of

pereopod | with 1, of pereopods 2-4 with 2 long

11 with 1, of yds 2-4 with 2 lon

spines. Pereonites 5-7 and pleotelson fused,

dorsal sutures visible, ventral sutures absent, each

pereonite bearing pair of dorsal spines located

relatively far from each other and anterolateral

long spines with narrow basis. Pleotelson with

several (1-6) dorsal and 3 lateral spines (2 spines

and a process between them in S. wolffi sp. nov.

and S. digitata) directed backward. Basal

article of antenna | same (equal) in both sexes,

elongated, with subparallel margins, flattened lat-

eral lobe and medial spine, exeeding in length

article width, article 2 length 0.3 article | length

and subequal in length to article 5, article 3

slightly longer than article 2, article 4 shortest

among first five, in males articles 2-5 broader and

stronger than in females, flagellar articles in male

short, numerous, in female flagellar articles less

numerous, longer, subequal in length to article 4.

Basal article of antenna 2 with long lateral spine,

article 2 without spines, article 3 with 2-3 spines.

Mandibular incisor process broad, flattened and

more or less curved, condyle shorter than molar

process, which narrowed distally, with row of

small denticles and setae on tip, palp normal,

long, thin, last article twisted. Maxilla 1 inner

lobe with numerous small, thin distal setae, disto-

medial seta largest. Maxilla 2 lobes subequal in

length, middle and outer lobes with 4 long spine-

like distal setae, inner lobe bears thin simple

marginal setae, distal setae more strong, some

pectinated. Maxillipedal palp inserting in mid-

length of basis, palp article 3 with semi-

circular medial margin, epipod elongated, lateral

margin smoth, lacking acuted rolling process.

Pereopod | much smaller than posterior pere-

opods, having only simple setae, carpus slightly

curved, dactylar dorsal claw finger-like, rounded

distally. Pereopods 2-4 progressively lengthen-

ing; straight carpi and propodi, bearing strong,

unequal bifid ventral setae and sparse

simple thin dorsal setae. All bases subequal in

length. Male pleopods 2 not connected to each

other, fully separated by insertion of pleopods 1;

pleopod 2 protopod semicircular, lateral and dis-

tal margins with plumose setae. Female pleopod 2

(operculum) oval, with plumose marginal setae,

median keel low, with or without spine.

Species included. All species except the new ones

were transfered from the genus Storthyngura

(Table 1).

Geographic distribution. Species of the genus

Storthyngurella have been found mainly in the

Southern Hemisphere: from Middle American

Trench, 12°N to 61°S in the Atlantic and in the

southeastern Indian Ocean in the depths of 2596

to 7200 m.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 269

Table 1. List of species of Storthyngurella gen. nov.

Species Locality: (A), Atlantic, (I), Indian, (P), Pacific Oceans Depths (m)

S. hirsuta sp. nov. (A) South Sandwich Trench: 56°07’-56°52’S, 24°56'24°59' W 5530-6150

S. wolffi sp. nov. (D South Australian Basin: 43°36.5’S, 144°04'E 4464

(P) Solander Depression: 54°57.5’S, 163°51'E 4590

(P) Hjort Trench: 57°17’S, 157°13'E 5460-5760

S. menziesi sp. nov. (A) Argentine Basin: 38°40'S, 48°08'W; 48°52'S, 26°36’ W 5225

4560-4570

S. benti (Wolff, 1956) (P) Kermadec Trench: 32°10'-36°07'S, 175°54°-178°32'W 5340-7000

S. spinosa (1) South Indian Basin: 54°S, 109°E 3567

(Beddard, 1885)

S. triplispinosa (P) Panama Basin: 16°12’S, 74°41°W 2856-2596

(George and (P) Middle American Trench: 12°11°N, 89°34"W 5680-5690

Menzies, 1962) (A) 45°34'S, 06°02" W 4588

S. digitata (A) Argentine Basin: 44°53.3’S, 51°26.5°W 5843

(Menzies, 1962) 36°12'-38°40'S, 48°08 —49°10° W 4630-5255

S. zenkevitchi (A) Romanche Gap: 01°01'S, 18°21.55'E 7200

(Birstein, 1969)

Etymology. Representatives of the new genus are

similar to but smaller and more elegant than

species of Storthyngura. The diminutive suffix

-ella is added to Storthyngura.

Remarks. Storthyngurella gen. nov. has clear

affinities with the genera Storthvngura and

Microprotus. From Storthyngura, the new genus

differs in smaller body size, longer body spines

and their arrangement, terminal spine of pleotel-

son, long spines on the first articles of both anten-

nae, narrow pereopods 5-7, long distal lobes of

male pleopods 1 which make an angle to each

other and extent far beyond the distal margins of

pleopods 2, and longer stylet of pleopods 2,

inserting at the midlength of the protopod,

Until now the presence of tubular pereopods

5-7 in Munnopsidae was described only for

Microprotus. At least two species of Storthyn-

gurella, S. menziesi and S. triplispinosa, have

such legs, while other Storthyngurella species

have carpi and propodi of pereopods 5-7 signifi-

cantly narrower than in all munnopsids except

Microprotus. Plumose setae in Storthyngurella

species show different stages of reduction: either

short plumose setae located on both margins of

narrow carpi and propodi, or carpi and propodi

bearing small plumose setae on dorsal margins

and unequal bifid setae on ventral margins, or the

plumose setae totally reduced. Wilson et al.

(1989) discussed the hypothesis that there is sec-

ondary reversion of natatory legs into walking

ones within this specialised swimming family. An

intermediate stage of such a transformation is

seen in Storthyngurella. Besides the narrow legs,

long spines of the body, pair long spines on the

coxae of pereopods 1-4, long spine on the article

1 and article 3 of antenna 2 and long tubular

uropods also make the new genus similar to

Microprotus.

In spite of external similarity between Stor-

thyngurella gen. nov. and Acanthocope (both

have a long terminal spine on pleotelson, body

spines, long uropods and narrow pereopods 5-7)

a more detailed analysis did not confirm their

close affinity. The monotypic subfamily Acantho-

copinae is well isolated from other spiny munnop-

sid genera, first by the different structure of

male genital apparatus (Malyutina, in press).

Therefore, Wägele's (1989) unification of

Acanthocope, Storthyngura and Microprotus into

Acanthocopinae is dubious.

Storthyngurella hirsuta sp. nov.

Figures 1-3

Material examined. Holotype. South-Sandwich Trench,

RV Akademik Kurchatov, cruise 11, stn 898 (56°47'S,

24°56'W), 6052-6150 m, ZMMU Me 1316 (male, 11.1

mm long).

Paratypes. Type locality, ZMMU Mc 1317a ( 8

males, 7.4-12.3 mm long; 6 females, 9.3-11,7 mm

long; 14 fragmented specimens), NMV J45733 (1 male,

| female). Stn 896 (56°52'S, 24°9’W), 5530-5651 m,

ZMMU Me 1317b (5 males, 12 females, 6.5-13.4 mm

long; fragments of 6 specimens).

270 M. V. MALYUTINA

Figure 1. Storthyngurella hirsuta sp. nov., male, holotype: a, b, dorsal and lateral views of body; c, d, frontal

and lateral views of cephalon; e, maxilla 1; f, maxilliped; g, antenna 1; h, maxilla 2; i, left mandible; j, right

mandible; k, peduncle of antenna 2.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 271

Figure 2. Storthyngurella hirsuta sp. nov.: a, b, female, paratype, dorsal and lateral views of body; male, holo-

type: c-g, pereopods 1-5; h, uropod.

272 M. V. MALYUTINA

Figure 3. Storthvngurella hirsuta sp. nov.: a, operculum of female, paratype; b-f, pleopods 1-5 of male,

holotype: g, antenna | of female, paratype; h, i, pereopods 6, 7 of male, holotype.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 273

Description of the holotype. Body (Figs la, b)

slender, length 3.4 times width, height 0.14 times

length. Cephalon length 0.56 times width, dis-

tance between antennae 1 is 0.1 cephalon width.

Frons (Figs Ic, d) sloping, concave medially,

frontal arch low, clypeus 1.9 times as long as

labrum, clypeus width 0.8 labrum width. Pere-

onite 1 with 1, pereonites 2-4 with 3 dorsal

medial long spines. Anterolateral spines of pere-

onite 4 perpendicular to body axis, half as long as

pereonite width. Coxae spines of pereopods 1—4

much shorter than lateral spines of pereonite 4.

Posterior body part 1.3 times as long as anterior

part. Pereonite 5 longer than pereonites 6 and 7,

the last 2 subequal in length, with long waist in

anterior part; pereonites gradually narrowing

from 5 to 7, their long anterolateral spines direct

backward and slightly upward distally.

Pleotelson subtriangular, its length 1.2 times

width, terminal spine length 0.7 pleotelson length,

spine turning up distally, its lateral sides with 3

spines: anterior and posterior long, slightly

curved upward, medial spine shorter, curved

downward, Dorsal surface with 2 long medial

spines and 2 short spinules on the sides and

slightly behind the posterior medial spine. Preanal

ventral process of pleotelson long, narrow,

slightly curved backward, with concave notch.

Lateral margins of pereonites, pleotelson, and

body spines covered with small acute tubercles.

All spines on body, coxae and antennae look like

bristle-scourer, with dense, long. hair-like bifid

setae (Fig. lk).

Antenna | (Fig. lg) length 0.3 times body

length, article 1 is 2.3 times as long as wide,

article medial length 1.3 article width, article 1

extends almost to article 2 distal end. Article 2

length 0.45 article 1 length, articles 3-S length

1.2, 0.3, and 0.9 article 2 length respectively,

flagellum of numerous short articles, some with

aesthetascs.

Antenna 2 (Fig. lk) broken, only basal part pre-

sent. Article 1 lateral spine equal in length to arti-

cle 1 width, article 2 twice as long as article 1,

article 3 almost twice as long as article 2, with

long lateral spine and 2 shorter medial spines.

Mandibles (Figs li, j). Incisor process with 4

cusps, lacinia mobilis of left mandible consider-

ably broadening distally, almost half as long as

incisor process, with 4 teeth, spine row with 11

and 12 members in left and right mandibles

respectively, palp thin, slightly longer than

mandibular body, article 2 is 3 times as long as

article 1, with few distal setae, article 3 narrow,

with numerous setules in inner surface and 5 stout

distal setae.

Maxilla | (Fig. le). Inner lobe half as long as

and narrower than outer lobe, the latter with 12

claw-like setae, some serrated.

Maxilliped (Fig. If). Basis with 7 coupling

hooks, endite with 6 fan-setae and numerous thin

setae distally, endite width 0.7 basis width, palp

article 2 lateral length 1.4 article width, and the

width 1.7 endite width, article 3 length 0.25

article 2 lateral length and 0.9 medial length,

article 4 medial lobe and article 5 with 5 simple

distal setae each. Epipod rounded distally, its

length 2.7 width.

All pereopods (Figs 2c—g, 3i, j) ambulatory,

thin, similar, but first thinner and disproportion-

ately smaller then others, their length gradually

increase from 2 to 4 and then decrease to 7. Pere-

opods 1—4 bases subequal in length, slightly

longer in pereopods 5-7. All bases with row of

small spines and plumose setae on dorsal margin

each. Ischia of pereopod 1 and 5-7 less than half

corresponding bases length (0.40, 0.42, 0.42,

0.47), in pereopods 2-4 more than half (0.54,

0.56, 0.60). Ratios of lengths of carpi 1-7 : bases:

0.80, 1.20, 1.60, 1.82, 0.94, 0.79, 0.84 respec-

tively; propodi : bases: 0.56, 1,05, 1.76, 1.96,

0.88. 0.82, 0.84; and dactyli : bases: 0,15, 0.66,

0.86, 0.96, 0.66, 0.64, 0.62. Carpi 5-7 slightly

flattened and broaded, in pereopod 7 less so.

Ventral margins of carpi and propodi 5-7 with

thin, unequally bifid setae, dorsal margins with

small plumose setae. Dactyli 2-7 with numerous

thin, simple dorsal setae.

Pleopod 1 (Fig. 3b) slightly tapering, its length

3.2 proximal width, midlength slightly narrowed,

ventral surface lacking setae, but with small

tubercules, distal lobes long, 30° to cach other at

distance 0.2 of protopod length from distal tip.

Inner lobes of distal margin subrectangular with

numerous thin long setae, outer lobes narrow,

produced, slightly curved inward.

Pleopod 2 (Fig. 3c) length twice width, with

dense row of plumose setae, endopod inserted at

distance 0.6 of protopod length from basal mar-

gin, stylet 0.8 protopod length, well tapering dis-

tally, extending not far beyond distal margin of

protopod, exopod extrinsic musculature occupy-

ing central part of protopod.

Pleopod 3 (Fig. 3d) length 0.8 pleopod 2

length, endopod subrectangular with 4 distal plu-

mose setae, length twice width, exopod narrow,

2-segmented, longer than endopod, with 16 distal

plumose setae and row of thin simple lateral setae.

Pleopod 4 (Fig. 3e). Endopod semicircular with

acute distomedial tip, exopod length 0.8 endopod

length and 0,5 endopod width, with 2 distal

plumose setae.

274

Uropod (Fig. 2h) 0.82 pleotelson length. Proto-

pod slightly broadening distally, with 1 long and

2 short distal setae, length 7.8 width. Endopod

about half width of protopod and 0.8 protopod

length, with 4 medial, 4 lateral strong unequal

bifid setae and 5 plumose distal setae.

Female paratype. In appearance and structural

details, the female is similar to the male. Antenna

| (Fig. 3g) shorter than in male, with articles 2-5

thinner , flagellum of about 10 clongated articles,

Pereonite 7 with ventromedial stout spine. Oper-

culum (Fig. 3a) oval, length 1.3 width. Keel

rounded ventrally, covered with small acute

tubercules, distal part elevated and folded, lateral

and distal margins with plumose setae.

Etymology. Hirsuta (Latin), hairy or shaggy,

referring to the long, dense hairs on the body

spines.

Remarks. The new species is most similar to

S. zenkevitchi, The two species share the same

number and arrangement of dorsal spines on pere-

onites, male pleopod 2 tapering distally, and 4

distal setae on the endopod of the pleopod 3 (all

other species have 3 setae). The new species is

distingished by longer body spines, longer ter-

minal spine on the pleotelson, different length of

the lateral spines of pleotelson and in having long,

dense hairs on the body spines.

Storthyngurella wolffi sp. nov.

Figures 4-6

Material examined. Holotype. RV Dmitrii Mendeleev,

cruise 16, stn 1349 (46°36.5'S, 144°04’E), 4464 m,

ZMMU Me 1318 (male, 9.1 mm long).

Paratypes. Type locality, ZMMU Me 1319a (4

females, 6.8-10.7 mm long), NMV J45732 (1 male, 1

female). Stn 1285 (54°57.5'S, 163°51.3/E), 4590-4560

m, ZMMU Me 1319b (1 male, 11 mm long) Stn 1299

(57°17'S, 157°13" E), 5760-5460 m, ZMMU Me 1319c

(1 male, 12.3 mm long).

Description of holotype. Body (Figs 4a, b) length

3.2 times width, height 0.2 times length.

Cephalon (Figs 4c, d) length 0.5 width, distance

between antennae | 0.12 cephalon width, frons

concave medially, frontal arch low, clypeus twice

as broad and half as long as labrum. Pereonites

1-4 with | dorsal medial long spine each. Antero-

lateral spines of pereonite 4 perpendicular to body

axis, equal in length to pereonite medial length.

Coxal spines of pereopods 1-4 almost twice as

long as anterolateral spines of pereonite 4. Nata-

some 1,3 times as long as anterior body part.

Pereonite 5 slightly longer than pereonites 6

and 7, the last 2 subequal in length, pereonites

M. V. MALYUTINA

gradually narrowing from 5 to 7, their long

anterolateral spines directed slightly forward,

Pereonites ventral side with small acute medial

spines.

Pleotelson pentagonal, separated by clearly

visible suture, length 1.1 width, terminal spine

almost straight, length 0.6 pleotelson length, lat-

eral sides bearing 2 long spines subequal in length

to terminal spine, and rectangular projection

between them directed slightly down. Dorsal

surface with 2 long medial spines and 2 short on

sides and slightly behind posterior medial spines,

Preanal ventral process of pleotelson long and

broad, turned backwards, visible in dorsal view,

with deep serrated medial notch and long lateral

spines almost reaching tip of terminal spine.

Lateral margins of pereonites with fine denticles,

margins of pleotelson and body spines with

prominent ones.

Antenna | (Fig. 6a) length 0.4 body length,

article | elongated, 2.3 times as long as wide,

article medial spine subequal in length to article,

lateral lobe subequal in length to article 2. Article

2 length 0.3 article 1 length, with distal denticles,

articles 3-5 length 1.6, 0.3, and 1.1 of article 2

length respectively, flagellum with many short

articles, some with aesthetascs.

Antenna 2 (Fig. 4a) almost twice as long as

body. Basal articles 1-3 gradually lengthening,

article 1 with long lateral spine almost twice as

long as article width, article 2 twice as long as

article 1, article 3 about 1.5 as long as article 2,

with 1 lateral and | medial spines, article 4 shorter

than article 3, length 0.6 width, article 5 twice as

long as all preceding articles, article 6 slightly

narrower and 1.2 times as long as article 5.

Flagellum with approximately 110 elongated

articles.

Mandibles (Figs 4e-g). Incisor process of left

mandible broad and very thin, so curved that in

frontal view looks narrow, with 4 cusps, lacinia

mobilis broadening distally, almost half as long as

incisor process, with 4 narrow dorsal and | broad

ventral tooth, spine row with 13 and 14 members

in left and in right mandibles respectively, molar

process with 6 denticulated distal setae, palp thin,

slightly shorter than mandibular body, first 2

articles equal in width, article 2 is 2.7 times as

long as article 1, with 4 distomedial setae, article

3 with 7 large distal setae.

Maxilla | (Fig. 4i). Inner lobe width 0.6 outer

lobe width, with 12 claw-like setae, some

serrated,

Maxilliped (Fig. 4h). Basis with 7 coupling

hooks, endite with 6-7 fan-setae and numerous

thin setae distally, endite width 0.7 basis width,

275

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS

mas

eso

Figure 4. Storthyngurella wolffì sp. nov., male, holotype: a, b, dorsal and lateral views of body; c, d, frontal

ght mandible; h, maxilliped:

and lateral views of cephalon; e, left mandible; f, distal part of left mandible; g, ri

i, maxilla 1; j, maxilla 2.

276 M. V. MALYUTINA

Figure 5. Storthyngurella wolffi sp. nov., male, holotype: a-g, pereopods l-7; h, antenna | of female,

paratype.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 277

Figure 6. Storthyngurella wolffi sp. nov., male, holotype: a, antenna 1, b-f, pleopods 1-5; g, uropod; female,

paratype: h, pleotelson, ventral view; i, operculum, lateral view.

27% M. V. MALYUTINA

lateral length of palp article 2 subequal to article

width, and its width 1.7 endite width, article 3

length 0.24 article 2 lateral length and 0.8 medial

length, article 4 medial lobe and article 5 with 7

simple distal setae each, Lpipod rounded distally,

S tines as long as wide, lateral margin with

smoothed midlength projection,

Percopods 1 4 (hips 5a d) ambulatory, thin,

similar in structure, first slightly smaller than

second, pereopods 3 and 4 much longer than all

other, percopods 5 7 (hips 5e p) subequal in size

and shape, All pereopod bases subequal in size, in

4 they are broadest articles. All

bases with small and sparse setae only, Ischia of

pereopods Land 5 7 about half of corresponding

bases length (O46, O48, 0.51, 0.42), and more

than hall in pereopods 2 4 (0.79, 0.66, 0.66)

Ratios of lengths of carpi 1 7: bases: 0,56, 1.34,

Lob, 1.95, 0.94, 0.90, 0.67 respectively; propodi

bases: 0.63, 1.57, 2.04, 2.42, 0.97, 0.95, 0 72: and

dactyly bases: 0.1, 0.6, 0.91, 0.79, OAL, 04, 0,32

Carpi 5 7 Hattened and slightly broadened, length

about 4 times width, propodi 5 7 length 7.1, 6.5,

Ot width respectively, both margins of carpi

and propodt 5 7 and dorsal margin of ischia 5 7

with thin short plumose setae, Dactylt 2 7 with

numerous thin, simple dorsal setae

Pleopod 1 (Fig. 6b) relatively broad, broaden

mge distally, length 2.7 proximal width, ventral

surface lacking setae, but with small high tuber

cules, distal lobes /0 to cach other at distance 0.2

of pleopod length trom distal tip. Inner lobes of

distal margin subrectanpular with numerous thin,

long setae, outer lobes narrow, projecting, slightly

curved inward

Pleopod 2 (Fig. Ge) length 1.6 width, endopod

inserted 0,6 protopod length trom basal margin,

stylet 0.9 protopod length, strongly tapering dis

tally, not extending beyond distal margin of pro

lopod, exopod extrinsic musculature occupying

bulk of protopod, coming close to lateral margin

Pleopods I and 2 much larger and thicker than

remainder

Pleopod 3 (big. Gd) length 0.6 pleopod 2

length, endopod subreetangular, length 1.4 width,

with 3 distal plumose setac, exopod narrow,

sliphtly longer than endopod, with 9 distal

plumose setae and row of thin simple lateral setac.

Pleopod 4 (big, 6e), Endopod semicircular with

acute distomedial tip, exopod length 0.8 endopod

length and 0.5 endopod width, with 1 distal seta

Uropod (Pip. 6g) 0.56 pleotelson length,

extending bevond terminal spine of pleotelson

Protopod shghtly broadening distally, with 4

medial setae, length 4.3 width. Endopod about

hall width of protopod and slightly shorter (0.97),

pereopods |

with 4 medial, 4 distal unequal bifid setae, and |

plumose lateral seta, exopod slightly narrower

than endopod, its length 0.8 endopod length, with

3 distal unequal bifid setae,

Female paratype. In appearance and structural

details, the female is similar to the male. Antenna

| (Fig, 5h) almost half as long as in male, article

| length 2.2 width, medial spine subequal in

length to article, articles 2 5 length ratio almost

same as in male, article 2 subequal in width to fol-

lowing articles, flagellum with & elongated arti-

cles. Operculum (Figs 6h, i) oval, length 1.3

width. Keel rounded ventrally, covered with small

acule tubercules with long, stout spine in

midlength, lateral and distal margins with

plumose setae

Etvmologv The species is named after the out-

standing Danish carcinologist, Prof. Torben

Wolff.

Remarks, The specimens varied little. Lateral

spines of the preanal process are broken in all

specimens except the holotype. The new species

is identified as Srorthynevurella by the combina-

tion of the following features: long spines on the

body, coxae | 4, and basal articles of both anten-

nae; long terminal spine and preanal process on

the pleotelson; narrow carpi and propodi of

pereopods 5 7; shape of male pleopods | and 2;

armament of pleopods 3; and long uropods. But

S. wolff is distinguished from the other species of

the genus by unusual shape of pleotelson and |

(not 3) dorsal spine on pereonites 2 4. The only

other species with I dorsal spine on perconites

2 41s. benti, but pleotelson shape in this species

is common for the genus, The presence of two lat-

eral spines on pleotelson makes S. wolffi similar

only to S digitata, but spines on the

pleotelson and pereonites of the latter species are

different,

Storthyngurella menziesi sp. nov.

Figures 7, 8

Material examined. Holotype. Argentine Basin, RV

Dmitrii Mendeleev, cruise 43, stn 4109 (38°40'S,

4R OR W), 5225 m, ZMMU Me 1320 (male, 10,2 mm

long)

Paratypes, Type locality, ZMMU Me 132 la (2 males,

92, and 94 mm; I female, 11.2 mm; fragments of 6

specimens), NMV J45734 (1 male, I female), RV

thademih Kurchatov, cruise LL, stn SSN (48°52'5,

26°34 W), 4560 4570 m, ZMMU Me 1321b (1 female,

14.3 mm long),

Description of the holotype. Body (Figs 7a, b)

length 3.2 times width, height 0,15 times length.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 279

Figure 7. Storthyngurella menziesi sp. nov. a, b, f-k, male, holotype; c-e, female, paratype. a, b, dorsal and

lateral views of body; c, d, pleotelson, ventral and lateral views; e, antenna 1; f, maxilliped; g, maxilla 1;

h, maxilla 2; i, left mandible; j, right mandible; k, uropod.

M. V. MALYUTINA

Figure 8. Storthyngurella menziesi sp. nov., male, holotype: a-c, pereopods 5-7; d, pereopod 1; e-i, pleopods

l-5; j, antenna 1.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 281

Cephalon length 0.5 width, distance between

antennae | is 0.15 cephalon width, antennal

sockets well marked, frons slightly concave

medially, clypeus 1.75 times as broad and half as

long as labrum. Pereonite 1-4 with 3 dorsal long

spines. Anterolateral spines of pereonite 4 per-

pendicular to body axis, equal in length to half of

pereonite width. Coxal spines of pereopods 1—4

shorter than anterolateral spines of pereonite 4.

Natasome 1.3 times as long as anterior body part.

Pereonites 5—7 subequal in size, their ante-

rolateral spines long, directed slightly backward.

Pleotelson triangular, length 1.1 width, ter-

minal spine straight, length 0.35 pleotelson

length, lateral sides with 3 long subequal spines

directed backward, parallel to lateral spines of

pereonites 5-7. Anterior part of pleotelson with

2 long dorsal medial spines, posterior part with 2

pairs of shorter spines. Preanal ventral process of

pleotelson broad, semicircular, turned backwards,

in ventral view almost completely covering anal

operculum. Body surface covered by small

acute tubercules, all spines on body, coxae and

antennae with long hair-like setae.

Antenna | (Fig. 8j) length 0.25 body length,

article 1 elongate, 2.2 times as long as wide,

article medial spine 1.6 times as long as article

width, lateral lobe extends almost to distal up of

article 2. Article 2 with 2 distal setae, length 0.4

article 1 length, article 3-5 lengths 1.1, 0.3, 0.9

article 2 length respectively, flagellum with

numerous short articles, some with aesthetascs.

Antenna 2 (Fig 7a). Basal articles 1-3 gradu-

ally lengthening, long lateral spine of article 1

subequal in length to coxal spines of pereopods

1-4, article 2 distinctly longer than article 1,

article 3 subequal in length to article 2, with 1 lat-

eral and 1 medial spines, article 4 half as long as

article 3, length 0.6 width, article 5 twice as long

as total length of all preceding articles, article 6

slightly narrower and 1.4 times as long as article

5. Flagellum with elongate articles, most broken

off.

Mandibles (Figs 71, j). Incisor process of both

mandibles with 4 cusps, lacinia mobilis broaden-

ing distally, almost half as long as incisor process,

with 5 teeth, spine row with 12 and 13 members

in left and right mandibles respectively, molar

process tip denticulated, with 6 setulose distal

setae, palp 1.2 times as long as mandibular body,

first 2 articles equal in width, article 2 twice as

long as article 1, with 4 distomedial setac, article

3 with setae row on medial margin.

Maxilla | (Fig. 7g). Inner lobe width 0.7 outer

lobe width, with 13 claw-like setae, some

serrated.

Maxilliped (Fig. 7f). Basis with 8 coupling

hooks, endite with 6 fan-setae distally, endite

width 0.6 basis width, lateral length of palp

article 2 almost equal to article width. its width

twice endite width, article 3 length 0.3 article 2

lateral length and 0.7 medial length. medial mar-

gin with 17-18 stout setae, article 4 medial lobe

and article 5 with 5 simple distal setae each. Epi-

pod rounded distally, lateral margin semicircular,

length 2.3 width.

Pereopods 2—4 broken, only bases present.

Pereopods 1—4 bases subequal in length, those of

pereopods 5-7 slightly longer, all bases with row

of small spines and plumose setae on dorsal

margin. Pereopod | (Fig. 8d) almost half as long

as pereopods 5-7, pereopods 5-7 (Fig. Sa-c)

subequal in size and shape, with tubular articles,

pereopod 6 longest. Ischia of pereopods | and 5-7

about half corresponding basis length (0.43, 0.50,

0.48, 0.41). Ratio of lengths (pereopods | and

5-7) carpi : bases: 0.76, 0.87, 0.84, 0.81 respec-

tively; propodi ; bases: 0.48, 0.87, 0.90, 0.86; and

dactyli : bases: 0.15, 0.56, 0.51, 0.47. Carpi 5—7

slightly and propodi half as wide as bases. Ventral

margins of carpi and propodi 5—7 with thin, short

unequal bifid setae, dorsal margins denticu-

lated, with thin short simple setae. Dactyli with

numerous thin simple dorsal setae.

Pleopod 1 (Fig. Se) relatively broad, broaden-

ing distally, length 2.8 distal width, ventral sur-

face lacking setae, but with small high tubercules,

distal lobes 70° to each other at distance 0,2 pro-

topod length from distal tip. Inner lobes of distal

margins semicircular, with numerous thin long

setae, outer lobes narrow, expanded, slightly

curved inward.

Pleopod 2 (Fig. 8f) length 1.7 width, endopod

insert in midlength of protopod medial margin,

stylet subequal in length to protopod, strongly

tapering distally, extending beyond distal margin

of protoped, exopod extrinsic musculature

extending almost throughout protopod. Pleopods

Land 2 much larger and thicker than remaining

ones.

Pleopod 3 (Fig. 8g) length 0.6 pleopod 2

length, endopod subrectangular, length 1.4 width,

with 3 distal plumose setae, exopod narrow, width

0.3 endopod width, 1.6 times as long as endopod,

with 9 distal plumose setae and row of thin

simple marginal setae,

Pleopod 4 (Fig. 8h). Endopod semicircular,

exopod 2-articulated, length 0.8 endopod length

and width 0.6 endopod width, with 1 distal setae.

Uropod (Fig. 7k) 0.46 pleotelson length,

extending slightly beyond terminal spine of pleo-

telson. Protopod slightly broadening distally, with

282 M. V. MALYUTINA

8 medial and 6 distal setae, length 4.2 width.

Endopod 0.4 width of protopod and slightly

shorter (0.97), with row of small medial and 6-7

distal unequal bifid setae, exopod slightly nar-

rower than endopod, its length 0.8 endopod

length, with 3 distal unequal bifid setae.

Female paratype. In appearance and structural

pattern, the female is similar to the male. Antenna

| (Fig. 7e) about 1.5 times shorter than in male,

article 1 length 2.1 width, medial spine length

0.65 article length, articles 2-5 length ratio simi-

lar to that of male, flagellum with about 30 elon-

gate articles. Operculum (Figs 7e, d) oval, length

1.1 width. Keel rounded ventrally, covered with

small acute tubercules, distal part turned upward,

Etymology. The species is named after the late

outstanding American carcinologist Dr Robert

Menzies who described, among others, many

species of the genus Storthyngura.

Remarks. The new species is most similar to S.

(riplispinosa by body shape, number and arrange-

ment of body spines, and tubular last three pairs

of legs lacking natatory setae. The major differ-

ence between the species is the number of dorsal

spines on the pleotelson: six in the new species

and four in S. friplispinosa. S. menziesi is also

distinguished by more slender body (body length

is 3.2 times body width in the new species and 2.8

times in S. triplispinosa); and a longer exopod of

uropod (endopod to exopod length ratio is 1.21 in

the new species and 1.64 in S. triplispinosa). Both

S. benti and S. zenkevitchi have a similar shaped

pleotelson with three lateral spines and a rela-

tively short terminal spine but these species have

different numbers and arrangements of dorsal

spines on the pleotelson: S. benti has two medial

spines and a pair of tubercules on both sides of

posterior spine, and S. zenkevitchi has one dorsal

medial spine on the anterior part of the pleotelson

and three tubercules in posterior part.

Storthyngurella triplispinosa (Menzies, 1962)

comb. nov.

Figures 9, 10d

Storthyngura triplispinosa Menzies, 1962: 149, fig.

38A-E. — George and Menzies, 1968a: 177. — George

and Menzies, 1968b: 298, fig. 12B.

Material examined. Argentine Basin, RV Dmitrii

Mendeleev, cruise 43, stn 4094 (60°42’S, 41°03'W),

4670 m (pleotelson of female); stn 4086 (60°50’S,

41°10'W), 6130-6420 m (female, 6.4 mm long; frag-

ments of 2 specimens); stn 4090 (60°52’S, 40°56'W),

6145-5550 m (fragments of 2 specimens); RV

Akademik Kurchatov, cruise 11, stn 914 (56°21'S,

50°48"W), 5650-6070 m (anterior part of female with

oostegites); cruise 43, stn 4904 (32°10'S, 63°05'W),

3880-3930 m (1 male, 7.5 mm long).

Remarks. These specimens were identified as

S. triplispinosa only on the basis of external sim-

ilarities with holotype drawings; there was no

opportunity to compare my specimens with type

material. Besides the dorsal view of an entire

specimen, Menzies presented only drawings of

some details: antenna 1, pereopod 1, distal part of

pleopod | and uropod. These details in my speci-

mens are the same. Here I provide drawings of

other appendages.

Storthyngurella digitata (Menzies, 1962)

comb. nov.

Figure 10e

Storthyngura digitata Menzies, 1962: 146, figs 37

FC,

Material examined. Argentine Basin, RV Dmitrii

Mendeleev, cruise 43, stn 4109 (38°40'S, 48°08'W),

5225 m (1 female without pleotelson); RV Akademik

Kurchatov, cruise 43, stn 4893 (36°12'S, 49°10'W),

4630 m (1 female, 6.5 mm; 1 male 6.8 mm long).

Remarks, Although the body processes of

S. digitata are neither long nor spine-like, but bul-

bous, and the pleotelson does not have a long ter-

minal spine, this species is placed in Storthyn-

gurella because of the typical arrangement of

processes on the body, the first arcticles of both

antennae, narrow carpi and propodi of pereopods

5—7, the morphology of first pairs of male

pleopods, and three distal setae on the endopod of

pleopod 3.

Acknowlegements

The project was undertaken with partial financial

support of RFBR (grant N 97-049724).

References

Beddard, F.E., 1885. Preliminary notice of the Isopoda

collected during the voyage of H.M.S. ‘Chal-

lenger’. - Part II. Munnopsidae. Proceedings of the

Zoological Society of London 1885: 916-925.

Birstein, Ya. A., 1969. Crustacea Isopoda from the

Romanche Trench, Byulleten’ Moskovskogo Obsh-

chestva Ispytatelei Prirody. Otdel Biologicheskii 3:

50-69. [in Russian]

Malyutina, M.V., in press. New information on Acan-

thocope Beddard, 1885 (Crustacea, Isopoda, Mun-

nopsidae). Russian Journal of Marine Biology,

George, R.Y. and Menzies, R.J., 1968a. Distribution

and probable origin of the species in the deep-sea

isopod genus Storthyngura. Crustaceana 15:

171-187.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 283

Figure 9. Storthyngurella triplispinosa (Menzies, 1962). All except f, male from stn 4904: a, dorsal view

of body; b, maxilla 2; c, maxilla 1; d, left mandible; e, antenna 1; f, antenna 1 of female from stn 4904;

g, maxilliped; h, pereopod 1; i-k, pereopods 5-7; 1, uropod; m-q, pleopods 1-5.

284 M. V. MAL YUTINA

i v Vis

Jr )

A A aN NA

Figure 10. Structure of male operculum in five species of Storthyngurella gen. nov. and one species of

Storthyngura: a, Storthyngurella hirsuta sp. nov. ; b, S. wolffi sp. nov.; c, S. menziesi sp. nov.;

d, S. triplispinosa (Menzies, 1962); e, S. digitata (Menzies, 1962); f, Storthyngura birsteini Menzies, 1962.

STORTHYNGURELLA, NEW GENUS OF ISOPOD FROM DEEP-SEA BASINS 285

George, R.Y. and Menzies, R.J., 1968b. Species of

Storthyngura (Isopoda) from the Antarctic with

descriptions of six new species. Crustaceana 14:

275-301.

Menzies, R.J., 1962. The isopods of abyssal depths in

the Atlantic Ocean. Vema Research Series 1:

79-206.

Wägele, J.-W., 1989. Evolution und phylogenetisches

System der Isopoda. Stand der Forschung und neue

Erkenntnisse. Zoologica (Stuttgart) 140: 1-262.

Wilson, G.D. and Hessler, R.R., 1980. Taxonomic char-

acters in the morphology of the genus Eurycope

(Isopoda, Asellota) with a redescription of

Eurycope cornuta G.O. Sars, 1864. Cahiers de

Biologie Marine 21: 241-263.

Wilson, G.D., Kussakin, O.G. and Vasina, G.S.,

1989. A revision of the genus Microprotus

Richardson with descriptions of two new

species, M. acutispinatus and M. lobispinatus

(Asellota, Isopoda, Crustacea). Proceedings of

the Biological Society of Washington 102(2):

339-361.

Wolff, T., 1956. Isopoda from depths exceeding 6000

meters. Galathea Report 2: 85-157.

Memoirs of Museum Victoria 57(2): 287-310 (1999)

REVISION OF LAPHYSTIOPSIDAE (CRUSTACEA: AMPHIPODA):

NEW AND OLD SPECIES FROM SOUTH CHINA SEA, SOUTHEASTERN

AUSTRALIA, FALKLAND ISLANDS AND WESTERN ATLANTIC OCEAN

J. L BARNARD*

National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA

Abstract

Barnard, J.L., 1999. Revision of Laphystiopsidae (Crustacea: Amphipoda): new and old

species from South China Sea, southeastern Australia, Falkland Islands and western Atlantic

Ocean. Memoirs of Museum Victoria 57: 287-310.

Collections from Bass Strait and eastern Tasmania reveal two new species of Laphystiopsis,

L. wulgi sp. nov., and L. zomerysis sp. nov. of the rarely encountered family Laphystiopsidae.

To verify differences between Prolaphystiopsis and the type of the family, Laphystiopsis, old

but unreported materials of Laphystiopsis planifrons Sars in the Smithsonian’s National

Museum of Natural History collections were reviewed and are presented herein. Prolaphys-

tiopsis platyceras, type of the genus, was borrowed and redescribed. Sars” (1895) and Schel-

lenberg’s (1931) differentiation of the genera was based on erroneous observations of the palp

of maxilla 1. A review of the family is presented, with updating of diagnoses and literature.

Prolaphystiopsis differs from Laphystiopsis in the narrow rostrum, bulging head, and enlarged

and lobate article | of antenna 1. Luphystiopsis planifrons is newly recorded from the western

Atlantic Ocean. Laphystiopsis iridometrae Shoemaker, 1919 from the South China Sea is

reviewed and illustrated for the first time. Prolaphystiopsis latirostris Ledoyer, 1986 is

removed to Laphystiopsis. Individuals of the family are presumed to be inquiline parasites or

commensals of sessile marine invertebrates, particularly crinoids, but are rarely noticed by

amphipodologists.

Introduction

While examining amphipods from Bass Strait in

Museum Victoria, Jean Just discovered a speci-

men of Laphystiopsidae, a family rarely col-

lected. Later [in 1990] I visited him and Gary

Poore at Museum Victoria and continued sorting

through similar collections and found one more

specimen from eastern Tasmania, The two speci-

mens are reported on here as two new Australian

species.

The remarkable morphological resemblance of

L. zomerysis sp. nov. and L. wulgi sp. nov. to the

North Atlantic Laphystiopsis planifrons Sars,

1895 in somatic aspect and the perfunctory

generic distinction (l- or 2-articulate palp of

maxilla 1) between Laphystiopsis and Prolaphys-

* Jerry Laurens Barnard died on 16 August 1991

shortly after completing a first draft of this manuscript.

The editor thanks Elizabeth Harrison-Nelson for send-

ing the paper to the Memoirs of Museum Victoria as

Jerry had intended. J.D. Thomas, J.K. Lowry, J. Just

and I made corrections and improvements to the origi-

nal draft but these have been slight. Gary C. B. Poore

287

fiopsis necessitated reexamination of specimens

of L. planifrons to verify the uniarticulate condi-

tion of the palp on maxilla 1, and to search for

other generic differences, In this examination,

L. planifrons was found to occur in the western

Atlantic Ocean. Maxilla 1 palp proved to be 2-

articulate despite the analysis of Sars (1895).

Thus, the only known difference between Prola-

phystiopsis and Laphystiopsis evaporated. After

examining the syntypes of Prolaphystiopsis

platvceras, type species of the genus, it was

determined that the narrow rostrum, bulging

head, and enlarged and lobate article 1 of antenna

l are new characters that diagnose Prolaphys-

tiopsis. Laphystiopsis ornitorhynchus Bulycheva

is transferred to Prolaphystiopsis.

Review of the scant literature of this family

revealed that Laphystiopsis iridometrae Shoe-

maker, 1919 from the “China Sea” had been only

briefly described but never illustrated, The holo-

type deposited in Smithsonian collections pro-

vided a detailed analysis of that species.

Although more than 20 juveniles were also pre-

sent in the Smithsonian collections, only the

holotype was fully adult.

KK J. L. BARNARD

Species in Laphystiopsidae are assumed to be

“parasites” on other organisms, much in the same

manner that Lafystiidae (Sars, 1895; Bousfield,

1987) are parasitic on fish, Indeed, L. iridometrae

was originally found by Dr Austin H. Clark

embedded in tissues of the crinoid /ridometra

melpomene ( Iridometra adrestine |A M1. Clark])

and given to Shoemaker who described it spar-

ingly. IF not for the excellent work of Bousfield

(1987), in collecting and soliciting specimens of

Lafystiidae from fish, very little would be known

Of its presumed sister family. Beeause they have

been collected secondarily from their hosts,

laphystiopsids appear in amphipod collections

only accidentally. Commensal amphipods. that

eling to the host initially on collection can be

easily lost during routine curatorial activities, c.g.,

alcohol and/or container changes. For a more

complete understanding of the group, active

searches should be undertaken focusing espe-

cially on sessile or slow moving invertebrates

such as gorgontans and erinoids.

Rach species is diagnosed; the drawings form

the principal descriptive material; the deseriptions

for cach are composed only of comments ampli-

lying the drawings, Diagnoses and descriptions

are nol congruous among species. The diagnosis

of the family is based on new observations, not all

of which have been confirmed in all species of the

family. Geographie codes listed in brackets lor

cach species taken from Barnard and Karaman

(1991). Length and width of coxae are the same as

length and width of any article on the pereopods:

the words deeper and broader are not appropriate.

Material is deposited in Museum Victoria,

Melbourne (NMV) and National

Natural History, Smithsonian

Washington (USNM).

Institution,

Laphystiopsidae Stebbing, 1899

Diagnosis, Rostrum present or absent, if present

well developed, reaching at least halfway along

peduncular article of antenna 1. Accessory flagel

lum absent or uniarticulate, Field of mouthparts

quadrangular. Mandibular molar scarcely tritura-

tive or not, rakers absent or L raker present. Inner

plate of maxilla I feeble, poorly setose (1 seta or

naked), outer plate with 5 7 spines; palp large,

2 articulate. Palp of maxilliped 4-articulate.

Coxae Let small, quadrate or anteroposteriorly

rectangular or ovate, Occasionally disjunct from

each other, Gnathopods 1-2 feeble, simple. Uro-

somites free. Uropod 3 biramous, outer ramus

longer than peduncle, Telson short, entire.

Description. Body broad, depressed, weakly to

strongly carinate on pleonites 3-4 or 1-3 only.

Museum of

Urosome slightly flattened, urosomite | elongate

or not. Head flat, rostrum large and spatulate [or

absent]; eyes present or absent; sides of head

bulging or not. Antenna | dominant, peduncle

short, flagellum elongate, article 1 of flagellum

pubescent or not or developed as a weak callyn-

ophore; antenna 2 often as long as antenna | but

of lesser thickness and lesser dominance, Mouth-

parts generally feebly armed. Upper lip broad,

incised. Mandibular incisors ordinary, not attenu-

ate for piercing, toothed; palp well developed, 3-

articulate, very poorly setose, armaments mostly

scales. Inner lobes of lower lip fleshy and well

developed or absent. Inner plate of maxilla |

small, ovate, with | small seta or none, outer plate

with 5 spines: palp 2-articulate (but see the dis-

cussion of Prolaphystiopsis — ornitorhynchus

below). Plates of maxilla 2 narrow, inner setose

medially or not. Maxillipeds small, feebly armed,

plates ordinary except basal article of outer plate

elongate; palp large or small. Coxae variable,

short and evenly extending or middle coxae

longer, occasionally coxae so small as to be dis-

junct, Gnathopods simple, carpi elongate. Pere-

opods 5 7 increasingly elongate or not, article 2

increasingly expanded. Epimeron 2 dominant.

Pleopod 3 significantly shorter than pleopods

| 2. Uropods | and 3 exceeding uropod 2 (as far

as known), outer rami slightly shortened or not:

peduncle of uropod 3 scarcely elongate. Telson

ovale,

Variables. Prolaphystius departs from the typical

characterization in the total absence of a rostrum,

lack of inner lobes on the lower lip, larger middle

coxae, excavate coxa 4, long carpus of pereopod

3, lysianassid-like pereopods 5 7, elongate tel-

son, and elongate urosomite 1. Specimens of this

antarctic genus have not been observed; it may

belong in another family. It keys out to the gener-

alized “melting pot” of Eusiridae but differs from

that group in the reduction of spines on the outer

plate of maxilla 1 to 5 (versus generally 9-11), the

feeble palp of the maxilliped, the elongated article

on which the outer plate of the maxilliped is

attached; the reduction of raker spines to 1 (versus

generally 5+), and the elongation of urosomite 1.

Relationship. See Barnard and Karaman (1991)

for comparisons with the families: Iphimediidae;

Lafystiidae; Eusiridae (Calliopiidae, Pleustidae);

Oedicerotidae; Stilipedidae; Astyridae; Colo-

mastigidae; Maxilliphimediidae. The family

differs from Lafystiidae in the 4-articulate (versus

2-articulate) palp of the maxilliped, the reduction

of spines on the outer plate of maxilla | from 7 to

5, the presence of a molar, and well developed

palp on maxilla 1, and except for Prolaphystius,

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 289

the presence of inner lobes on the lower lip, and

the poorly developed and poorly setose (1 versus

2+ setae) inner plate of maxilla 1.

The Laphystiopsidae are similar to the Eusiri-

dae, Calliopiidae and Pleustidae but with feeble,

simple gnathopods; feeble, and poorly setose

maxillipedal palps. The lower lip of the type

genus is like the characteristic labium of Pleusti-

dae, and the inner lobes of Laphystiopsis are large

and fleshy, whereas the inner lobes are lost in

Prolaphystius.

Like the Lafystiidae, the Laphystiopsidae may

have some roots in Iphimediidae. Lafystiidae

have an acuminate coxa 4 and otherwise are

similar to the Iphimediidae except for the reduced

palp on the maxilliped. The Laphystiopsidae are

so diverse that they must be characterized indi-

vidually. Laphystiopsis and Prolaphystiopsis

differ from Iphimediidae in the very short

nonacuminate coxae and flattened (though

strongly rostrate) head; Prolaphystius differs in

the nonrostrate head and thin geniculate urosome.

The Stilipedidae and Astyridae bear large outer

lobes on the maxillipeds and strongly dominant

carpi on the gnathopods.

All but one genus of Oedicerotidae have elon-

gate peduncles on uropod 3, and that genus,

Metoediceros, like other oedicerotids, has slightly

or strongly subchelate gnathopods, a dispropor-

tionately elongate pereopod 7, strongly sctose

pereopods, and unnotched upper lip.

Corophioids have triturative molars and usually

subchelate or strongly setose and specialized

gnathopods.

The Laphystiopsidae could be confused with

Phoxocephalidae that have an elongate, flat, spat-

ulate rostrum, but Laphystiopsidae differ in the

non-fossorial pereopods and antennae, and the

large biramous uropod 3 lacking article 2 on the

outer ramus (occasionally true in Phoxocephali-

dae, but only with short uropod 3), the small

coxae, the weak antenna 2 and the uncleft telson.

The uniformity of Laphystiopsidae is broken by

the loss of the rostrum in Prolaphystius. That

genus has a long urosomite | as in Dulichtidae,

but otherwise has little similarity to that family.

The Laphystiopsidae are analagous to Maxil-

liptidae in that both are inquilinous on sessile or

semi-sessile invertebrates. Maxilliptidae occupy

gorgonians (Thomas, 1996) while the Laphys-

tiopsidae are definitely known from crinoids but

possibly also from coelenterates (collected in

fields of Lophelia in Norway). The two families

resemble each other in what may be superficially

convergent characters as follows: similar coxal

shapes, bulging ocular regions, presence of cal-

lynophore, feeble maxillae and maxillipeds, fec-

ble gnathopods, huge oostegites, and weak telson

(except Prolaphystius). In contrast, the Maxillipi-

idae are characterized by an enormously elongate

pereopod 6, serrate spines on outer plate of max-

illa 1, thickened article 2 of maxilliped palp, elon-

gate peduncle of uropod 3, unnotched upper lip,

and one or both mandibles with 21 slender rakers.

Maxillipiids are free-living on gorgonians where

they gather in large numbers with their clongate

sixth pereopods stretched outward laterally and

frequently rotating whereas the only known eco-

logical observation of laphystiopsids ts that they

make surficial burrows on crinoids.

Key to genera of Laphystiopsidac

l. Rostrum absent; coxa 4 almost as long as broad, deeply excavate posteriorly

ENE IEOR E T EER ETAT Prolaphystius

Rostrum elongate, apex exceeding first peduncular segment; coxa 4 small,

much wider than long, not excavate posteriorly … nnee ren 2

Article 1 of antenna 1 enlarged, lobate; rostrum apically constricted, margin

rounded; eye lobes strongly bulging

Prolaphystiopsis

Article | of antenna | not enlarged or lobate, lacking apical projection; ros-

trum not constricted, truncate; eye lobes not bulging … ……… Laphystiopsis

Laphystiopsis Sars

Laphystiopsis Sars, 1895: 386.

Type species. Laphystiopsis planifrons Sars, 1895

(monotypy).

Diagnosis. Rostrum well developed, very broad;

ocular lobes not bulging. Article | of antenna |

not grossly lobate. Mandibular molar conical,

unridged. Inner lobes of lower lip present. Palp of

maxilla | uniarticulate. Coxae short, broad, ante-

rior coxae ovate, often not touching serially, coxa

4 much wider than long, not excavate posteriorly.

Pereopods 3-4 alike, with carpus very short.

Pleonite 3 dorsally carinate but not forming hori-

zontal shelf, urosomite | carinate and saddled.

Telson short, oval.

Description, Article | of antenna | weakly cari-

nate to strongly produced apically and massive.

290 J, L. BARNARD

Coxae produced forward or not. Articles 2-7 of

pereopod 3 like pereopod 4, Pleonites 3-4

carinate and pleonite 4 saddled, or only pleonites

1-3 carinate.

Included species.

L. wulgi sp. nov., eastern Tasmania, Australia

[782]

L. zomerysis sp. nov., eastern Bass Strait,

Australia, bathyal [782]

L. species “k”, herein [254]

Transferred species.

1952 to

L. iridometrae (Shoemaker, 1919) L Sie ents T Bul cheve

(Vader, 1978), near Hong Kong, inquilinous < Ornitöraynehus WENEN,

[6521] Prolaphystiopsis.

L. latirostris Ledoyer, 1986, Geyser Bank,

Indian Ocean, abyssal [618A]

L. planifrons Sars, 1895 (Stephensen, 1926,

1931, 1938; Gurjanova, 1951), north boreal

Atlantic, sublittoral to bathyal [240 + B]

Distribution. Marine; South China Sea; boreal

North Atlantic, 167-900 m; northeast of Geyser

Bank near Madagascar, Indian Ocean,

2300-2500 m; and southeastern Australia,

102—1000 m; often on crinoids or corals.

Key to species of Laphystiopsis

l. Pleonite 4 lacking dorsal process … nana. L. latirostris

— Pleonite 4 with dorsal process … nanne ereen venen nner 2

2. Epimeron 2, posteroventral corner truncate „s... L. zomerysis

3, Dorsal cusps of pleonites 3-4 pointed … en. L. planifrons

Dorsal cusps of pleonites 3-4 rounded … eneen. 4

4. Rostrum reaching or exceeding second peduncular segment; ventral margin

of coxa 4 even, broadly emarginate … naren eneen L. iridometrae

Rostrum barely reaching end of first peduncular segment; ventral margin of

coxa 4 produced midventrally .....

Laphystiopsis planifrons Sars

Figures 1-3

Laphystiopsis planifrons Sars, 1895: 386, pl. 135.—

Norman, 1895: 488— Stebbing, 1906: 209

Stephensen, 1926: 73; 1928: 185, fig. 36 (6-10); 1929:

106, fig. 25 (156): 1931a: 208; 1938b: 182.—Oldevig,

1933: 113 — Gurjanova, 1951: 495, fig. 323.—Oldevig,

1959: 48.

Material examined, U.S. Fish Commission Lot 547,

which in written records (not on label) is: Gloucester

Fisheries Expedition, on schooner Proctor Brothers,

Sable Island Bank off Nova Scotia, 7 Nov 1879,

43° 16'N, 60°35’ W, 350 fm (written records cite as 640

m), [?dredge], sample includes sea-anemone, pectens,

crinoid Acanthogorgia pinnata, and Alevonium multi-

forum and Pennata borealis USNM 36124 (female

“p” 8.25 mm).

U.S. Fish Commission Steamer A/batross stn 2429,

off Newfoundland, 42°55'30"N, 50°51’00"W, 471 fm,

23 Jun 1885, gray mud, large beam trawl, bottom tem-

perature 38.7°F USNM 31966 (male “o” 6.43 mm).

U.S. Fish Commission Albatross stn 2540, northwest-

ern Atlantic Ocean, off Massachusetts, 39°58’20°N,

70°52'00"W, 7 Aug 1885, 144 fm, large beam trawl,

green sand, bottom temperature 46.7°F (note in vial

reads: “many frags of crinoids were found in bottle

from which these specimens were taken C.R.S[hoe-

maker]”) USNM 230425 (young male “m” 5.06 mm).

woop cs Bea EE | PERRE TE T L. wulgi

Diagnosis. Head slightly longer than pereonites

1-3, scarcely bulging laterally (as seen from dor-

sal view); pleonites 1—4 each with conspicuous

dorsal bulge or carina; pleonite 4 not longer than

pleonite 3; coxa 5, 50% as long as wide; coxa 6

bilobed and wide anteroposteriorally (80% as

wide as coxa 5 and 160% as wide as coxa 7),

60+% as long as wide; epimeron 2 rounded pos-

teroventrally; article 5 of pereopods 3-4, 1.5

times as wide as long; article 2 of pereopod

7 broadly pyriform, articles 3-7 together about

3 times as long as article 2 (from Sars, 1895:

articles 5-7 absent on present material).

Description of female “p”. See illustrations. Head

with apparent glandular tissue in place of eye.

Antennae | broken apically but probably not

longer than antenna 2, flagellum with 13+

articles, callynophore with 6 groups of aes-

thetascs, proximal to distal = 2rudimentary-2-5-4-

3-7, following articles 1-8 aesthetase formula =

2-3-3-3-0-3-0-2. Antenna 2 ordinary, flagellum

20-articulate. Callynophore articles without

ridges besides insertion points for rows of aes-

thetascs. Accessory flagellum very poorly devel-

oped. Ventral surface of article 1 on antenna | and

medioventral surfaces on articles 3-4 of antenna

2 with sparse ridges, stiff setules, weak scales.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 291

oXP |

Figure 1. Laphystiopsis planifrons, unattributed figures = female “p” 8.25 mm; “o” = male “o” 6.43 mm.

Capital letters in figures refer to parts; lower case letters to left of capital letters refer to specimens and to the

right refer to adjectives as follows: B, body; C, coxa; D, dactyl; E, epimeron; F, accessory flagellum;

G, gnathopod; H, head; L, labium; M, mandible; O, oostegite; P, pereopod; PL, pleopod; R, uropod;

T, telson; U, upper lip; V, palp; W, pleon; X, maxilla; Z, gill; d, dorsal; i, inner; m, medial; o, outer; r, right;

t, left.

292 J. L. BARNARD

Figure 2. Laphystiopsis planifrons, female “p” 8.25 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA)

Figure 3. Laphystiopsis planifrons, unattributed figures = female “p” 8.25 mm; “o”

male “o” 6.43 mm.

293

294 J. L. BARNARD

One raker present on left mandible. Right lacinia

mobilis very transparent, with 3 teeth, in direct

view showing as column. Comb of setae on palp

article 3 of complexity shown by Sars (1895).

Pleopods 1-2 alike but pleopod 3 smaller, like

pleopods of L. zomerysis; all inner rami with

12-13 articles, outer with 13. Most spines of

uropods 1-2 and setae of uropod 3 missing and

marked with pits. Cuticle with saw-tooth ridges

and denticles similar to L. zomerysis below.

Male “o”. Like female but callynophore twice

as long, right mandibular palp like female but left

with primary marginal setae absent, other scales

present.

Illustrations. Pereopod 6 not enlarged, see body;

oostegite of pereopod 4 not enlarged, see body;

drawing of antenna 2 reduced in relation to

antenna 1; apices of maxillae badly eroded; max-

illipeds missing (see drawing taken from male

bean

0°; most spines on uropod 3 broken.

Relationship. Because this is the type species of

the genus, it will serve as model for comparison

of the other species.

Distribution. Northeast Atlantic generally from

the Skagerrak northward along Norwegian coast

and into Norwegian Sea, 50-900 m; here

recorded for first time from northwestern Atlantic

Ocean off Massachusetts, Newfoundland and

Nova Scotia, 264-861 m.

Laphystiopsis iridometrae Shoemaker

Figures 4-6

Laphystiopsis — iridometrae Shoemaker, 1919:

245 246. — Vader, 1978: 126-127 (discussion only).

Material. Holotype. U.S. Fish Commission Albatross

stn 5310, South China Sea, near Hong Kong, 21°33'N,

116°13°E, 100 fm, 4 Nov 1908, 12 ft tanner beam trawl,

sand, shell, bottom temperature 65.5°F, parasitic upon

crinoid /ridometra melpomene A.H. Clark |= I. adres-

tine A.H. Clark], USNM 49599 (male “g” (newly

designated letter) 4.55 mm, with penial processes of

regular size).

Other material. Albatross stn 5311, South China Sea,

near Hong Kong, 21°33°N, 116°15'E, 88 fm, 4 Nov

1908, coarse shell sand, 12 foot tanner beam trawl, mud

bag on /ridometra melpomene, USNM 49801 (3

subadults and 5 tiny juveniles, including subadults “h”

3.11 mm and “i” 3.15 mm; lacking penes and ooste-

gites). USNM 49600, same data as 49599 (1 subadult

and 13 tiny juveniles).

Diagnosis. Head equally as long as pereonites

1-3, scarcely bulging laterally (as seen from dor-

sal view); pleonites 1-4 each with conspicuous

dorsal bulge or carina; pleonite 4 not longer than

pleonite 3; coxa 5 60% as long as wide: coxa 6

bilobed and wide anteroposteriorally (more than

90% as wide as coxa 5 and 135% as wide as coxa

7), 70+% as long as wide; epimeron 2 rounded

posteroventrally, corner rounded; article 5 of

pereopods 3-4 1,2 times as wide as long; article 2

of pereopod 7 broadly ovate, articles 2-7 together

about equally as long as article 2.

Description of holotype male “g” 4.55 mm. See

illustrations. Head with apparent glandular tissue

in place of eye. Left antenna 1 broken apically but

right one not longer than antenna 2, flagellum

with 15 articles, callynophore with 3 groups of 3

setae each, following articles 2-8 aesthetasc for-

mula = 2-3, 2, 1-2, 0, ?, 0, 1-22 (damaged).

Antenna 2 rather short and stunted on left side,

but normal on right (see body illustration, with

left antenna 2 replaced by right), flagellum 15-

articulate. Callynophore article without ridges

besides insertion points for rows of aesthetascs.

Accessory flagellum very poorly developed. Ven-

tral surface of article 1 on antenna | and

medioventral surfaces on articles 3-4 of antenna

2 with sparse ridges, stiff setules, weak scales.

Maxilliped, inner plate with 2 short marginal

setae (1 stout) and | stout + 2 thin apical setae,

outer plate with 3 marginal setae. One possible

raker present on left mandible. Right lacinia

mobilis very transparent, with 4 teeth. Comb of

setae on article 3 of palp much less complex than

shown by Sars (1895) for Laphystiopsis plani-

frons and L. zomerysis to follow. Pleopods 1-2

alike but pleopod 3 smaller, drawing of pleopod 1

like pleopod 2, lengths of peduncle and rami of

uropod 3 shown; however, all inner rami with 10

articles, outer with 11. Most spines of uropods

1-2 and setae of uropod 3 missing and marked

with pits. Cuticle with saw-tooth ridges and

denticles similar to L. zomerysis below.

Illustrations. Palp of mandible reduced in relation

to body of mandible.

Relationship. See L. wulgi for differences. Differ-

ing from other members of the genus in the very

short set of articles 3-7 on pereopods 5-7.

Distribution, South China Sea near Hong Kong,

on crinoids; 161—183 m.

Laphystiopsis zomerysis sp. nov.

Figures 7-9

Material examined. Holotype. Australia, Victoria, S of

Point Hicks, 38°21.90’S, 149°20.00’ E, 1000 m, 23 Jul

1986, WHOI epibenthic sled, G.C.B. Poore et al. on RV

Franklin (SLOPE stn 32), NMV J18521 (female “f”

4.60 mm).

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 295

Figure 4. Laphystiopsis iridometrae, unattributed figures = holotype male “g” 4.55 mm; “i” = subadult “i”

3.15 mm.

296 J. L. BARNARD

Figure 5. Laphystiopsis iridometrae, unattributed figures = holotype male “g” 4.55 mm; “i” = subadult “i”

3.15 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 297

Figure 6. Laphystiopsis iridometrae, unattributed figures = holotype male “g” 4.55 mm; “i” = subadult “i”

3.15 mm.

J. L. BARNARD

298

Figure 7. Laphystiopsis zomerysis, holotype female “f? 4.60 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 299

Figure 8. Laphystiopsis zomerysis, holotype female “f” 4.60 mm.

300 J. L. BARNARD

Figure 9. Laphystiopsis zomerysis, holotype female “f” 4.60 mm. Stick figures of pleopods show lengths of

parts.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 301

Diagnosis. Head about 1.25 times as long as pere-

onites 1-3 together, scarcely bulging laterally (as

seen from dorsal view); pleonites 1—4 each with

conspicuous dorsal bulge or carina; pleonite 4 not

longer than pleonite 3. Coxa 5 about half as long

as wide. Coxa 6 bilobed and wide anteroposterio-

rally (more than 80 % as wide as coxa 5 and

170% as wide as coxa 7), about 60% as long as

wide. Article 5 of pereopods 3—4 about 1.3 times

as long as wide. Article 2 of pereopod 7 narrowly

pyriform, articles 2-7 together about 3.25 times

as long as article 2. Epimeron 2 not beveled,

almost quadrate, Posteroventral corner quadrate.

Description. See illustrations. Individual very

close to ecdysis, with duplicated parts easily seen

inside appendages. Wholemount view with uro-

somite 3 slightly tilted towards observer to show

telson. Setae of oostegites rudimentary. Ventral

surface of article 1 on antenna | and article 3 on

antenna 2 with dense, minute, sharp and ragged

ridges and projecting scales and short stiff

embedded setules, forming a surface resembling

velcro. Left accessory flagellum illustrated, right

with 2 setae. Inner (or ventral) surface of cal-

lynophore with 4 crosswise sharp meandering

ridges, similar ridges present on next 7 flagellar

articles, ridge numbers on articles 2-8 as follows:

1-2-2-2-1-1-1, vestigial ridge on article 9; cal-

lynophore also with 3 sets of 1-2 aesthetases,

next instar apparently to develop many more mul-

tiple sets. Antenna | broken apically, thus rela-

tionship to length of antenna 2 unknown. Right

mandible only with deeply bifid lacinia mobilis,

no raker spines; left lacinia mobilis broadly

toothed like incisor, not distinctly separated from

mandibular body (not a next-instar duplicate of

incisor because that complex observable deep

within body of mandible), therefore 2 spines

proximal to incisor called rakers. Outer plate of

maxilla 1 with 6 spines, the sixth smallest and

thinnest overlapping face of fourth spine from

medial margin; inner plate lacking seta,

Pleopods: see stick-figures of lengths, pleopods

1-2 alike, pleopod 3 much shorter, outer rami

with 11 articles, inner with 10, coupling hooks 2,

peduncles not setose. All setae of uropod 3

broken off, their insertions marked as circles at

the bases of the hooked serrations on margins of

rami.

Illustrations. Drawing of left mandibular palp

reduced compared with body of mandible but

apex of right palp drawn at same magnification as

body of mandible.

Etvmology. From Greek, zomerysis, meaning

“spoon” or “ladle”, referring to the rostrum of this

animal.

Relationship. The new species differs from

L. latirostris in dorsal armament and coxae. The

outer and inner plates of maxilla 1 of L. latirostris

were not described. The lower lip of L. zomerysis

only about half as long as coxa 5 and shorter than

coxa 7.

There is an uncanny resemblance of L. zomer-

ysis to Laphystiopsis planifrons in general body

aspect but L. zomervsis differs from L. planifrons

in the shorter and less robust rostrum, the

stronger dorsal protrusions on pleonites 1-2, a

blunter protrusion on pleonite 4, a broader notch

on the upper lip, a definite triturative area on the

molar, lack of a seta on the inner plate of maxilla

1, and, the squared posteroventral corner of

epimeron 2. Differing from L. iridometrae in the

squared epimeron 2 and the longer set of articles

3-7 on pereopods 5-7.

Distribution. Australia, eastern Bass Strait, 1000 m.

Laphystiopsis wulgi sp. nov.

Figures 10-12

Material. Holotype. Tasman Sea, 15 km E of Maria

Island, Tasmania, 42°37'S, 148°20°E, 9 Oct 1984, 102

m, WHOL epibenthie sled, R.S. Wilson (stn SOS-84-1),

NMV J18530 (female “j” 4.28 mm).

Diagnosis. Head only 80% as long as pereonites

1-3, scarcely bulging laterally (as seen from dor-

sal view); pleonites 1-4 each with conspicuous

dorsal bulge or carina; pleonite 4 not longer than

pleonite 3; coxa 4 with midventral projection;

coxa 5 60% as long as wide; coxa 6 bilobed and

wide anteroposteriorally (more than 80 % as wide

as coxa 5 and 200% as wide as coxa 7), 70% as

long as wide; epimeron 2 beveled posteroven-

trally, corner rounded; article 5 of pereopods 3—4

1.1 times wider than long; article 2 of pereopod 7

broadly ovate, articles 2-7 together about 2.2

times as long as article 2.

Description, See illustrations. Setae of oostegites

well developed, oostegites huge, oostegite on

coxa 2 absent. Ventral surface of article 1 on

antenna | and article 3 on antenna 2 with dense,

minute, sharp and ragged ridges and projecting

scales and short stiff embedded setules, forming a

roughened surface. Inner (or ventral) surface of

callynophore without crosswise sharp meander-

ing ridges, no ridges present on flagellar articles,

Callynophore also with 3 sets of 2 aesthetascs

each, next instar not visible. Flagellum of antenna

1 with 19 articles, of antenna 2 with 17, distal

reach of antenna 2 two flagellar articles shorter

than antenna | when stretched out together. Right

302 J. L. BARNARD

Figure 10. Laphystiopsis wulgi, holotype, female “j” 4.28 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA)

303

AAA TTI yore

rr utt in U!

P3 \

Figure 11. Laphystiopsis wulgi, holotype, female “j” 4.28 mm.

304 J. L. BARNARD

ars De =

ZES YY)

ANN

SNN

)

Figure 12. Laphystiopsis wulgi, holotype, female “j” 4.28 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 305

mandible only with deeply bifid lacinia mobilis,

no raker spines, drawing showing incisor flat and

molar only slightly offset unnaturally; left lacinia

mobilis broadly toothed like incisor, not distinctly

separated from mandibular body, therefore 1

spine proximal to incisor called raker. Outer plate

of maxilla | with 5 spines, the fifth smallest and

thinnest overlapping face of third spine from

medial margin; inner plate lacking seta, Pleopods

1-2 alike but pleopod 3 smaller, drawing of pleo-

pod 1 like pleopod 2, lengths of peduncle and

rami of uropod 3 shown as stick-drawing; how-

ever, inner rami of pleopods 1-2 with 12 articles,

outer with 13 and 14 respectively; pleopod 3 inner

ramus with 11, outer with 13 articles. Margins of

rami on uropods densely and minutely serrate (see

offset enlargement on uropod 2). Most spines of

uropod 3 broken off, their insertions marked as

circles at the bases of the hooked serrations on

margins of rami. Telson covered apically and lat-

erally with cuticular ridges.

Illustrations, Wholemount view with urosomite 3

slightly tilted towards observer to show telson.

Left accessory flagellum illustrated, right similar.

Drawing of left mandibular palp reduced in size

compared with body of mandible.

Etymology. Australian Aboriginal, an Aboriginal

spirit.

Relationship. Laphystiopsis wulei differs from

L. latirostris in the parts of the diagnosis involv-

ing dorsal armament and coxa 6. The outer and

inner plates of maxilla 1 of L. latirostris were not

described. The lower lip of L. zomerysis is also

poorly known. In L. latirostris coxa 6 is only

about half as long as coxa 5 and shorter than

coxa 7.

Laphystiopsis wulgi differs from L. zomerysis

in the shorter head, wider coxae 5—6, much

shorter pereopods 5-7, posteroventrally rounded

epimeron 2, and thick articles 4-5 of pereopods

3—4. In addition L. wulgi, based on a single spec-

imen, has much larger oostegites, a character of

uncertain value because the oostegites of the

single specimen of L. zomerysis are immature

(setae rudimentary). It differs from L. iridometrae

in the longer combined set of articles 3-7 on pere-

opods 5-7, and the ventrally produced coxa 4, and

the smaller posterior lobe of coxa 6.

Distribution, Australia, off eastern Tasmania; 102 m.

Laphystiopsis species k

Material, U.S. Fish Commission Fish Hawk stn 1026,

off Marthas Vineyard, Massachusetts, 39°50’30°N,

71°23'W, 182 fm, 8 Sep 1881, green mud and sand, on

Hathrometra tenella (Retzius) (also a label “on Antedon

dentata (Say)”), USNM 38155 (specimen “k”).

Notes. Specimen in poor condition with shriveled

antenna 1, legs 1-4, some broken posterior pere-

opods; pleonite 4 with giant hood, pleonite 3

scarcely produced dorsally, coxae 5-7, head, uro-

somal appendages, pleonites 1-2 and pereonite 7

like L. planifrons; coxae 1-4 tiny, 3-4 with gaps

in front and behind, 1-2 very short but overlap-

ping. Clearly a new species whose formal

description is reserved for better material.

Prolaphystiopsis Schellenberg

Prolaphystiopsis Schellenberg, 1931: 115.

Type species. Prolaphystiopsis

Schellenberg, 1931 (monotypy).

platyceras

Diagnosis. Rostrum well developed but much

narrower from dorsal view than in Laphystiopsis,

sides of head thus bulging laterally on each side

almost as much as width of rostrum. Article | of

antenna | formed into huge lobe. Mandibular

molar conical, weakly ridged. Inner lobes of

lower lip present. Palp of maxilla | biarticulate,

outer plate with 6-7 spines. Coxae short, broad,

anterior coxae ovate, barely touching or weakly

disjunct serially, coxa 4 much wider than long,

not excavate posteriorly. Pereopods 3—4 alike,

with carpus very short, Pleonite 3 very weakly

carinate dorsally, but not forming horizontal

shelf, urosomite 1 carinate and saddled. Telson

short, oval,

Description. Coxae produced forward or not.

Articles 2-7 of pereopod 3 like pereopod 4.

Relationship. Differing from Laphystiopsis in the

narrower rostrum, laterally bulging head and huge

lobe on article 1 of antenna 1.

Included species.

P. ornitorhynchus (Bulycheva, 1952) Sea of

Japan, 167-510 m [391 + B] comb. nov.

P. platvceras Schellenberg, 1931, Falkland Is,

197 m [831]

Distribution. Sea of Japan, and Falkland Islands,

167-510 m.

Prolaphystiopsis platyceras Schellenberg

Figures 13-15

Prolaphystiopsis platvceras Schellenberg, 1931: 115,

fig. 62.

Material examined. “Svenska Siidpolar exp. 1901-03

No 58 11/9 1902, 179 m, Bodentemp. 4.1°, Sand und

Kies, 52°29'S, 60°36'W, S von W Falkland, Typen,

306 J. L. BARNARD

Typsaml. 712”, Swedish State Museum 6622, (8 syn-

types). Lectotype selected herein: female “a” 7.64 mm.

Paralectotypes: “b” female 4.3 mm, “c” broken, sex not

determined 3.2 mm, “d” in 2 pieces ?sex, 2.0 mm, “e”

juvenile 1.55 mm, “x” female 5.6 mm, “y” female 6.2

mm, “z” 4.6 mm female; sex of certain specimens not

determined to avoid further breakage, probably no

males present; lectotype measured to nearest 0.01 mm,

others to nearest 0.1 mm.

Diagnosis. (In same context as Laphystiopsis

spp.) Head scarcely longer than pereonites 1-2,

strongly bulging laterally (as seen from dorsal

view); pleonites 2-3 each with indistinct dorsal

bulge or carina; pleonite 4 shorter than pleonite 3;

coxa 5 45% as long as wide; coxa 6 bilobed and

not very wide anteroposteriorally (66% as wide as

coxa 5 and 162% as wide as coxa 7), 45% as long

as wide; epimeron 2 rounded posteroventrally,

corner rounded; article 5 of pereopods 3-4, 0.9

times as wide as long; article 2 of pereopod 7

narrowly pyriform, articles 2-7 together

[unknown, broken].

Description of lectotype female “a” 7.64 mm. See

illustrations. Head with apparent glandular tissue

in place of eye. Antenna | longer than antenna 2,

about as long as head plus pereonites 1—4

together, flagellum with 22 articles, callynophore

with 4 groups of 3-4 aesthetascs cach (not all

shown in figure), following articles 2-9 aes-

thetase formula = 4-3-2-2-1-1-1-0. Antenna 2

short, about 60% as long as antenna 2, flagellum

with 14 articles. Callynophore article without

ridges besides insertion points for rows of aes-

thetascs. Accessory flagellum very poorly devel-

oped. Ventral surface of article 1 on antenna | and

medioventral surfaces on articles 1-3 of antenna

2 with sparse ridges, stiff setules, weak scales.

One raker present on each mandible. Right lacinia

mobilis very transparent, with 2 teeth. Comb of

setae on article 3 of palp about as complex as

shown by Sars (1895) for Laphystiopsis plani-

frons and L. zomerysis. Pleopods 1—2 alike but

pleopod 3 slightly smaller, stick drawing of pleo-

pod 1 like pleopod 2, lengths of peduncle and

rami of uropod 3 shown, appearance of pleopods

like those drawn herein for one or more species of

Laphystiopsis; however, inner rami of pleopod 1

with 11 articles, outer with 13, inner of pleopods

2-3 with 13, outer with 15; coupling hooks 2 per

pleopod, no accessories. Most spines of uropods

1-2 and setae of uropod 3 missing and marked

with pits. Cuticle with saw-tooth ridges and

denticles similar to L. zomerysis above.

Juvenile “e” 1.55 mm. Smallest available; very

similar to adult, principal noticeable external dif-

ference: coxae 5 and 6 narrower, longer, posterior

lobes slightly longer than anterior lobes;

appendages much less spinose and setose, for

example, outer rami of uropods 1-3 with 4-2-4

spines only; flagellum of antenna 2 with 9

articles, antenna | broken.

Illustrations. Uropod 1 enlargement reduced to

75% of uropods 2-3-telson, Two views of lower

lip from female “x” are anterior and posterior, one

tilted, other flatter; upper and lower lips of lecto-

type damaged and not analyzed. Drawings of

mandibles purporting to show more than raker but

other projections apparently are scales or setules.

Distribution. South of West Falkland Islands; 179 m.

Prolaphystiopsis ornitorhynchus (Bulycheva)

comb. nov.

Bulycheva,

Laphystiopsis — ornitorhynchus

1952: 199-201, fig. 4.

Remarks. No body view of this species was given

in the original description so many elements of a

diagnosis cannot be calculated (sizes of coxae, for

example). I presume this species to be distinct

from P. platyceras but there is not much charac-

ter information to go on: the long setae on the

huge lobe of antenna 1; and slightly different

shapes and setation patterns on gnathopods, pere-

opod 7, mandibular palp and other mouthparts.

Distribution. Petra Velikogo (Peter the Great

Bay), Sea of Japan; 167-510 m.

Prolaphystius K.H. Barnard

Prolaphystius K.H. Barnard, 1930: 342.

Type species. Prolaphystius isopodops K.H.

Barnard, 1930 (monotypy).

Diagnosis. Rostrum absent, head not bulging.

Article 1 of antenna 1 not grossly lobate.

Mandibular molar columnar, scarcely triturative,

raker spine = 1. Inner lobes of lower lip absent.

Palp of maxilla | biarticulate. Coxae of ordinary

size, touching serially, coxa 4 about as long as

wide, excavate posteriorly. Pereopod 3 with elon-

gate carpus. Pleonite 3 dorsally flattened and

forming horizontal shelf projecting posteriorly;

urosomite | unmodified (young) or in adult

weakly saddled. Telson elongate, linguiform.

Description. Article | of antenna | thick, not car-

inate. Coxae not produced forward. Pereopod 4

distinct from pereopod 3. Pleonites 1—2 not cari-

nate.

Included species.

P. isopodops K.H. Barnard, 1930 [876B]

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 307

Figure 13. Prolaphystiopsis platyceras, lectotype female “a” 7.54 mm.

J. L. BARNARD

Figure 14. Prolaphystiopsis platyceras, unattributed figures = lectotype female “a” 7.54 mm;

“x” = female

“x” 5.6 mm.

REVISION OF LAPHYSTIOPSIDAE (AMPHIPODA) 309

Figure 15. Prolaphystiopsis platyceras, unattributed figures = lectotype female “a” 7.54 mm; “x” = female

“x” 5.6 mm. Note two views of lower lip.

310 J. L. BARNARD

Distribution. Marine, Antarctica, McMurdo

Sound; 406-441 m.

Acknowledgments

Thanks to Dr. F. M. Bayer for information about

the names of crinoids. I acknowledge the exten-

sive help given by Drs G.C. B. Poore, Jean Just

and Robin Wilson at Museum Victoria. Mrs

Elizabeth Harrison-Nelson at Smithsonian Insti-

tution assisted me in preparation of this work.

Mrs. Linda B. Lutz, of Vickburg, Mississippi,

inked the illustrations. Lennart Sandberg of

Naturhistoriska Riksmuseet, Stockholm, kindly

loaned the syntypes of Prolaphystiopsis

platyceras.

References

Barnard, J.L. and Karaman, G.S., 1991. The families

and genera of marine gammaridean Amphipoda

(except marine gammaroids). Parts 1 and 2.

Records of the Australian Museum, Supplement

13: 1-866. xxx.

Barnard, K.H., 1930. Amphipoda. British Antarctic

(“Terra Nova”) Expedition, 1910. Natural History

Reports, Zoology 8: 307- 454, 63 figures.

Bousfield, E.L., 1987. Amphipod parasites of fishes of

Canada. Canadian Bulletin of Fisheries and

Aquatic Sciences 217: 1-37, 10 figures.

Bulycheva, A.I, 1952, Novye vidy bokoplavov

(Amphipoda, Gammaridea) iz Japonskogo Morja.

Akademiia Nauk SSSR, Trudy Zoologicheskogo

Instituta 12: 195-250, 39 figures.

Gurjanova, E., 1951. Bokoplavy morej SSSR i sopre-

del’nykh vod (Amphipoda-Gammaridea).

Akademiia Nauk SSSR, Opredeliteli po Faune

SSSR 41: 1-1029, 705 figures.

Ledoyer, M., 1986. Crustacés amphipodes gammariens.

Faune de Madagascar 59(2): 599-1112, figures

227-415.

Norman, A.M., 1895. A month on the Trondhjem Fiord.

Annals and Magazine of Natural History (6) 15:

476-494.

Oldevig, H., 1933. Sveriges Amphipoder. Goteborgs

Kunglia Vetenskaps- och Vitterhets-Samhalles

Handlingar (B) 3 (4): 1-282 [all copied figures

after Sars, 1895].

Oldevig, H., 1959. Arctic, subarctic and Scandinavian

amphipods in the collections of the Swedish

Natural History Museum in Stockholm.

Goteborgs Kunglia Vetenskaps-Vitterhets-

Samhalles Handlingar (6B) 8 (2): 1-132, 4 plates.

Sars, G.O., 1895. Amphipoda. An account of the

Crustacea of Norway with short descriptions and

figures of all the species 1: i-vili, 1-711, 240

plates, 8 supplementary plates.

Schellenberg, A., 1931. Gammariden und Caprelliden

des Magellangebietes, Siidgeorgiens und der Wes-

tantarktis. Further Zoological Results of the

Swedish Antarctic Expedition 1901-1903 2 (6):

1-290, | plate, 136 figures.

Shoemaker, C.R., 1919. A new amphipod parasitic on a

crinoid. Proceedings of the Biological Society of

Washington 32: 245-246.

Stebbing, T.R.R., 1899, Revision of Amphipoda (con-

tinued). Annals and Magazine of Natural History

(7) 4: 205-211.

Stebbing, T.R.R., 1906. Amphipoda I. Gammaridea.

Das Tierreich 21: 1-806, 127 figures.

Stephensen, K., 1926. Revideret Fortegnelse over Dan-

marks arter af Amphipoda. 2. Del. (Gammaridea:

Fam. Stegocephalidae til Fam. Eusiridae). Viden-

skabelige Meddelelser fra Dansk Naturhistorisk

Forening 82: 43—101.

Stephensen, K., 1928. Storkrebs IL. Ringkrebs 1. Tan-

glopper (Amfipoder). Danmarks Fauna, Dansk

Naturhistorisk Forening: 1-399, 93 figures.

Stephensen, K., 1929, Amphipoda. Die Tierwelt der

Nord- und Ostee, Leipzig 14 (10, f): 1-188, 43 fig-

ures.

Stephensen, K., 1931. Crustacea Malacostraca. VII.

(Amphipoda. III.) Danish Ingolf-Expedition 3:

179-290, figures 54-81.

Stephensen, K., 1938. The Amphipoda of N. Norway

and Spitsbergen with adjacent waters. Tromso

Museums Skrifter 3:141-278, figures 20-31.

Thomas, J.D. 1996 Ecology and behavior of Maxillip-

ius commensalis, a gorgonophile amphipod from

Madang, Papua New Guinea (Crustacea:

Amphipoda: Maxillipiidae). Bulletin of Marine

Science, 58(1):314-326, figures 1—5.

Vader, W., 1978. Associations between amphipods and

echinoderms. Astarte 11: 123-134.

Memoirs of Museum Victoria 57(2): 311-317 (1999)

DESCRIPTION OF A NEW SPECIES OF THE PACIFIC SHRIMP GENUS PARACRANGON

(CRUSTACEA: DECAPODA: CRANGONIDAE) FROM SOUTHERN AUSTRALIA,

WITH A KEY TO THE GENUS

Yukio HANAMURA!, VICTORIA WADLEY2 AND JOANNE TAYLORS

‘National Research Institute of Fisheries and Environment of Inland Sea, Ohno-cho, Hiroshima 739-0452, Japan

(hanamura@nnf.affre.go.jp)

2CSIRO Marine Research, GPO Box 1538, Hobart, Tasmania 7001, Australia

(vicki. wadley@marine.csiro.au)

3Museum Victoria, 71 Victoria Crescent, Abbotsford, Victoria 3067, Australia

and Zoology Department, University of Melbourne, Parkville, Victoria 3052, Australia

(jtaylor@mov.vic.gov.au)

Abstract

Hanamura, Y., Wadley, V. and Taylor, J., 1999. Description of a new species of the Pacific

shrimp genus Paracrangon (Crustacea: Decapoda: Crangonidae) from southern Australia,

with a key to the genus. Memoirs of Museum Victoria 57: 311-317.

A sixth species of the genus Paracrangon, P. australis sp. noy. (Crustacea: Decapoda:

Crangonidae), is reported from a seamount off southern Australia and from Macquarie Island.

The Australian species is unique in having four—five teeth on the dorsal median margin of the

carapace, which is associated with a distinct reticulate structure on the supraventral part. All

species of the genus recorded to date have four or fewer teeth on the dorsal median margin of

the carapace. The present record is the first occurrence of the genus in the southwest Pacific

and greatly extends its known geographical range. A key for identification of all species is

presented with data on geographical and bathymetric ranges.

Introduction

The genus Paracrangon is a small group of cran-

gonid shrimps unique in having no second pere-

opods. It currently contains five species in the

Pacific Ocean, from Japan to off Peru, along the

North Pacific subarctic waters (Méndez, 1981;

Ohé and Takeda, 1986). Among the species,

P. echinata Dana, 1852 is a trans-North Pacific

species, occurring from the west coast of North

America to Japan and the Tsushima Strait in the

Sea of Japan, with a wide depth distribution from

the sublittoral to over 1000 m. The remaining

four species exhibit comparatively limited distri-

butions. Paracrangon abei Kubo, 1937 and P.

furcata Kubo, 1937 are endemic to Japan, while

P. areolata Faxon, 1893 has been recorded in the

eastern tropical Pacific from off Mexico to Peru.

In recent cruises (SS01/97, SS01/99) in south-

ern Australia and Macquarie Island of FRV

Southern Surveyor epibenthic sled surveys pro-

duced several shrimps including six specimens of

the genus Paracrangon. These specimens differ

from other species of Paracrangon in having

311

four-five dorsal median teeth on the carapace,

and undoubtedly belong to an undescribed

species.

This paper reports on this sixth species of

Paracrangon and provides evidence of an impor-

tant extension of the known geographical range

of the genus to the southwestern edge of the

Pacific. The specimens are deposited in collec-

tions of the Tasmanian Museum and Art Gallery,

Hobart (TM) and Museum Victoria, Melbourne

(NMV).

Paracrangon australis sp. nov.

Figures 1-3

Material examined. Holotype. Tasmania, approxi-

mately 84 km SSE of South East Cape (44°16’S,

147°20’E), 987 m, epibenthic sled, FRV Southern Sur-

veyor, 27 Jan 1997 (stn SS 01/97 36), TM G3656

(ovigerous female, 15.0 mm carapace length).

Paratypes. Tasmania, 84 km SSE off South East

Cape (44°16.2'S, 147°19.8’E), “J1” seamount, 1300 m,

epibenthic sled, T.N. Stranks et al. on FRV Southern

Surveyor, 27 Jan 1997 (stn SS01/97 37), NMV J41279

(2 ovigerous females, c. 15, 15.7 mm carapace length).

312 Y. HANAMURA, V. WADLEY AND J. TAYLOR

Vigure 1. Paracrangon australis sp. nov., holotype ovigerous female (cl. 15.0 mm), lateral view.

Other material, Macquarie Island, North end of Gap

(52°59.4" 53°2.0'S, 159°590 159°58.2'E), 1422 m,

Benthic Dredge, FRV Southern Surveyor, 31 Jan 1999

(stn 5801/99 130), TM G3756 (1 male 13.5 mm cara-

pace length). Macquarie Island, Beer Garden

(53055,9 53°54.9'S, 159°5,9'-159°2.2'E), 363.6 m,

Benthic Dredge, FRV Southern Surveyor, 26 Jan 1999

(stn §501/99 97), TM 64330 (1 male, 12.5 mm cara-

pace length, 1 ovigerous female, 20.0 mm carapace

length),

Diagnosis. Rostrum moderately long, directed

obliquely upwards, armed ventrally with 2 teeth

of normal shape, not furcate, Carapace with dor-

sal median margin bearing 5 teeth, and supraven-

tral carina forming irregular reticulate structure.

Description of holotype. Rostrum nearly straight

(broken off distally), extending obliquely

upwards, dorsal margin smooth, without tooth or

spine, ventral margin with strong tooth situated

just anterior to cornea and slightly smaller tooth

placed distally (Fig. 1).

Carapace with dorsal margin carinate for

almost entire length, armed with § teeth, size

varying considerably, first notably larger than

second, third tooth robust and fourth tooth sube-

qual to fifth; antennal tooth sharp, reaching

midlength of cornea; pterygostomian tooth larger

than antennal tooth; branchiostegal tooth set back

from anterolateral margin of carapace, strong,

flared anterolaterally; distinct carina supporting

branchiostegal spine extending backwards to

posterolateral margin (lateral carina), widely

reticulated posteriorly; 2 teeth decreasing in size

posteriorly arising along this carina; carina sup-

porting antennal tooth extending to near

midlength of carapace (dorsolateral carina), with

small tooth at posterior end; relatively weak

carina running between ventral margin and lateral

carina (supraventral carina), connecting with lat-

eral carina at both anterior and posterior ends,

somewhat reticulate and small tooth present near

posterior end; anterior vertical carina running

ventrad from base of third dorsal tooth and meet-

ing with dorsolateral carina; posterior vertical

carina weak, running ventrad from base of fifth

dorsal median tooth, slightly curving anteriorly

near ventral end (Figs 1, 2a).

Abdomen with somite 1 rounded or weakly

ridged dorsally, but not forming distinct carina;

somites 2-5 sharply carinate dorsally, with high-

est carina on somite 3; somite 6 1.78 times as long

as somite 5, 2 median dorsal carinae converging

into posterior end, ventrolateral margin with

small anterior tooth, and large, developed tooth

posteroventrally, posterolateral margin ending in

sharp tooth; ventral surface of somite 5 with pos-

teriorly curving sharp, long process near postero-

median part and somite 6 with pair of anteriorly

directed sharp processes at anterior end of ventral

surface; pleura of somites 1-5 acutely produced

ventrally, increasing in size towards posterior,

accompanying median pleural carina supporting

ventral spine or process, without additional tooth

or spine on anterior margin (Figs 1, 2b). Telson

broken off distally, but at least 2 pairs of dorso-

lateral spines present. Exopod of uropod shorter

A NEW SPECIES OF THE SHRIMP PARACRANGON FROM SOUTHERN AUSTRALIA 313

ded

be mm,

Cai mm a

2mm

5mm

f 0.5 mm

Figure 2. Paracrangon australis sp. nov., holotype ovigerous female (cl. 15.0 mm). a, carapace and eye, dor-

sal view. b, posterior part of body, dorsal view. c, antennule. d, antenna. e, pereopod 1. f, vestigial pereopod

2. g, pereopod 3. h, pereopod 4. i, pereopod 5.

314 Y. HANAMURA, V. WADLEY AND J. TAYLOR

ef 1 mm

an õm , b-d — 2mm,

Figure 3. Paracrangon australis sp. nov., holotype ovigerous female (cl. 15.0 mm). a, maxilliped 3.

b, maxilliped 2. c, maxilliped 1. d, maxilla 2. e, maxilla 1. f, mandible (dorsal view in right figure).

A NEW SPECIES OF THE SHRIMP PARACRANGON FROM SOUTHERN AUSTRALIA 315

than endopod, with roundly produced distolateral

lobe (Fig. 2b).

Eye with cornea well pigmented, slightly wider

than eye-stalk (Fig. 2a),

Antennular peduncle relatively slender, with

first segment 1.44 times as long as second, latter

2.81 times as long as third; upper flagellum short,

composed of 19 articles, flattened towards dis-

tally, lower flagellum with 9 articles; stylocerite

very short, distally rounded (Fig. 2c),

Antennal scale 0.53 times as long as carapace,

242 times as long as wide, distolateral spine

falling short of end of lamella; carpocerite long,

extending well beyond end of lamella (Fig. 2d).

Mouthparts as illustrated (Fig. 3b-f). Maxil-

liped 3 extending beyond end of antennular

peduncle by length of whole distal segment, latter

1.36 times as long as penultimate (Fig. 3a).

Pereopod 1 subchelate, extending beyond end

of antennular peduncle by length of distal third of

propodus and dactylus, movable finger sharp,

curving inward, fixed finger sharp (Fig. 2e). Pere-

opod 2 vestigial (Fig. 2f). Pereopod 3 slender,

extending as far as end of pereopod 1, dactylus

about third length of propodus, with long, sharp

terminal seta (Fig. 2g). Pereopod 4 extending

beyond end of antennular peduncle by about dis-

tal half length of propodus and dactylus; propodus

with 8 long ventral spines; dactylus sharp, curv-

ing posteriorly, quarter—fifth length of propodus

(Fig. 2h), Pereopod 5 extending beyond end of

antennular peduncle by about distal third length

of propodus and dactylus; propodus with 12-13

ventral spines, including 3 close-set distal spines;

dactylus slightly less than fifth length of propodus

(Fig. 2i).

Note on paratyes. The paratypes are more or less

damaged in the carapace so the number of teeth

on the dorsal median margin could not be counted

accurately. The dorsal margin of abdominal

somite 1 is more weakly ridged in the paratypes

than in the holotype but not forming an acute

carina as in following somites. Other external

features including the carinal structure and

ornamentation agree well with those of the

holotype.

Colour in fresh condition. The background colour

is basically light-red, with a slightly darker red on

the rostrum and the posterior part of the abdomen.

Egg size. Non-eyed eggs are nearly spherical,

moderately large, diameter 1.8—1.9 mm, and

eyed eggs are oval, approximately 2.5 mm along

longer axis.

Distribution. The type ovigerous females were

collected from 987—1300 m depth approximately

84 km SSE off South East Cape, Tasmania. The

more recently discovered material from

Macquarie Island was at 363-1422 m. The char-

acteristic red colour suggests this species to be a

typical deepwater inhabitant.

Etymology. The specific name “australis” (=

southern in Latin) indicates that the species is the

southernmost inhabitant of the genus.

Remarks. Unlike the five described species of the

genus Paracrangon, P. australis is unusual in

possessing four-five dorsal median teeth on the

carapace; all others have four or fewer. An irreg-

ularly reticulated structure of the supraventral part

of the carapace noted in P. australis is similar to

that in P. areolata and P. okutanii, This structure

is not found in the remaining three species.

In addition to the carapace spine counts of the

dorsal median margin, P. australis differs from P.

areolata in having relatively shorter dactyli of the

posterior two pereopods (fifth—sixth length of

propodus vs third), and from P. okutanii by hav-

ing proportionately shorter, obtuse ventral projec-

tions of the first two pleura, and the rostrum being

directed more upward (50° vs 35°).

Although the disposition and number of teeth

on the carapace display some intraspecific varia-

tions (Brashnikov, 1907; Hayashi, 1986), they

have a specific pattern (Table 1). This may simply

be due to lack of basic information about this fea-

ture partly reflecting the rarity of the species and

future study may alter this table. In addition, an

exact description of carinal sturucture of the cara-

pace will be useful for definite identification of

each species.

A microscopic chitinous lobe between the first

and third perepods is considered a second pere-

pod. Confirmation of this is needed in other

species.

Discussion

Paracrangon is considered primarily a North

Pacific genus with highest species richness in the

waters around Japan, four of the five hitherto

known species having been recorded there (Fig.

4). Prior to our finding, Paracrangon areolata

was thought to be the only species to occur in the

southern hemisphere. It occurs in the tropical east-

ern Pacific as far south as 17°S (Méndez, 1981;

Hendrickx, 1995). The finding of a sixth species

in southern Australian and Subantartic waters is a

significant extension to the known geographical

316 Y. HANAMURA, V. WADLEY AND J. TAYLOR

Table 1. Disposition and number of teeth on carapace of species of Paracrangon

Species dorsal median dorsolateral part lateral part mainly ventrolateral part

margin btween median along lateral between lateral

and lateral carinae carina including carina and ventral

branchiostegal margin

and hepatic teeth

P. abei 3 2

P. areolata 4 2

P. australis 4-5 |

P. echinata 4 2-6

P. furcata 2-3 0

P. okutanii 4 1

4 0

3 l

3-5 2

2 0

3 l

range of the genus to 54°S. It is curious that the

genus has not so far been found in the tropical

western Pacific despite well organized intensive

deepwater studies in the region over a hundred

years.

Paracrangon is a well established crangonid

genus and identification can be made using the

following key. The key is modified from those

provided by Kubo (1937) and Ohé and Takeda

(1986), with a view to providing more con-

servative characters than the rostrum. The geo-

graphical and bathymetric ranges are included.

Key to species of Paracrangon

Supraventral carina of carapace forming distinct irregular reticulate structure

EE eN elles dh ace 2

Supraventral carina of carapace not forming distinct irregular reticulate

BUUOWIES nn seinen orden aan oetan Bleser une teas iay cag reei genera 4

Dactyli of posterior 2 pereopods about third length of propodi..... P. areolata

Faxon, 1893 (eastern tropical Pacific, from off Mexico to Peru; 650-1250 m)

Dactyli of posterior 2 pereopods about fifth—sixth length of propodi........... 3

Rostrum shorter than carapace, obtuse ventral projection of first two pleura

equal to or greater than depth of carapace „n...

rard P. australis sp. nov. (southern Australia, Macquarie Island; 360-1422 m)

Rostrum longer than carapace; obtuse ventral projection of first two pleura

shorter than depth ofcarapacen. mnd boren ERE NA

P. okutanii Ohé and Takeda, 1986 (Central Pacific coasts of Japan; 425-1205 m)

Dorsal median margin of carapace with 4 teeth; all abdominal somites cari-

nated dorsally ......... P. echinata Dana, 1852 (California to central Japan and

Sea of Japan, throughout North Pacific boreal waters; sublittoral to 1380 m)

Dorsal median margin of carapace with 2 or 3 teeth; abdominal somite 1 at

leastetounded doreen bt Huan heersten eeen omer meee mtn tian 5

First 2 median dorsal teeth of carapace denticulate distally; dorsal margin of

carapace with 2 teeth and basal ventral spine simple, not furcate; abdominal

somite 2 carinate dorsally … … ……… … … P. abei Kubo, 1937 (Japan, central

to SW Pacific coasts, along Tushima Current in Sea of Japan; 150-300 m)

Median dorsal teeth of carapace, simply tapering distally; dorsal margin of

rostrum smooth and basal ventral spine furcate; abdominal somite 2 rounded

OOTSAIN srt teven dures pdm er

yt, P. furcata Kubo, 1937 (Japan, central to SW Pacific coasts; 320—400 m)

A NEW SPECIES OF THE SHRIMP PARACRANGON FROM SOUTHERN AUSTRALIA 317

Acknowledgements

We are grateful to Dr M. Hendrickx, Mazatlan

Station, Universidad Nacional Autonoma de

México, Mexico, for morphological information

about the eastern Pacific species. One of authors

(YH) thanks Drs K. Hayashi, National Fisheries

University, Shimonoseki and T. Komai, Natural

History Museum and Institute, Chiba, Japan for

expert comments on terms and morphology. This

study is supported in part by grants from the Aus-

tralian Industry Research and Development Cor-

poration Grant (95/058), Environment Australia

(Australia Nature Conservation Agency), and

the Science and Technology Agency of Japan

(1996).

References

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v P. areolata

X P. australis

@ P. echinata

m P. furcata

@ P. okutanil

CONTENTS

Systematics and biology of Macquarie Island echinoderms

T. DOCG ee rn ET Rn

New species of the water mite genus Arrenurus from eastern Australia (Acari: Hydrachnidia:

Arrenuridae)

H, Smit sss ABe Tan rr) tole ee ed NR

A second species of Atriplectides Mosely from Australia (Trichoptera: Atriplectididae)

A. Neboiss sunt Eman eten te et EO a

A new genus of subcortical coccoids (Hemiptera: Coccoidea: Eriococcidae) on Eucalyptus

P J. Gullan re er oant wii MK EN

A new species of Ogyris Westwood (Lepidoptera: Lycaenidae) from southern inland Australia

R. Field. … saber etn A en ee ee

Replacement names for five Australian species of Lasioglossum (Chilalictus) (Hymenoptera:

Halictidae)

KL. Walkers Teyi NE TT Bnr Et EE

Designation of type species for the genera of Australian paropsine beetles (Coleoptera:

Chrysomelidae)

PG Kelly and C. A.M, Reid … noten vende terr Be ne rs on On ER

Storthyngurella, new genus of Munnopsidae (Crustacea: Isopoda), with descriptions of three

new species from deep-sea basins of the Southern Hemisphere

M. V Malyutitid: wrote coen Bro erneer Beene oe er es et ea en Nn

Revision of Laphystiopsidae (Crustacea: Amphipoda): new and old species from South

China Sea, southeastern Australia, Falkland Islands and western Atlantic Ocean

J L. Barnard: zit seacte gb I tg EE RO ORAS, ie ere et ear EE

Description of a new species of the Pacific shrimp genus Paracrangon (Crustacea: Decapoda:

Crangonidae) from southern Australia, with a key to the genus

Y Hanamura, -V Wadley. and J Taylor’ es secr cs ae Sen ca Aes Re Pree eee