MEMOIRES

DU MUSEUM

NATIONAL

D’HISTOIRE

NATURELLE

Result at s des Campagnes MUSORSTOM

Volume 12

Coordonne par

Alain CROSN1ER

1 3 PEC, 199 4

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resultats des campagnes

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ISSN : 1243-4442

© Editions du Musdum national d'Histoire naturelle, Paris, 1994

MEMOIRES DU MUSEUM NATIONAL D’HISTOIRE NATURELLE

TOME 161

ZOOLOGIE

EDITIONS

DU MUSEUM

PARIS

1994

Source : MNHN, Paris

SOMMAIRE

1.

Pages

La campagne CALSUBsur les pentes bathyales de la Nouvelle-Caledonie (en anglais) . 9

Michel Roux

2.

3.

4.

5.

6.

7.

8.

9.

10.

li.

12.

Protozoa, Xenophyophorea Granuloreticulosa : Psammina zonaria sp. nov. du Pacifique ouest.

Considerations sur quelques aspects de la croissance des xenophyophores (en anglais) .

Ole Secher Tendai.

Bryozoa : Especes vivantes et fossiles des sous-familles catenicellides : Ditaxiporinae Stach et

Vasignyellinae nov. (en anglais) .

Dennis P. Gordon & Giampetro Braga

Cirripedia Thoracica : Verrucomorpha de Nouvelle-Caledonie, d'Indonesie et des iles Wallis

et Futuna (en anglais) .

John S. BUCKERIDGE

Crustacea Amphipoda : Lysianassoides du Sud-Ouest Pacifique tropical (en anglais)

James K. Lowry & Helen E. Stoddart

Crustacea Isopoda : Bopyridae des collections Musorstom recoltes dans l'lndo-Pacifique

tropical. I. Sous-familles Pseudioninae (en partie), Argeiinae, Orbioninae, Athelginae et

Entophilinae (en anglais) .

John C. Markham

Crustacea Decapoda : Les Metapenaeopsis indo-ouest-pacifiques avec un appareil stridulant

(Penaeidae) .

Alain CROSNIER

Crustacea Decapoda : Observations complementaires sur les Metapenaeopsis indo-ouest-

pacifiques sans appareil stridulant (Penaeidae). Description de deux especes nouvelles .

Alain CROSNIER

Crustacea Decapoda : Penaeoidea recoltes lors de la campagne KARUBAR en Indonesie .

Alain CROSNIER

Crustacea Decapoda : Penaeoidea, a ('exclusion des Sicyoniidae, recoltes dans la zone

economique des iles Wallis et Futuna, lors de la campagne MUSORSTOM 7 .

Alain CROSNIER

Crustacea Decapoda : Pagures d'eau profonde (Parapaguridae) de la Polynesie frangaise.

Description de quatre especes nouvelles (en anglais) .

Rafael LEMAITRE

Crustacea Decapoda : Le genre Munida Leach, 1820 (Galatheidae) dans les eaux

neo-caledoniennes et avoisinantes. Description de 56 especes nouvelles (en anglais) .

Enrique MACPHERSON

. 49

. 55

87

127

225

255

339

351

367

375

421

CONTENTS

Pages

1. The Calsub cruise on the bathyal slopes off New Caledonia . 9

Michel Roux

2. Protozoa, Xenophyophorea Granuloreticulosa : Psammina zonaria sp. nov. from

the West Pacific and some aspects of the growth of xenophyophore . 49

Ole Sccher TENDAL

3. Bryozoa : Living and fossil species of the catenicellid subfamilies Ditaxiporinae

Stach and Vasignyellinae nov . 55

Dennis P. Gordon & Giampetro Braga

4. Cirripedia Thoracica : Verrucomorpha of New Caledonia, Indonesia, Wallis

and Futuna Islands . 87

John S. BUCKERIDGE

5. Crustacea Amphipoda : Lysianassoids from the tropical western South Pacific

Ocean . 127

James K. Lowry & Helen E. Stoddart

6. Crustacea Isopoda : Bopyridae in the Musorstom collections from the tropical

Indo-Pacific. I. Subfamilies Pseudioninae (in part), Argeiinae, Orbioninae,

Athelginae and Entophilinae . 225

John C. Markham

7. Crustacea Decapoda : The Indo-West Pacific species of Metapenaeopsis with

stridulating organs (Penaeidae) (in french) . 255

Alain CROSNIER

8. Crustacea Decapoda : Complementary observations on the Indo-West Pacific

species of Metapenaeopsis without stridulating organs (Penaeidae) with

description of two new species (in french) . 339

Alain CROSNIER

9. Crustacea Decapoda : Penaeoidea collected in Indonesia during the Karubar

cruise (in french) . 351

Alain CROSNIER

10. Crustacea Decapoda : Penaeoidea, Sicyoniidae excluded, collected in the

economic zone of the Wallis and Futuna Islands during the Musorstom 7 cruise

(in french) . 367

Alain CROSNIER

11. Crustacea Decapoda : Deep-water Hermit crabs (Parapaguridae) from French

Polynesia with descriptions of four new species . 375

Rafael Lemaitre

12. Crustacea Decapoda : Studies on the genus Munida Leach, 1820 (Galatheidae)

in New Caledonia and adjacent waters with description of 56 new species . 421

Enrique Macpherson

RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES <

The CALSUB cruise

off New

on the bathyal slopes

Caledonia

Michel ROUX

Laboratoire des Sciences de la Terre et URA 157 du CNRS

Universite de Reims Champagne-Ardenne

B.P. 347 - F 51062 Reims Cedex

ABSTRACT

The CaLSUB cruise took place in 1989 off the coast of New Caledonia. The diving saucer Cyana enabled the

exploration of the bathyal environment from the edge of the coral platform down to a depth of 3000 m. In this paper an

account is given of how the study was carried out, along with the material gathered during the course of the study which

may be consulted (in particular the photographs and the videos). The various sectors explored, and the principal

observations made are discussed. Apart from the wide diversity of ecological niches observed, what makes this sector of

the Pacific Ocean original, is the existence of bathyal benthic communities rich in "living fossils", giving us an idea of

the fauna which populated the Tethys Ocean during the Mesozoic. The use of the Cyana diving saucer allowed direct

observation of the nature of the sea beds and the associated fauna, in particular of rocky slabs, and environments affected

by deep currents, where fields of hydraulic bioclastic dunes develop along fault scarps covered with dense populations

dominated by Echinoderms. We were able to establish the bathymetrical distribution of the bathyal populations.

RESUME

La campagne CaLSUB sur les pentes bathyales de la Nouvelle-Caledonie.

La campagne CALSUB s'est deroulee en 1989. au large de la Nouvelle-Caledonie. Elle a permis une exploration, a l'aide

de la soucoupe plongeante Cyana, de l'environnement bathyal, depths la bordure de la plate-forme corallienne jusqu'a pres

de 3000 m de profondeur. On presente ici le deroulement de la campagne, les documents recueillis susccptibles d'etre

consultes (notamment les photographies et les videos), les differents secteurs explores, et les principalcs observations

effectuees. Outre la grande diversite des niches ecologiques observees, l'originalite de ce secteur de l'ocean Pacifique reside

dans la presence de communautes benthiques bathyales, riches en "fossiles vivants" et evoquant celles qui peuplaient la

Tdthys au cours du Mesozoique. L'utilisation de la soucoupe Cyana a donnd acces a l'observation directe des fonds et des

associations fauniques, et particulierement, d'environnements sur dalles rocheuses et de ceux parcourus par des courants

profonds ou se developpent des champs de dunes hydrauliques bioclastiques le long d'escarpements de faille couverts de

peuplements tres denses domines par les echinodermes. Letagemcnt bathymetrique des peuplements bathyaux a pu etre

precise.

ROUX, M., 1994. — The CALSUB cruise on the bathyal slopes off New Caledonia. In : A. CROSNIER (ed.), Resultats des

Campagnes MUSORSTOM, Volume 12. Mem. Mas. natn. Hist, nat., 161 : 9-47. Paris ISBN 2-85653-212-8.

10

M. ROUX

INTRODUCTION

The exploration of the deep sea fauna of New Caledonia began with test cruises from 1977 to 1979 on beds

ranging in depth from 200 m to 1000 m off the coast of Grande Terre, off the Isle of Pines and off the Loyalty

Islands (Int£s, 1978). As of 1985, the scope of the exploration was widened and intensified within the framework

of the Musorstom cruise program (Richer df. Forges, 1990) and the Envimarges program (Roux, 1991),

which enabled sedimentologists, paleontologists and biologists to collaborate.

As the Musorstom cruises developed, paleontologists began to play an important role in the study of the

material gathered. The "living fossils" theme, at the forefront since Musorstom 1, led by J. Forest (1981),

which set out to find new examples of the only living representative of the glypheid crustaceans, Neoglyphea

inopinata, was reinforced when the MUSORSTOM cruises moved from the Philippines to New Caledonia. With the

Envimarges program, the widening of interdisciplinary cooperation was aimed at improving understanding of the

ecological and geodynamic factors which, in the course of geological history, have enabled the survival of ancestral

forms amongst the current bathyal fauna of the Western Pacific and. in particular, the fauna of New Caledonia.

The Envimarges program, overseen by the INSU-CNRS with the support of TOTAL-CFP, concentrated

primarily on a transect perpendicular to the axis of the basin of the Loyalty Islands from Thio to the west and the

north-west of Lifou. Two complementary sectors were also studied : the slope situated to the south of Noumda and

the slope situated between the Isle of Pines and the large southern reef, reasonably regularly inclined towards the

south-east. This program was carried out by three separate cruises : Biocal. Biogecal and Calsub.

Fig. 1. — Recovery of SP 3000 Cyana at the end of a dive during the CALSUB cruise.

FIG. 1 . — La SP 3000 Cyana recuperee a la fin d'une plongee de la campagne CaLSUB.

In 1985, during the Biocal cruise on board the R/V Jean Charcot (mission led by Claude Lfivi), five sectors of

the bathyal slope (SB 1 to 5) were mapped using the multibeam sounder. Sea Beam. Samples of sediment (using

the Kullenberg and Usnel coring systems) and fauna (using beam trawls and Warren's epibenthic dragnet) were also

taken. In 1987, the BIOGECAL cruise, on board the R/V Coriolis (mission led by Pierre COTILLON) enabled the

CALSUB CRUISE OFF NEW CALEDONIA

1 1

completion of the sedimentary and biological sampling of the transect on flat ground and moderate inclines.

Finally, in 1989, the Calsub cruise, on board the R/V Suroit (mission led by Michel Roux) used the Cyana

diving saucer (fig. 1) to explore the steeper and rockier beds. Moreover, a liaison with the Proppac program,

conducted by ORSTOM (Study of the effects of climatic variation on pelagic production in the high seas of the

SW Pacific) allowed us to obtain samples of plankton in order to study the current nannoflora between 20°S and

10°N by 165°E, and thereby to compare this to the fossilised calcareous nannoplankton in the Quaternary

sediments of the Loyalty basin.

The principal results of the Envimarges program, with special reference to the Earth Sciences, have been

compiled in a special, illustrated report (Lambert & Roux, co-ordinators, 1991). which biologists will be able to

consult usefully.

THE CALSUB CRUISE AND RESULTING DOCUMENTATION

1. - Summary of how the cruise was carried out

The Calsub cruise took place between 18 February and 13 March 1989 and consisted of two legs. Twenty-two

dives were made (fig. 2) from the outer edge of the platforms to the abyssal zone at a depth of around 3000 m

(fig. 3; table 1).

12

M. ROUX

FlG. 3. — Bathymetric range of dives into different areas investigated.

Fig. 3. — Elagement bathymetrique des plongees dans les differents secteurs explores.

Four areas had been chosen : (1) the region to the NE of Lifou. including the area shown on the Sea Beam map

SB3; (2) Sea Beam map SB 1 off the coast of Thio; (3) the basin which runs between the Isle of Pines and the

Great Southern Reef; (4) Stylaster Bank, a guyot situated to the SE of the Isle of Pines. As for the areas which had

not been mapped during the Biocal cruise, we were able to benefit from readings taken by the Marine

Hydrographic Service of Noumea in Santal Bay (to the W of Lifou) and on the SE edge of the Isle of Pines.

The initial dive program was changed during the course of the cruise because of the rough condition of the sea

which often only allowed us to dive in sheltered sectors. Santa! Bay and the Ouvea-Lifou rise (10 dives), as well as

the basin situated to the SW of the Isle of Pines (5 dives), were explored more intensively than initially intended,

whereas the number of much deeper targets was reduced (SB 1 and SB 3), and the plan to dive on the Stylaster

Bank had to be abandoned. Thanks to the excellent cooperation between the crew of the Suroit and the technical

team of the Cyana saucer, we were able on a number of occasions to make two dives in a single day to targets of a

moderate depth.

The scientific team of each leg was composed of 6 members of the Ecoprophyce Research Group of the CNRS

and one guest from the ORSTOM centre in Noumea. Michel Roux (Earth Sciences Laboratory, University of

Reims), who led the cruise, was involved in both legs. The first leg, from 18 February until 1 March, brought

together Jean-Paul Bourseau and Michel Rio (Centre for Earth Sciences, University of Lyon), Rene

Grandperrin (ORSTOM, Noumea). Bernard Laurin (Centre for Earth Sciences, University of Dijon), Claude

Monniot (National Natural History Museum, Department of Invertebrate Marine Biology, Paris) and Jean

Vacelet (CNRS, Marine Station at Endoumc, Marseille). The second leg brought together Philippe Bouchet

and Alain GuiLLE (National Natural History Museum, Department of Invertebrate Marine Biology, Paris),

Christian GAILLARD (Centre for Earth Sciences, University of Lyon), Bertrand RICHER DE FORGES (ORSTOM,

Noumea), Michel Segonzac (IFREMER, Brest) and Helmut Zibrowius (CNRS. Marine Station at Endoume,

Marseille).

Each dive produced the following documentation : (1) a complete video recording on VHS tape; (2) several

hundred photographic slides; (3) a detailed report written by the diver once back on board. Umatic video sequences,

which are of very high quality, were only occasionally taken; they have been converted to VHS, thereby making

them easier to consult.

CALSUB CRUISE OFF NEW CALEDONIA

13

The sampling capacity of the Cyana saucer is limited, particularly when working on steep slopes. This was

improved by using autonomous shuttles (lifts) and ballasted sampling cans carried aboard the saucer. The big draw-

lift was only used on the first dives at the deepest site (SB 3). Its ability to carry baskets of dimensions 60 cm x

40 cm x 30 cm enabled us to collect complete specimens of some size (sponges, stalked crinoids). A double-box

lift, which is more maniable, was more frequently used.

In general, this lift was placed halfway along the route : we placed in it a ballasted can full of samples taken

during the first part of the dive, substituted by an empty can used during the second part. Moreover, one of the

boxes is capable of keeping large specimens collected nearby. When the lift could be cast off in the evening,

around 10 or 12 hours before a dive, bait (tuna, chicken) was attached to the ballast and protected from sharks by

wide-meshed wire netting. It was in this way that nautilids and various crustaceans could easily be observed and, in

some cases, even captured.

Since the suction system did not function properly, we were unable to collect small macrofauna, despite its

abundance on certain epibathyal beds.

On board, a few samples were prepared and conserved in liquid nitrogen, with a view to performing comparative

phylogenetic analyses, using information provided by ribonucleic acid (RNA) sequencing.

Table 1. — Cyana dives during the Calsub cruise.

Tableau 1. — Plongees de Cyana lors de la campagne CALSUB.

Source

14

M. ROUX

2. - Documentation available for consultation

The video and photographic documents, along with the reports of the dives can be consulted at the Earth

Sciences Laboratory at the University of Reims. The biological specimens have been listed and sent out to

specialists by the CENTOB (1FREMER - Brest). The samples of sediment or rock have been listed and sent out

for study by the Centre for Earth Sciences at the University of Lyon. Copies of slides and a video of the cruise

(produced by FR3 Champagne-Ardenne and available only for strictly scientific or pedagogic purposes) are

available by request to the head of the mission.

The data chamber which appears on the slides (fig. 4) and video films, indicates the depth, bearing, time and

numerical order of the photograph (or. on the videos, the number of the dive). For the lesser depths, the

bathymetric readings tend to be overestimated and must be corrected by reckoning that from 500 m up to the

surface, the overestimation of the depth varies progressively from 0 to 30 m.

depth (1850m)

/

IIH

a 3

<— photo number (527)

<-— hours (12h)

<— minutes (5 5mn)

<--- seconds (13s)

/

course (065)

Fig. 4. — Data chamber printed on the lower left side of pictures taken by Cyana during Calsub cruise.

Fig. 4. — Chambre de donnees apparaissant en bas et a gauche sur les photographies prises par Cyana lors de la campagne

Calsub.

The reports of the dives include :

- detailed maps of the route taken and its position in the study zone (for example, figs 5, 6 and 7);

- diagrams indicating the main substrates and environments observed;

- a list of the useful photographs, with a brief commentary and sometimes preliminary names for organisms;

- a selection of the dive's more striking photographs:

- the list of the samples taken with the corresponding video sequences and photographs;

- a summary of the dive, written following the course of the VHS tape, with a temporal guide from the data

chamber; here the observations and interpretations have the advantage of having been written 'on the spur of the

moment- by the observer;

- complementary information provided, once on board, by the members of the scientific team, or after the

mission by specialists who had examined the documents.

Video and photographic documentation gathered during cruises such as this, is often underused. This is

probably partly due to the fact that not enough information is available as to its nature and content, and partly to

its mere bulk, which means that a thorough preliminary study is necessary before the required information can be

found. We have attempted to facilitate the work of specialists interested by the extremely diverse documentation

gathered by the Calsub mission and hope that it will be frequently consulted.

3. - The different sectors explored

3-1. - Around the island of Lifou

The Loyalty Islands, ot which Lifou is one of the three largest, are ancient atolls, raised up by the lithospheric

bulge which affects the Pacific Plate, before its subduction at the Vanuatu Trench. There is no platform at the

Source

CALSUB CRUISE OFF NEW CALEDONIA

15

island's edge, and very rapidly we reach depths in the thousands of meters. The three sectors explored off the coast

of Lifou (fig. 5) correspond to ( 1) a sheltered bay. (2) a rise and (3) the flank of an ancient underwater volcano.

3-1-1. - In Santal Bay

Santal Bay is situated on the western coast of Lifou. A narrow projecting edge supports coral pinnacles. The

bed of the bay is deep, around a thousand meters, and is reached by a moderate to steep ramp covered in coral sands.

By the headlands which enclose the bay to the north and south, the slope is much steeper, even abrupt in places,

and rocky beds are predominant. The morphology of this bay might indicate that it is the scar of a collapse which

affected Lifou as it rose.

To the north, near cap Aime, four dives explored the talus between 1 150 m and 130 m. The first two (dives 5

and 6) were carried out in the mouth of the bay. in an area swept by currents, and the following two (dives 7 and 8)

on the side of a more sheltered spur.

To the south, near the mouth of the bay at cap Lcfevrc, three dives (9 to 1 1), at between 600 m and 700 m,

studied the external flanks, often abrupt, of a small coral group, the Shelter reef (fig. 6).

Coral or algal material, and even blocks of rock loosened by erosion, are in constant movement on these steep

epibathyal slopes (Rio el al. . 1991).

The benthic fauna only develops beyond areas frequently affected by these sedimentary movements. On the

whole, the density of benthic fauna observed in Santal Bay was rather low.

FlG. 5. — Location of dives conducted off Lifou island.

FlG. 5. — Localisation des plongees mcnees au large de file de Lifou.

16

M. ROUX

Santal Bay.

Fig. 6. — Carte avec la plongee 9 effectuee sur le flanc NE du recif Shelter. Ce type de document existe pour chacune des

plongees en baie du Santal.

3-1-2. - On the Ouvtia-Lifou rise

Dives 14. 15 and 16 explored, between 900 m and 400 m, the flanks of the Ouvea-Lifou rise, just to the west

of the Jouan reef, a small coral outcrop. At around 500 m. the first two dives discovered a very peculiar submarine

morphology, probably linked to ancient hydrothermal activity (VANNEY, 1991; VANNEY et al„ 1992), containing

indurated ferromangamferous rock, dissected by erosion into "mushroom rocks". The other hard ground beds are,

tor the main part, large calcareous surfaces, often suffering biocorrosion or covered in a thin film of pteropod

shells. The flanks ol the rise are affected by faults which form the limit of the sedimentary channels swept by

currents and which are bordered by subvertical walls (even overhangs in some places). Dive 16, near to Jouan reef,

observed substantial quantities of rocks and algal gravel, at times covering the entire bed for around 400 m or

500 m, limiting the benthos to a predominantly vagile fauna. Dive 15, to an area more sheltered from the effects

ol nentic sediments, observed an abundant fauna attached to the hardened beds.

3-1-3. - In the deeper zone to the North of Lifou

The Sea Beam map SB3 (fig. 7) allowed us to accurately plan four dives (1 to 4), between 1100 m and 2900 m

in depth, on the flank of a volcanic seamount with a summit at around 1000 m deep, and slightly detached from

Source :

CALSUB CRUISE OFF NEW CALEDONIA

17

the talus situated to the north of Lifou. Here, rocky slabs or rather coarse volcanic breccia are predominant. In

those areas where the slope is less steep and on the few flattish ledges, accumulations of sediment, are seen to

develop.

Fig. 7. — Location of dives 1 to 4 off the North of Lifou island. Map without extrapolation, from multibeam sounder of

R/V J. Charcot.

Fig. 7. — Localisation des plongees 1 a 4 au Nord de Lifou. Carte sans extrapolation, levee avec le Sea-Beam du

N/O J. Charcot.

These are the dives which best demonstrate the New Caledonian abyssal benthos on hard substratum. Dive 2

(fig. 8) provides us with the clearest picture of the fauna and of its distribution. However, the navigational

difficulties we faced with the saucer during dive 3 (the deepest) meant we were unable to localise the route it took,

which involved, in part, the sedimentary beds at the foot of the talus.

3-2. - Off the coast of Thio

This is a mcsobathyal area, dominated by sedimentary beds in the process of being eroded, and therefore

generally much less stable than those in the SB3 area. Weather conditions only allowed us to carry out three dives

(12, 13 and 17) on the bathyal talus. It was not possible to explore the fringe of the reef immediately below the

barrier reef situated off the coast of Thio. Sea Beam map SB 1 (fig. 9) shows a series of trenches and edges parallel

to the slope. The dives were performed in stages along the same edge in each case. It would appear that this

morphology is dominated by the effects of regressive erosion starting from the base of the talus, since the trenches

become less marked towards the top of the talus. This would primarily come about as the result of erosion

processes in a sedimentary material that has undergone little or no lithificaton.

We may consider then that the fauna observed is able to adapt to an unstable bed where erosion is active on

most surfaces, with a substratum ranging from crumbly to compact, containing slightly more resistant.

18

M. ROUX

ferromanganiferous layers. We only observed more indurated sites on dive 12 near the cpibathyal region. Ancient

or indeed current bioturbation affects the compact sediments as much as the crumbly sediments; we were able to

analyse this in greater detail during dive 17 (GAILLARD, 1991 a and b). The second half of dive 12, which included

more indurated substrates, merits a comparison with dives 1 and 2 of SB3.

Fig. 8. — Map of the area of dive 2 with the different substrates observed on the north slope of Lifou.

Fig. 8. — Carte dc la zone parcourue par la plongee 2 avec les principaux types de fonds observes.

3-3. - Between the Isle of Pines and the Great Southern Reef

Five dives (18 to 22) allowed us to describe the environment of this epibathyal gulley (fig. 10).

Dives 18 and 19 examined the slope which links the outer edge of the coral platform with the Isle of Pines

between 60 m and 450 m deep. There is no barrier reef in this place, and the carbonate ooze which accumulates at

the edge of the platform leads to mass slides at those areas where the slope becomes steeper.

The ramp which slopes towards the SE and which forms the bottom of the gulley is periodically swept by

substantial currents, which create bioclastic hydraulic dunes in areas of sedimentary accumulation, and types of

"reg" covered by rocky particles which are a result of the loosening of indurated ferruginous crusts in the non¬

deposit zones (Rio el al„ 1991).

Dive 21, at around 350 m depth, showed some relatively stabilised dunes, separated by rather wide gaps of the

"reg" type, rich in well-diversified sessile and vagile benthos.

Dives 20 and 22, between 575 m and 620 m, demonstrated an interesting biosedimentary ensemble, linked to a

fault escarpment which channels the currents (figs 1 1 and 12). Detailed cartography of the area was undertaken us¬

ing a 3.5 kHz sounder and the bathymetric data collected during the Cyana dives. The dunes, in this area, are active

with very soft sediment, devoid of sessile epifauna, even in the gaps between dunes. At the foot of the scarp.

Source

CALSUB CRUISE OFF NEW CALEDONIA

19

Fig. 9. — Location of dives 12, 13 and 17 on the hathyal slope off the north east of Thio. Map without extrapolation

from multibeam sounder of R/V J. Charcot.

Fig. 9. — Localisation des plongees 12, 13 et 17 sur le talus bathyal situe au NE de Thio. Carte sans extrapolation, levee

avec le Sea-Beam du N/O J. Charcot.

Fig. 10. — Location of dives 18 to 22 on the epibathyal gulley between the Isle of Pines and the Great Southern Reef.

Fig. 10. — Localisation des plongees 18 a 22 dans la gouttiere epibathyale entre Hie des Pins et le Grand Recif Sud.

Source

20

M. ROUX

the stronger force of the current excavates a depression several meters deep, covered with blocks of rock, pebbles

and very crude bioclastic sediments. The upper end of the subvertical wall is a hardened ferruginous layer, slightly

overhanging, which supports a very dense population of suspension-feaders, predominantly Echinoderms (Roux el

al„ 1991 a and b).

3-4. - Stylaster Bank

Stylastcr Bank is a guyot situated to the southeast of the Isle of Pines. 23°38' S, 167°43' E. Since the high

swell prevented us from using the Cyana saucer, we carried out instead a series of bathymetric profiles using the

3.5 kHz sounder, in order to gain a precise idea of the morphology and cartography. This guyot is rooted to

surrounding beds situated at a depth of around 1000 m (fig. 13). Its flanks arc relatively regular and very steep. The

summit rises to between 450 m and 500 m depth, and has a very irregular and dissymetric topography. This type

of morphology would seem to lend itself well to testing the possible effects of currents on the density of benthic

populations and the resulting halieutic resources.

Fig. 11. — The fault escarpment environment observed off the South of the Isle of Pines with its benthic fauna.

A: hemipelagic pliocene chalk; B : bioclastic sand; C : "reg" at the foot of the fault escarpment; F : fault;

HG : ferruginous hard ground.

1 : crab ( Geryon ); 2 : actinian; 3 : large bivalve ( Acesta)\ 4 : spherical siliceous sponge; 5 : zoantharian;

6 : ophiacanthid brittle stars with uplifted arms; 7 : suspension-feeding sea stars (Brisingidae); 8 : ophiacanthid

brittle stars with arms crawling on the substrate; 9 : bourgueticrinid stalked crinoid ( Porphyrocrinus ); 10 : regular

soft sea urchin ( Phormosoma ); 11 : spatangid sea urchin ( Taimanawa ). Arrows indicate direction of currents.

This site is also illustrated in fig. 22-2 to 4 and fig. 25-6.

Fig. 11. — L'escarpement de faille observe au sud de File des Pins et sa faune benthique.

A : craie hemipelagique dage pliocene; B : sable bioclastique; C : "reg" en pied d'escarpement; F : faille ; HG :

surface durcie ferrugineuse.

1 : crabe ( Geryon ); 2 : actinie; 3 : bivalve limide ( Acesia)\ 4 : eponge siliceuse globuleuse; 5 : zoanthaire;

6 : ophiure ophiacanthidee aux bras dresses; 7 : asterie brisingidee; 8 : ophiure ophiacanthidee aux bras plaques sur

le substrat; 9 : crinoide pedoncule (Porphyrocrinus)-, 10 : echinide regulier ( Phormosoma ); 11: echinide irregulier

(Taimanawa). Les fleches indiquent le sens des courants.

Ce site est egalemem illustre fig. 22-2 a 4 et fig. 25-6.

Source . MNHN, Paris

CALSUB CRUISE OFF NEW CALEDONIA

21

FlG. 12. — Map of Ihe fault escarpment with hydraulic dunes area off the South of the Isle of Pines, and location of dives

20 and 22.

FlG. 12. — Carte du relief de faille et dc la zone a dunes hydrauliques situes au sud de Hie des Pins, et localisation des

plongees 20 et 22.

DISTANCE in kilometres

Pig. 13. — Bathymetric profile of the Stylaster Bank (3.5 kHz sounder).

Pig. 13. — Profil bathymetrique du banc Stylaster (sondeur 3,5 kHz).

Source :

22

M. ROUX

MAIN OBSERVATIONS AND REMARKS

1. - Remarks on climate and hydrology

During the austral summer. New Caledonia is subject to the influence of equatorial depressions and may at

times suffer from tropical cyclones which arise in the Coral Sea and to the north of the arc formed by Vanuatu and

the Salomon Islands.

The violent winds and the strong rainfall which go hand in hand with tropical depressions and cyclones are

particularly strongly felt by the marine environment, causing strong swells, drop in salinity, serious perturbations

of the currents, homogenisation of the shallow sea water layer, exceptionally high tidal ranges, and resuspension

of sediments from the bed, with a consequent strong increase in the turbidity of the sea water. They probably also

have consequences, for the environment at a much deeper level, as we shall see later. Depressions such as these,

which result in fierce storms, pass close to Grande Terre on average once or twice a year (Rougerie, 1986).

New Caledonia is set within the hydrological context of the Coral Sea, which is very open to the great oceanic

currents running through the central and southwestern Pacific. The hydrology in the basin of the Loyalty Islands

during the winter of 1981 was analysed by D. Guevel (1983). In addition, the Cyana diving saucer recorded

temperature contours at the time of the Calsub dives at the end of the summer of 1989 (fig. 14).

The vertical thermohaline structure in winter (fig. 14C) shows a salinity maximum of 35.68 % between 150

and 170 m, thereby separating the southern layer of subtropical water from the superficial layer of water

Sea water temperature in°C Salinity %o

Fig. 14. — Vertical hydrological profiles near the east edge of Loyalty ridge (A) and basin (B and C).

A and B : data recorded by the diving saucer Cyana during Calsub cruise ; C : modified from Guevel, 1983.

CS : shallow water layer; ESS : southern subtropical water layer; EAI : intermediate deep antarctic water layer.

Fig. 14. — Profils hydrologiques au large de la Nouvelle-Caledonie. Structure thermique et thermohaline verticale a Test

de la ride (A) et dans le bassin des Loyaute (B et C) .

A et B : donnees enregistrees par la soucoupe Cyana lors de la campagne CALSUB; C : modifid d'apres Guevel, 1983.

CS : couche superficielle; ESS : eau subtropicale sud; EAI : eau antarctique intermediate.

Source

CAL.SU B CRUISF. OFF NEW CALEIX)NIA

23

which is saltier to the South (subtropical influence) than to the North (equatorial influence). The intermediate

antarctic water, which is travelling north, has an average 34.50% salinity and a 5°C temperature in this area.

Minimal salinity values occur between 700 m and 1 100 m. Its upper limit has been defined at nearer 600 m close

to the barrier reef off the coast at Noumea (Rougerie. 1986). This conclusion is also suggested by the Calsub

thermal readings (fig. 14 A and B) and it may even rise further towards the end of summer off the coast of Thio,

which would explain the rapid fall in temperature between 400 m and 500 m.

The cartography of the water masses in the Loyalty basin (Guevel, 1983) demonstrates the following : (1) a

superficial frontal zone to the west of Mare which separates warmer and less salty water in the north from that in

the south; (2) between 0 to a depth of 200 m, a geostrophic current running towards the SE, which dominates the

NW drift resulting from the trade winds; (3) at a depth of 1000 m, a geostrophic current running towards the NW.

Seasonal changes in the principal surface currents are more noticeable on the western edge of Grande Terre, as

well as to the S and SE of the Isle of Pines. The strength of the currents observed between 300 m and 650 m by

the CALSUB dives between the Isle of Pines and the Great Southern Reef may be due to seasonal conditions which

could cause to the upper part of the antarctic water layer to rise, channelled by the morphology of the gulley. It

may also be due to the effect of internal waves being propagated along the upper limit of the antarctic water layer.

The moving hydraulic dunes observed during dive number 22. as well as the presence of Echinoderms which we

would normally expect to encounter much deeper (see later) all indicate the frequent, if not almost permanent

influence of antarctic water at a depth of around 600 m. The more stabilised dunes observed at around 350 m during

dive 21 , are probably due to a more exceptional activity of deep currents likely to give rise to them (fig. 15).

NW

SE

Rocky

substrate

water

Fig. 15. — Axial profile of the epibathyal gulley between the Isle of Pines and the Great Southern Reef.

A : frequent upwellings of antarctic water explaining active hydraulic dunes. B : exceptional upwellings of antarctic

water allowing the formation of stabilised hydraulic dunes and development of a diversified benthic fauna.

FIG. 15. — Profil axial de la gouttiere epibathyale situee entre Hie des Pins et le Grand Recif Sud.

A : remontees frequentes de l'eau antarctique intermediate entretenant des dunes hydrauliques vives. B : remontees

exceptionnelles permettant la stabilisation des dunes hydrauliques et la colonisation par une faune benthique

diversifiee.

2. - Bathymetric distribution of communities

Because these results have already been presented and discussed in some detail elsewhere (Roux et al, 1991 a

and b), we shall restrict ourselves to a summary of the main aspects here. In the bathyal domain of

New Caledonia, it appears that the changes in fauna, as we head deeper down into the ocean, are closely linked to

the limits of the different water layers.

24

M. ROUX

It is at depths of around 100 m to 150 m that the break in the slope limiting the platform usually occurs, and

that the fall in luminosity is most noticeable. In Santal Bay. encrusting green algae, of the Palmophyllum- type,

are associated with rhodolites at around 1 10 m, whereas Melobesia-lype algae are found as deep as 145 m.

Compact algal structures, supporting a rich fauna of large gorgonians, sponges and antipatharians (figs 21-1 and

21-2), develop between 60 m and 100 m. Algal slabs with scoriaceous surfaces tend to lose their blanket of sessile

megafauna, particularly when the incline becomes steeper, at around 80 m to 100 m (fig. 21-3). Beyond 100 m,

algal balls dominate, together with a primarily vagile epifauna (fig. 21-4). The substratum’s biological cover

becomes more dispersed and its bathyal character becomes evident from 150 m downwards, when the layer of

southern subtropical water is entered.

The epibathyal domain (from approximately 150 m to 700 m) is characterized by a wide variety of beds. Rocky

slabs, covering soft sediment, undermined by the burrowing of crustaceans or fish, conserve caverns sheltering a

number of sessile and vagile organisms (figs 22-1 and 23-4). The hard beds alternate with sedimentary beds°of

varying stability and granulometry (ranging from masses of fallen rock to hemipelagic ooze). They arc frequently

covered by vegetal debris, at times of some considerable size. This debris becomes rarer as we go deeper; however,

we did observe a tree, probably uprooted during a cyclone, at a depth of about 2000 m (fig. 23-6).

On the upper part of the talus, the rocky surfaces are often swept by avalanches of sediment, which is at times

quite crude, coming from the outer edge of the platform (fig. 23-2). On these unstable substrates, only a vagile or

semi-vagile fauna develops (holothurians, sea stars, comatulid crinoids, gasteropds). Those surfaces better sheltered

from these movements of sediment (figs 21-5 and 23-5) show a great diversity of siliceous sponges, some of

which (fistulous sponges, clionids) seem to participate in the cupulatc erosion of the substratum (Gaillard

1991 a).

Between 300 m and 700 m. large steep indurated surfaces support different communities of suspension-feeders

(gorgonians, echinoderms, sponges) (fig. 22-5). On the Ouvea-Lifou rise, each community is clearly dominated by

one or two species, and it is possible to observe groups consisting of a single species, particularly the

echinoderms (fig. 23-3). In the area to the SW of the Isle of Pines, the richness and abundance of the macrofauna

reaches high levels. Between 320 m and 410 m. in the zone of the relatively stabilised dunes (fig. 22-6), the

epifauna is extremely diverse (brachiopods, gasteropods. Stylaster corals, sponges including lithistids, echinids,

stalked crinoids, crustaceans), with a regular and fairly homogeneous spread. At around 600 m. the fault scarp, at

the foot of which moving hydraulic dunes develop (fig. 25-6) demonstrates, in a relatively reduced space, how the

intensity of the currents and the nature of the substrate can affect the composition of benthic communities

(fig. 11). The summit of the scarp supports very dense communities of suspension-feeders, dominated by

echinoderms (fig. 22-2 to 4). A large part of this fauna, particularly among the echinoderms ( Phormosoma and

bnssinginds, for example), is in great abundance, and normally more characteristic of depths below 1000 m. In

fact, this site marks the beginning of the transition zone between the beds washed by subtropical water and those

washed by intermediate antarctic water.

This transition generally occurs between 700 m and 1000 m. The communities in this zone, which are often

rather peculiar, may still be quite dense. What singles them out is (he presence of hexactinellid sponges, notably

the abundance of Monorhaphis (figs 24-1 and 24-2). These communities already contain taxa which, as we shall

see, dominate the deepest areas explored (e. g. hyocrinid crinoids. Euplectella, Hyalonema , Iridogorgia -fig. 16A-,

xenophiophorids, etc.).

Beyond 1000 m. the macro- aid megafauna become definitely more scarce, while remaining quite diversified.

Individual specimens are apt to grow very lage, in some cases over one metre. The stalked hexactinellid sponges

become very diversified (fig. 24-6). The sponges and dead branches of gorgonians frequently support suspension-

feeders (comatulid crinoids, sea stars, brittle stars, actinians), at times in thick bunches (fig. 24-3). Between

1000 m and 1600 m, the megafauna on the rocky surfaces is characterised by the simultaneous presence of three

forms : two homeomorphic stalked crinoids (. Proisocrinus and Guillecrinus) (figs 24-4 and 24-5) and a gorgonian

of the genus Metallogorgia (fig. 16B). At around 2000 m. we see the first examples of large Euplectella , attached

to the gently sloping rocky beds. In the sedimentary zones, the relative importance of bioturbation increases with

depth (fig. 26-1), and different ichnofacies arc the distinguishing markers for lower or upper beds at a depth of

around 1800 m (Gaillard, 1991).

Source

CALSUB CRUISE OFF NEW CALEDONIA

25

Fig. 16. — Two bathyal gorgonians with a typical morphology.

A : a large specimen of the genus Iridogorgia on the slope off Thio (dive 12. depth 880 m, observer H. ZlBROWius);

B : a specimen of the genus Melallogorgia on the mesobathyal slope off Thio (dive 13, depth 1766 m, observer

M. Segonzac).

Fig. 16. — Deux gorgones bathyales avec une morphologie typique.

A : un grand individu du genre Iridogorgia sur le talus au large de Thio (plongee 12, profondeur 880 m, observateur

H. ZlBROWius); B : un individu du genre Melallogorgia sur le talus mesobathyal au large de Thio (plongee 13,

profondeur 1766 m, observateur M. SEGONZAC).

3. - Notes on a few particular taxa

3-1. - Behaviour of the cephalopods

Solitary specimens of the New Caledonian naulilus (Nautilus macromphalus) were encountered several times

between 300 m and 545 m during dives carried out during the day around the Loyaly Islands. A piece of bait

(chicken), placed on the sea bed 10 to 12 hours prior to dives 19 (413 m) and 22 (610 m), to the SW ol the Isle of

Pines, attracted a group of around fifteen nautilus specimens during dive 19 (fig. 23-1); only crustaceans (Geryon

and Heterocarpus) came to the bait for dive 22. Our observations of the bathymetric distribution of the nautili

confirm the data acquired through use of lobster pots between 1976 and 1978 (Int£s, 1978; Rancurel, 1990).

The specimens encountered by the Cyana saucer were very active and swam efficiently, without crashing

against the rocky walls. Their speed can reach between 20 and 30 cm/s (Roux, 1990). which indicates that their

physiological optimum would seem to be superior in an epibathyal environment, rather than under experimental

conditions on the surface, where their ability to oxygenate does not seem to allow them to exceed a speed ot

16 cm/s (Wells & Wells, 1985; Wells. 1987 and in lit/.).

Two very different types of behaviour were noted in the cirrates (Cirroctopoda). The most spectacular recorded

and photographed sequences were those of dives 3 (Cirrothauma) and 16 (Opisthoteuthis).

26

M. ROUX

Ai a depth of 2880 m, a specimen (which may belong to the genus Cirrothauma), when touched by the claw of

the Cyana, quickly inflated his interbrachial membrane into a turgescent balloon, reminiscent of a pumpkin

(figs 17A and 17B) (Boletzky el a!.. 1992). This may be a means of capturing prey which, once spotted, would

be engulfed in the brachial ensemble. However, in an environment where light docs not penetrate, tactile stimuli

are likely to be important and the ballooning response might have a stunning or disorientating effect on a potential

predator in a first (possibly accidental) contact with the cirrate. This could thus be a defense mechanism.

At a depth of 812 m, a specimen of the genus Opisthoteuthis demonstrated that it was able to pass unnoticed

on a sloping rocky substratum by retracting itself into the form of an hemisphere (figs 17C and 17D). These two

octopods are capable of "gliding" by stretching out their arms and interbrachial membranes, thereby moving like a

parachute between two active phases of swimming.

3-2. - Behaviour of some of the fish encountered

B. S£ret gives a list (table 2) and pictures (figs 18 and 19) of the main species of fish observed during the

course of the Calsub cruise and of which we have photographs or video sequences taken aboard the Cyana saucer.

Here he also gives additional observations and remarks on the behaviour of a few kinds of fish we observed.

Numerous pictures of fish were taken during the Cyana dives, but only about sixty could be identified to

species, genus or family from the morphological characters observed. The identifications of taxa arc listed in table

2, with doubts remaining for a few attributions because of the difficulties of basing determinations on photographs

only. However, two specimens were photographed and collected by A. Guille during dive 20 : one specimen

(362 mm LT) of the macrourid rat-tail fish Coelorhinchus anatirostris (photo 023, collection MNHN 1989-965)

and one specimen (613 mm LS) of the trichiurid silver scabbardfish Lepidotus caudatus (photo 061, collection

MNHN 1989-966).

We have selected the following remarks on the behaviour of a number of species observed from Cyana.

Numerous sharks, mainly the shortnose spurdog Squalus megalops (fig. 18D) and the kitefin shark Dalatias

licha (fig. 18E) were observed. These observations are corroborated by the shark specimens frequently collected

during recent investigations on several seamounts off New Caledonia. Photograph 385 of dive 19 is a good

illustration of the opportunistic scavanging behaviour of the kitefin shark Dalatias licha : attracted by a bait, they

wait for the Nautilus to finish feeding (fig. 23-1) without attacking any of them. Another observation of interest is

that of the behaviour of Squalus megalops : this demersal shark was resting on the sea floor, evidence that its

respiration is not dependent on permanent movement as had frequently been assumed. Specimens of a new species

of cat shark belonging to the genus Galeus or the genus Halaelurus were observed, two genera never before

recorded in New Caledonian waters. During dive 22 at a depth of 613 m. a specimen of the skate genus Notoraja

was photographed (photo 093) resting on the flank of a bioclastic sandy dune; this is probably a new species, a

few juvenile specimens of which were collected during Biogeocal cruise.

Fig. 17. — Behaviour of two cirrate octopods.

A : just before contact with Cyana' s claw, a specimen having affinities with the genus Cirrothauma gliding through

the water, arms outstretched and showing the well-developed interbrachial membrane. B : the same specimen reacting

to contact with the claw by inflating its interbrachial membrane into a turgescent balloon (dive 3, depth 2880 m,

observer M. Rio).

C : a specimen of the genus Opisthotheulis swimming. D: the same specimen several minutes after the previous shot,

rolled into a ball on the rocky substrate (Dive 16, depth 813 m, observer M. ROUX).

Fig. 17. — Comportement de deux octopodes (Cirrates).

A : juste avant le contact avec la pince de Cyana. un individu proche du genre Cirrothauma "planant" entre deux caux,

les bras etendus et montrant la membrane interbrachiale bien developpee. B : le raeme individu reagissant au contact

de la pince en gonflant sa membrane interbrachiale en un ballon turgescent (plongee 3, profondeur 2880 m,

observateur M. Rio).

C : un individu du genre Opisthotheulis nageant. D: le memo individu, quelques minutes apres la vue precedente, pose

sur le substrat rocheux el retracte en boule sur lui-meme. (Plongee 16, profondeur 813 m, observateur M. Roux).

Source : MNHN, Paris

CALSUB CRUISE OFF NEW CALEDONIA

27

Source MNHN. Paris

28

M. ROUX

TELEOSTEANS

Synaphobranchidae

Source : MNHN, Paris

CALSUB CRUISE OFF NEW CALEDONIA

29

T rachichlhyidae

Table 2. — Identification of fishes from photographs taken by Cyana during the CALSUB cruise (listed by B. Sf-RET).

Tableau 2. — Liste des poissons identifiables photographies par Cyana lors de la campagne CALSUB (determinations par

B. SERET).

Among the teleostean fishes, several individuals of about three species were observed. The first example in

New Caledonia of Coelorhitichus anatirostris was photographed and collected during the Calsub cruise, a species

which was only previously known from Japanese waters and the China Sea.

Fig. 18. — A few fishes observed during the Calsub cruise.

A : oilfish belonging to the genus Rexea (dive 06; depth 1137 m; observer B. LAURIN). B : conger eel attributable to

Conger cf. cinereus (dive 17; depth 2033 m; observer C. Gaillard). C : ruby snapper of the species Etelis coruscans,

circling the saucer before coming to rest on the sea bed within range of the spotlights (dive 16; depth 512 m;

observer M. ROUX). D : spurdog of the species Squalus megalops, resting on the sea bed (dive 21; depth 332 m;

observer B. Richer de Forges). E : kitefin shark, Dalatias licha, lurking around the manipulating arm of the saucer

(dive 19; depth 415 m; observer M. SEGONZAC).

Fig. 18. — Quelques poissons observes durant la campagne CALSUB.

A : individu attribue au genre Rexea (plongee 6, profondeur 1137 m, obscrvateur B. LAURIN). B : congre attribuable a

Conger cf. cinereus (plongee 17, profondeur 2033 in, observateur C. Gaillard). C ; vivaneau de l'espece Etelis

coruscans, tournant autour de la soucoupe avant de s'immobiliser, post; sur le fond dans le champ des projecteurs

(plongde 16, profondeur 512 m, observateur M. ROUX). D : requin de l'espece Squalus megalops se reposant sur le fond

(plongee 21, profondeur 332 m, observateur B. RICHER DE FORGES). E : requin de l'espece Dalatias licha, rodant autour

du bras manipulateur de la soucoupe (plongee 19. profondeur 415 m, observateur M. SEGONZAC).

30

M. ROUX

Source : MNHN, Paris

CALSUB CRUISE OFF NEW CALEDONIA

31

The two wonderful species of ruby snapper observed exhibit very different behaviour on the approach of the

saucer. We were only able to take photographs of Etelis carbunculus on exceptional occasions because it swam off

rapidly every time the saucer came close. By contrast. Etelis coruscans moves more slowly and willingly circles

around the saucer. During dive 16 at a depth of 512 m. one individual came to rest just in front of Cyana and

remained motionless, as if immobilised by the spotlights (fig. 18C).

During dive 23 at a depth of 2015 m. a beautiful example (fig. 19) of the iripodfish Bathypterois cf. guentheri,

a species typical of the abyss, was filmed swimming and coming to rest in its classic position, perched on its long

pectoral fins and caudal fin. like a tripod, lying in wait for its prey.

Fig. 19. — An abyssal tripodfish attributable to Bathyplerois cf. guentheri about to come to rest on a sedimentary bed

(dive 02; depth 2015 m; observer M. Roux).

Fig. 19. — Un poisson abyssal "tripode" attribuable a Bathyplerois cf. guentheri pret a se poser sur un fond sedimentaire

(plongee 2. profondeur 2015 m, observatcur M. ROUX).

Numerous fish, of all sizes, burrow in the sediment and are therefore agents of bioturbation. Some, generally

of small to medium size, were seen to dig head-firsl into the sand, or to work the sand along a stretch of several

decimetres (fig. 25-5); it is clear that they feed on the superficial endofauna in this way. Others, often slightly

larger (groupers, ruby snappers) shelter in niches dug by the movement of soft sediment under an indurated level.

This burrowing can lead to the creation of cavernous substrates which are home to a rich, vagile fauna (VANNEY,

1991).

3-3. - The importance of bioturbation and biocorrosion

An inventory of the principal biogenic tracks has been drawn up (fig. 25 and 26) (Gaillard. 1991 a and b). Il

is at depths of between 1800 m and 2000 m that bioturbation seems to be at its most diversified (fig. 20). The

most original discoveries were the observation of the ichnogeni Paleodictyon and Urohelminthoida, exceptional in

this area, and lhal of a new track ("track FC"). which was very common and characteristic of ihe bathymetric level

between 1600 m and 2000 m (fig. 26-6 and 26-8). The bioturbation of the soft sediment is variable since it

depends to a large extent on the rhythm and the degree of external elements, parlicularly turbiditic faecal elements.

Biocorrosion of the rocky substrata is substantial towards the lop ot the slopes, but is more difficult to evaluate in

the semi-indurated oozes of the talus, since the numerous cavities witnessed here may be of recent origin

(biocorrosion) or former burrows in the process of being eroded (bioturbation). Fish and crustaceans sweep soft

sediment away from under indurated surfaces, thereby contributing to the formation of cryptic ecological niches.

Whatever the answer, it is certain that the action of organisms plays a role in the general process of erosion.

32

M. ROUX

Fig. 20. — Diversity and bathymetric range of traces and burrows off New Caledonia (from GaILLARD, 1991b).

Fig. 20. — Diversity et distribution bathymetrique des traces et terriers au large de la Nouvelle-Caledonie (d'apres

Gaillard, 1991b).

Generally speaking, the role played by both biocorrosion and bioturbation would seem to be essential to the

sedimentary dynamics of the zones we explored, and to the evolution of benthic populations, particularly as regards

to the relative importance of the sessile epifauna.

Source MNHN, Paris

CALSUB CRUISE OFF NEW CALEDONIA

33

4. - A word about "living fossils"

One of the most interesting aspects of the bathyal fauna of New Caledonia is the apparently high number of

tax a with what we might call an "archaic" stamp to them, "living fossils", relics of the fauna of the jurassic and

cretaceous Tethys Ocean. This phenomenon is particularly spectacular among the stalked crinoids (AmSziane-

Cominardi el al., 1987 and 1990; Bourseau el al., 1991) (figs 24-4 and 24-5), among the sponges, with the

presence of sphinctozoans (Vacelet el al., 1992) (fig. 21-6) and among the pterobranches. The pterobranch

species Cephalodiscus graptoloides, collected during the Calsub cruise, and identified as a specimen of interest by

H. Zibrowius, has an astonishing ressemblance to graptolites, a group considered to have been extinct since the

Carboniferous (Dilly, 1993; Rigby, 1993). Note also the discovery of a new line of evolution among the deep

sea ascidians, Fimbrora calsubia. which shows a new type of adaptation to macrophagy (Monniot & Monniot,

1991)

Up until the Lower Cretaceous, New Caledonia formed part of the eastern edge of Gondwana. When Gondwana

broke up. New Caledonia separated from Australia and then drifted towards the lower latitudes which, from a

climatic point of view, may have in some way made up for the very noticable cooling during the second half of

the Ccnozoic, which was to cause the Quaternary glaciation. During the most recent glacial maximum, the level

of the sea dropped by 120 m, causing the New Caledonian lagoon to emerge completely and destabilising the

upper portion of the bathyal slope. In any case, it is difficult to imagine that conditions in the New Caledonian

environment have remained unchanged since the Mesozoic.

The Calsub cruise demonstrated that there is a great variety of ecological niches, but also a relative instability

(both sedimentological and hydrological) in the epibathyal zone of New Caledonia (Lambert & Roux, 1991).

From 1000 m downwards, the environment seems to be more stable and the populations more homogeneous, as in

the Central Pacific.

When we talk of "living fossils", we may be talking of two things; on the one hand, they may be veritably

panchronic forms (slow or imperceptible evolution), or, on the other hand, forms that have undergone a more re¬

cent paedomorphic evolution, leading to a phenotypic convergence with ancient forms (A.m£ziane-Cominardi &

Roux, 1994). Only a certain permanence in the conditions of the environment allows us to assume the first pos¬

sibility, unless we are dealing with extremely opportunist taxa, capable of surviving great environmental changes.

In fact, the conditions for survival of ancestral forms depend a great deal on the autecology of each taxon and

we must therefore beware of making generalizations on the basis of the analysis of a single group of organisms.

CONCLUSION

The bathyal domain of New Caledonia is currently one of the best known in the Pacific, as much in terms of

the inventories of fauna as in terms of the direct observation of its populations by submersible. The Calsub

cruise has shown the great diversity of the ecological niches, particularly on the epibathyal beds, of which it

explored only a very small portion. It remains to explore in more detail the populations of fauna which develop at

the summit of guyots.

In spite of the considerable exploration efforts made, and given the diversity of the substrate and of the

environmental conditions, our knowledge of the New Caledonian bathyal fauna and its ecology still remains very

sketchy, particularly in regard to the rocky beds, where only the use of a submersible allows significant progress.

We see therefore how difficult it is to gain an idea of the real originality of the fauna in this region of the Western

Pacific. Its apparent originality is due in part to the absence or rarity of similar data from other areas of the same

ocean. The "living fossils" aspect, however spectacular it might seem, is probably more a reflection of a lack of

knowledge about the current bathyal benthos and of gaps in paleontological records, than an indication of the

existence of a New Caledonian sanctuary for fauna directly descended from the most distant geological eras.

However, we must not exclude from our study the unusual historical and climatic factors. It remains now for

further cruises to demonstrate these things, cruises which will be better targeted thanks to the invaluable direct

observations made by the Calsub cruise.

34

M. ROUX

ACKNOWLEDGEMENTS

The author is indebted to the other members of the Calsub cruise and especially to the Cyana team for their

contribution to the success of our investigations off New Caledonia. Many illustrations (fig. 21 to 26) are

reproduced here with the persmission and the courtesy of C. Montenat (I.G.A.L.). Thanks to A. Beaudon

(Reims) for drawings. Calsub cruise was supported by INSU-CNRS and this paper is a contribution of the

Ecoprophyce research group and of the UR A 157 of the CNRS.

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MONNIOT, C. & MONNIOT, F.. 1991. — Decouverte dune nouvelle lignee evolutive chez les ascidies de grande profondeur :

une Ascidiidae carnivore. C.R. Acad. Sci. Paris, ser. Ill, 312 : 383-388.

Pascal, A., Rio, M. & VaNNEY, J.R., 1992. — Les surfaces durcies des pentes bathyales de Nouvelle-Caledonie (Pacifique

du SW). Morphologies et diagenese. Bull. Soc. gdol. France, 163 (3) : 271-280.

RaNCUREL, P., 1990. — Contribution a la connaissance de la repartition bathyntetrique de Nautilus macromphalus

Sowerby. Haliotis, 10 : 71-81.

RICHER DE Forges, B„ 1990. — Les campagnes d'exploration de la faune batliyale dans la zone economique de Nouvelle-

Caledonie. In : Resultats des Campagnes Musorstom, Volume 6. Mem. Mus. natn. Hist, nat., Paris, (A), 145 : 9-54.

Rigby, S., 1993. — Graptolites come life. Nature, 362 : 209-210.

Rio, M., RoOx, M., GuERIN, H. et 1'equipe Calsub, 1991. — Le substrat geologique et les processus sedimentaires sur les

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Source MNHN. Paris

CALSUB CRUISE OFF NEW CALEDONIA

35

ROUGERIE, F., 1986. Le lagon Sud-Ouest de la Nouvelle-Caledonie : specificity hydrologique, dynamique et

productivity. Etudes et theses, ORSTOM (ed.), Paris, 234 p.

Roux, M., 1990. — Underwater observations of Nautilus macromphalus off New Caledonia. Chambered Nautilus

Newslett., 60 : 1.

ROUX, M., 1991. — La Nouvelle-Caiydonie et ses alentours. Cadre gdologique et ocyanographique du programme

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Nouvelle-Caiydonie. Doc. et Trav. IGAL, Paris. 15 : 22-36.

Roux, M., Bouchet, P„ Bourseau, J.-P„ Gaillard, C„ Grandperrin, R„ Guille, A., Laurin, B„ Monniot, C., Richer

DE FORGES, B., Rio, M., Sf.GONZAC, M., Vacelet, J. & ZlBROWius, H., 1991a. — L'environnement bathyal au large de

la Nouvelle-Caledonie : resultats pryiiminaires de la campagne Cai.SUB et consequences paleoecologiques. Bull. Soc.

geol. France. 162, (4) : 675-685.

Roux, M., Bouchet, P., Bourseau, J.-P„ Gaillard, C., Grandperrin, R„ Guille, A., Laurin, B., Monniot, C„ Richer

DE FORGES, B., Rio, M., SEGONZAC, M., Vacelet, J. & ZlBROWius, H., 1991b. — L'etagement du benthos bathyal en

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Caledonie. Doc. et Trav. IGAL. Paris, 15 : 151-165.

Vacelet, J., Cuif, J.-P., Gautret, P., Massot, M„ Richer de Forges, B. & Zibrowius, H., 1992. — Un sphinctozoaire

colonial apparente aux constructeurs de recifs triasiques survivant dans le bathyal de Nouvelle-Caledonie. C R Acad

Sci. Paris, ser. Ill , 314 : 379-385.

Vanney, J.R., 1991. — Le modele des pentes sous-marines observees par submersible lors de la campagne Calsub. In :

B. Lambert et M. Roux (coord.), L'environnement carbonate bathyal en Nouvelle-Caiydonie. Doc. et Trav. IGAL

Paris. 15 : 38-55.

Vanney, J.R., Rio, M., Roux, M., GufiRlN, H. et l'equipe Calsub, 1992. — Morphologies sous-marines particuliferes lides

a des circulations hydrothermales sur la ride des Loyaute (Nouvelle-Caledonie). Bull. Soc. gdol. France. 163 (3) : 255-

262.

Wells, M.J., 1987. — Ventilation and oxygen extraction by Nautilus. In : Nautilus. The biology and paleobiology of a

living fossil, Saunders W.B. & Landman N.H. edit., Topics in Geobiology, Pleneum Press, 6 : 339-350.

Wells, M.J. & Wells, J., 1985. — Ventilation and oxygen uptake by Nautilus. J. exp. Biol., 118 : 297-312.

36

M. ROUX

FIGURE 21

1. — Algal bed supporting large gorgonian communities. S of Santal Bay. W of Lifou. Depth 80 m. Dive 1 1;

observer J.-P. BOURSE AU.

1. — Trottoir algaire supportant une prairie de grandes gorgones. Sud de la baie du Santal a l'ouest dc Lifou.

Profondeur 80 m. Plongee 11; observateur J.-P. Bourseau.

2. _ Algal bed covering a dome and supporting lithistid sponges, gorgonians and anthipatharians. SW of the Isle

of Pines. Depth 60 m. Dive 18; observer P. BOUCHET.

2. — Dome couvert d'une construction algaire supportant des eponges lithistides, des gorgones et dcs

antipathaires. SW de Pile des Pins. Profondeur 60 m. Plongee 18; observateur P. Bouchet.

3. — Algal bed. S of Santal Bay. Depth 80 m. Dive 11; observer J.-P. Bourseau.

3. _ Trottoir algaire. Sud de la baie du Santal. Profondeur 80 m. Plongee 11; observateur J.-P. Bourseau.

4. — Large holothurian (about 80 cm long) on a slope with algal balls and bioclastic sand. S of Santal Bay.

Depth 130 m. Dive 11; observer J.-P. Bourseau.

4. — Grande holothurie (environ 80 cm) sur une pente h boulets algaires et sable bioclastique. Sud de la baie du

Santal. Profondeur 130 m. Plongee 11; observateur J.-P. Bourseau.

5. — Large coniform sponges on an escarpment encrusted with small sponges (biocorrosion?). N of Santal Bay.

Depth 380 m. Dive 05; observer R. Grandperrin.

5. — Grandes Sponges en comet sur une paroi incrustce de petites Sponges (biocorrosion ?). Nord de la baie du

Santal. Profondeur 380 m. Plong6e 5; observateur R. GRANDPERRIN.

6. — Small decimetrical structure of sphinctozoan sponge ( Vaceleiia ) observed at a depth of around 240 m in

Santal Bay (photograph taken by J. Vacelet).

6. — Petit massif decim6trique construit par les sphinctozoaires (Vaceleiia) observ6s vers 240 m dans la baie du

Santal (cliche J. Vacelet).

Source MNHN. Paris

Source : MNHN, Paris

38

M. ROUX

FIGURE 22

1. — Cavern carpeted with gorgonians. bryozoans and comatulid crinoids, home to a dense shrimp population of

the genus Plesionika. Sanlal Bay. Depth 180 m. Dive 10; observer C. Monniot.

1. — Grotte tapissee de gorgones. bryozoaircs, comatules et abritant une population de crevettes du genre

Plesionika. Sud de la baie du Santal. Profondeur 180 m. Plongce 10; observateur C. Monniot.

2. — Overhang at the top of a fault escarpment (see fig. 12} - side view. On the lower surface, zoantharians; on

the edge, brissingid sea stars; on the upper surface, ophicanthid and euryalid brittle stars. S of the Isle of Pines.

Depth 584 m. Dive 20; observer A. Guille.

2. — Surplomb au sommel d'un escarpement de faille, vue de profil (cf. fig. 12}. En dessous ; zoanthaires ; a

l'extrdmite : asteridcs brissingiddes ; au-dessus : ophiures ophiacanthiddes et euryales. Sud de l’tle des Pins.

Profondeur 584 m. Plongce 20; observateur A. Guii.le.

3. — Dense echinoderm population at the top of the same fault escarpment (see fig. 12). Same location. Depth

597 m. Dive 22; observer M. Roux.

3. — Agregat d'echinodermes au sommet d'un escarpement de faille (cf. fig. 12). Meme lieu. Profondeur 597 m.

Plongee 22; observateur M. Roux.

4. — Upper surface of the same escarpment entirely covered by ophiacanthid brittle stars. Same dive. Same depth.

4. — Meme lieu, population d'ophiures ophiacanthidees sur le replat sommital. Meme profondeur. meme

plongde.

5. — Slab of rock covered at regular intervals by small gorgonians (in the background), and, in the foreground, a

large gorgonian of the genus Calypirophora supporting an euryalid brittle star with partially retracted arms (on

the left). N slope of the Ouvda-Lifou rise. Depth 355 m. Dive 15; observer A. Guille.

5. — Dalle regulierement couverte de petites gorgones (en arricre-plan) et de plus grandes du genre Calypirophora

(au premier plan) supportant une euryale aux bras en partie retractes (a gauche). Versant nord du seuil Ouvea-

Lifou. Profondeur 355 m. Plongee 15; observateur A. Guille.

6. — Lithistid sponge ( Corallistes undulatus), diameter 30 to 40 cm. The sea floor is covered by terebratulid

brachiopods (small shadows). SW of the Isle of Pines. Depth 330 m. Dive 21; observer B. Richer de Forges.

6- — Eponge lithistide (Corallistes undulatus), 30 a 40 cm de diametre, sur un fond riche en brachiopodes

(terebratules) dont la presence est ici soulignee par leur ombre. SW de Tile des Pins. Profondeur 330 m.

Plongee 21; observateur B. Richer de Forges.

Source MNHN. Paris

Source : MNHN, Paris

40

M. ROUX

FIGURE 23

1. — Group of Nautilus macromphalus aitracied to a bait. SW of the Isle of Pines. Depth 413 m. Dive 19;

observer M. Segonzac.

1. — Groupe de Nautilus macromphalus attires par un appat. SW de 1’tle des Pins. Profondeur 413 m.

Plongee 19; observateur M. Segonzac.

2. — Allochthonous blocks and pebbles from a collapsed algal bed mixed with a group of autochthonous nautilus

shells. Northern slope of the Ouvda-Lifou rise. Depth 360 m. Dive 16; observer M. Roux.

2. — Blocs eboules et graviers provenant d'un trottoir algaire melanges & une concentration de coquilles de

nautiles autochtones. Versant septentrional du seuil Ouvea-Lifou. Profondeur 360 m. Plongde 16; observateur

M. ROUX.

3. — Population of cidarid sea-urchins on a steeply inclined slab of rock. Same location. Depth 370 m. Dive 15;

observer A. GuiLLE.

3. — Population de cidarides (6chinides rcguliers) sur une dalle pentue. Meme lieu. Profondeur 370 m.

Plongee 15; observateur A. GuiLLE.

4. — Dense macrofauna in a sheltered area (bryozoans, sponges, scleractinians, ascidians. etc.). S. of Santal Bay.

Depth 525 m. Dive 09; observer J. Vacelet.

4. — Macrofaune plus dense dans une zone abritde (bryozoaires. eponges, scldractiniaires, ascidies. etc...). Sud de

la baie du Santal. Profondeur 525 m. Plongee 09; observateur J. Vacelet.

5. — Hexactinellid sponge attached to a slab of rock by a thick cushion of spicules. Northern slope of the Ouvea-

Lifou rise. Dive: 16; observer M. Roux.

5. — Eponge hexactinellide sur une dalle rocheuse fixee par un coussin de spicules. Versant septentrional du seuil

Ouvda-Lifou. Plongde 16; observateur M. Roux.

6. — Tree uprooted by a storm lying on the sea floor at a depth of 1834 m. Notice the galatheid shrimps

(Munidopsis) grasping the roots. NE of Thio. Dive 17; observer C. Gaillard.

6. Epave d un arbre deracine par un cyclone. On distingue des galathees (Munidopsis ) accrochees aux racines.

NE de Thio. Profondeur 1834 m. Plongee 17; observateur C. Gaillard.

Source MNHN, Paris

Source : MNHN, Pahs

42

M. ROUX

FIGURE 24

1. — Hexactinellid sponge (Monorhaphis), 50 to 80 cm long. N of Santal Bay. Depth 875 m. Dive 08; observer

B. Laurin.

1. — Eponge hexactinellide (Monorhaphis ). longueur 50 a 80 cm. Nord de la baie du Santal. Profondeur 875 m.

Plongee 08; observateur B. Laurin.

2. — Dense population of Monorhaphis. The spicule of a number of dead specimens is covered by epibiontic

macrofauna. Depth 816 m. Same dive.

2. — Champ de Monorhaphis , comportant de nombreux individus morts dont le spicule est couvert par les

dpibiontes. Profondeur 816 m. Meme plong6e.

3. — Ophiacanthid and euryalid brittle stars perched on the axis of a dead gorgonian. NE of Thio. Depth 1627 m.

Dive 13 ; observer M. Segonzac.

3. — Ophiurcs (ophiacanthidees et euryales) perchdes sur un axe de gorgone morte. NE de Thio. Profondeur

1627 m. Plongee 13; observateur M. Segonzac.

4. — Stalked crinoid with jurassic affinities ( Proisocrinus ruberrimus). N of Lifou. Depth 1254 m. Dive 02;

observer M. Roux.

4. — Crinoi'de pedoncule d'affinitds jurassiques (Proisocrinus ruberrimus). Nord de Lifou. Profondeur 1254 m.

Plongee 02; observateur M. Roux.

5. — Stalked crinoid with paleozoic affinities (Guillecrinus neocaledonicus). Depth 1276 m. Same dive.

5. — Crinoi'de pedoncule d’affinites paleozoiques (Guillecrinus neocaledonicus ). Profondeur 1276 m. Meme

plongee.

6. — Hexactinellid stalked sponge (Hyalonema) with a comatulid crinoid attached to the upper part of the stalk.

N of Lifou. Depth 1820 m. Dive 01; observer C. Monniot.

6. — Eponge hexactinellide pedonculee (Hyalonema ) avec une comatule fixee au sommet du pedoncule. Nord de

Lifou. Profondeur 1820 m. Plongee 01; observateur C. MONNIOT.

7. — Euplectellid sponge (Hexactinellid sponge) about 1,5 m in height. N of Lifou. Depth 2116 m. Dive 02;

observer M. Roux.

7. — Euplectelle (Eponge hexactinellide) d'environ 1,5 m de haut. N. de Lifou. Profondeur 2116 m. Plongee 02;

observateur M. Roux.

Source MNHN. Paris

Source . MNHN, Paris

44

M. ROUX

FIGURE 25

(from Gaillard, 1991a, courtesy of the author)

'■ ~ Two heavily eroded beds at the top of a cliff. The cavities are in part ancient burrows, but may also be recent

borings, testifying to active bioerosion. Hemipelagic semi-indurated calccreous ooze. Thio slope

Depth 1871 m. Dive 17; observer C. Gaillard. '

1. — Deux bancs en sommet de relief soumis a une intense erosion. Les cavites sont pro parte des anciens terriers

ma|s peut-etre dgalcment pro parte des perforations rccentes temoignant d'une biocorrosion active. Boue calcaire

hemipelagiquc semi-indurec. Pente de Thio. Profondeur 1871 m. Plongde 17; observateur C. Gaillard.

2' TT- RoJ CLhtf showin8 a dense sponge community and traces of bioerosion. Santal Bay. Depth 376 m

Dive 09; observer J. Vacelet. f

2. — Paroi rocheuse montrant un peuplement dense d'eponges incrustees dans la roche et des traces de

biocorrosion. Baie du Santal. Profondeur: 376 m. Plongee 09; observateur J. Vacelet.

3. — Part of a high clilf (around 20m) with well stratified semi-indurated calcareous oozes. Note the abundance of

biogenic cavities on vertical walls. Slope off Thio. Depth 1886 m. Dive 17; observer C. Gaillard.

3' ^IUr?Ktie!e d’T falaisex iT,P°rtante (environ 20m> dans des b°^ calcaires semi-indurees bicn stratifies.

Vi oof abotid^ce des cavitds d onginc biologtque sur les parois verticales. Pente au large de Thio. Profondeur

1886 m. Plongee 17; observateur C. Gaillard.

4' se"1':,ndurated blocks- se,,lcd on hemipelagic muds. Note the numerous biogenic cavities on

Slone o ff” Th i o'* h ^ ifC ! 70 a” a,,!,ched *° lhc smal,cr one- Echinoids (?) trails are visible on the muddy area.

Slope off Thio. Depth; 1726 m. Dive 13; observer M. Segonzac.

4' 7oHPTneSooindUr6S fe°Ule> iS°ldS aU SCin des boues hcmipelagiques. No.er les nombreuses cavi.ds

ImSSlTi qU 1 S rKen,crmen' el 1 aclinie flx6c sur !e plus petit d'entre eux. Des pistes d'dchinides (?)

M Segonzac a Z°ne 0UeUSC' Pen’e au large dc Thi0- Profondeur; 1726 m. Plongee 13; observateur

5. — A crab at the entrance of its burrow. S of Santal Bay. Depth 603 m. Dive 09; observer J Vacelet

5' 7 VrAacELdETVant 1 entr6e de S°" ,Crrier- S de 13 baie dU Sanlal- Profondeur 603 m- Ploagde 09; observateur

6.

6.

Gullev^ff ^ 'wSi"8 ^ Tuai'™nawa a* 'he end of its track. Megaripple covered with ripple-marks.

Gulley ofl SW of Isle of Pines. Depth 596 m. Dive 22; observer M. Roux.

Oursin irregulicr du genre Taimanawa a l'extremitc de

au SW de Die des Pins. Profondeur 596 m. Plongee 22;

sa piste. M£garide couverlc de ripple marks. Gouttierc

observateur; M. Roux.

Source MNHN. Paris

Source : MNHN, Paris

46

M. ROUX

FIGURE 26

(from C. Gaillard, 1991a, courtesy of the author)

1. Superficial muddy sediment wholly bioturbated by holothurians (aproximate width 10 cm). North-Lifou

slope. Depth 2459 m. Dive 3; observer M. Rio.

1. — Sediment superficiel boueux entierement bioturbe par des holothuries (Iargeur des pistes 10 cm environ)

Pente N. de Lifou. Profondeur 2459 m. Plongee 03; observatcur M. Rio.

2. — Tumulus encircled by burrows (approximate total diameter 30 cm). Slope off Thio. Depth 1851 m Dive 17-

observer C. Gaillard.

2. — Tumulus entoure de terriers (diametre total 30 cm environ). Pente de Thio. Profondeur 1851 m Plonede 17-

observateur C. Gaillard.

3. — Holothurian trail (approximate width 6-10 cm). Slope off Thio. Depth 1663 m. Dive 13; observer

M. SEGONZAC.

3. — Piste d'holothurie (Iargeur approximative 6-10 cm). Talus de Thio. Profondeur 1663 m Plongee 13-

observateur M. Segonzac. ' b

4. — Circle of holes (approximate diameter 30 cm). Slope off Thio. Depth 2032 m. Dive 17- observer

C. Gaillard.

4. — Terriers en cerclc (diametre 30 cm environ). Talus de Thio. Profondeur 2032 m. Plongee 17; observateur

C. Gaillard.

5. — Straight groove (approximate lengih 50 cm). Slope off Thio. Depth 1865 m. Dive 17- observer

C. Gaillard.

5. — Souille rectiligne (longueur environ 50 cm). Talus de Thio. Profondeur 1865 m. Plongee 17- observateur C

Gaillard.

6’ ~ PC lrace ichnofacies. Hemipelagic calcareous ooze. Slope off Thio. Depth 1798 m. Dive 17- observer

c. Gaillard. ’

6. — Ichnofacies a traces FC. Boues calcaires hemipelagiques. Talus au large de Thio. Profondeur: 1798 m

Plongee 17; observateur C. Gaillard.

7. — Cluster of lumuli (approximate diamcier of one tumulus 10 to 20 cm). Slope off Thio. Depth 1693 m

Dive 13; observer M. SEGONZAC.

7irUloU,S 8r0.upfl (d'am^tre de chaque lumulus 10 a 20 cm environ). Pente au large de Thio. Profondeur

1693 m. Plongee 13; observateur M. Segonzac.

8 Tomm) FC <approxima,e lolaI len8th 2°-30 cm>- SW slope of New Caledonia (Biogeocal, CB 203, depth

8. — Trace FC Oongueur hors-tout 20 a 30 cm environ). Pente SW Nouvelle-Calddonie (Biogeocal, CB 203,

prorondeur 2000 m). ’ ’

(approximate ien8,h °f ,he ^ i5-2° cm>- ^ sw si°p<= »f

9- ^‘^TSoSoct1 ?B ““ COn,S I5‘20 Cm Cnv,r0n)' BaS dC la

Source . MNHN. Paris

Source : MNHN, Paris

Source : MNHN. Paris

_TATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RfiSULTATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULTATS DES CA

Protozoa Xenophyophorea Granuloreticulosa :

Psammina zonaria sp. nov. from the West Pacific

and some aspects of the growth of xenophyophores

Ole Secher TENDAL

Zoological Museum, University of Copenhagen

Universitetsparken 15, DK-2100 Copenhagen 0

Denmark

ABSTRACT

Psammina zonaria sp. nov. is the first member of this genus described from bathyal depths of the West Pacific. It is

characterized by strong compartmentalization of the interior space and strictly directional growth. It appears to live

attached to hard substrates, with the stiff but fragile test sticking up in the near-bottom water layer.

RESUME

Protozoa, Xenophyophorea Granuloreticulosa : Psammina zonaria sp. nov. du Pacifique ouest.

Considerations sur quelques aspects de la croissance des xenophyophores

Psammina zonaria sp. nov. est la premiere espfece du genre Psammina decrite des profondeurs bathyales du Pacifique

Ouest. Elle se caracterise par un fort compartimentage de l'espace interieur et une croissance strictement directionnelle.

Elle semble vivre fixee a des substrats durs, avec son test, rigide mais fragile, dresse dans la couche d'eau se trouvant prfes

du fond.

INTRODUCTION

Xenophyophores are macro- and megafauna-sized agglutinating rhizopods, mainly found in the deep sea

(Tendal, 1972, 1989). Although the present class Xenophyophorea was erected as an order, Xenophyophorida, of

the Rhizopodea early in the century (Schulze, 1912), it is only over the last two decades that its wide

geographical occurrence and ecological significance have become acknowledged beyond a narrow circle of deep-sea

workers. The generalities, including terminology, are now accessible in larger textbooks (Marshall, 1979,

Tendal, O. S„ 1994. — Protozoa Xenophyophorea Granuloreticulosa : Psammina zonaria sp. nov. from the West

Pacific, and some aspects of the growth of xenophyophores. In : A. CROSNIER (ed.). Resultats des Campagnes

MUSORSTOM, Volume 12. Mim. Mus. naln. Hist. nat.. 161 : 49-54. Paris ISBN 2-85653-212-8.

50

O. S. TENDAL

PAGE, 1982; TENDAL. 1989; Gage & Tyler, 1991; Gooday & Tendal, in press). The ecology of the group

has attracted special interest in recent years (Gooday et al., 1993; Levin, 1991; Levin & Thomas 1988- Levin

& Gooday. 1992; Riemann et al., 1993; Tendal, 1985).

About 50 species of xenophyophores are formally named, and quite a few await formal description. Because

most species are known from just a few records, and many only as fragments, any new find has the potential to

bring substantial new information about the group.

The material secured during the Musorstom 7 Expedition was collected from bathyal depths on the flat top of

a seamount in the central West Pacific, north of the Fiji Islands, between the Combe and the Bayonnaise banks

(Richer de Forges & Menou. 1993); this is the first record of a *cnophyophorc from that region. It also adds to

our knowledge of the growth pattern of xenophyophores in general and the basic organization of members of the

genus Psamnvna in particular.

MLIHUD5

The samples were preserved in alcohol shortly after recovery.

analvzer^EDSl cnnnprf^dCt°nStitUen1^ ^ ^ S) WaS performed with an energy-dispersive X-ray spectrographic

analyzer (EDS) connected to a scanning electron microscope (CamScan), which was run at 15 kV.

.. V* .h0l0,>!Pf/nd one Paratype are in the Museum national d'Histoire naturelle, Paris, and two paratypes are in

the Zoological Museum, University of Copenhagen. 1

SYSTEMATICS

Family PSAMMfNIDAE Haeckel, 1889

Genus PSAMMINA Haeckel, 1889

Psammina zonaria sp. nov.

Fig. 1-2

-WalliS and Fu,una Islands Musorstom 7 : St. DW 620, 12°34 4’S 178°11 O'W

1280 m, clay, 28 May 1992. Waren dredge : Fourteen specimens and large fragments. ’ * 110 W’

Ud S 3°mm Shfe* ^ SPa,Ula‘e Psammina whh even increase in width from narrow to broad end.

IhP ° cl m white r tef space conspicuously compartmentalized by bars running across the whole width of

the test. Colour white, consistency hard and brittle. Xenophyae planktonic foraminiferal shells of all sizes.

Type Material. - Two fragments in the sample fit together and form a large entire specimen selected as the

489UvT if is sDaHiL fe^T^ ‘ol M“f na,i°na' d'His,oire naIurelle- Paris (registration number MNHN-

489UV). It is spatulate in form. 26 mm (16 + 10 mm) long, 13 mm wide at one end and 4 mm a. the other A

SLena S° T1 ,here/MNHN-490UV). Two paratypes in the Zoological Museum. University of

Copenhagen, are without registrauon number. y

DESCR'PTIGN - There are three other nearly complete specimens in the sample, measuring in length and

greatest and smallest width, 15, 1 1. and 4 mm. 12, 9. and 4 mm and 10, 7, and 5 mm. respectively8 The Skness

Test : The tubular part is in all cases broken at the narrow end. As the tubular end gradually widens the test

xenoplfyac wltha s^cdeK^nat,ainin8 ** CharaC,eris,is,ic Psammina ^nization of two parallel plates of

Source MNHN. Paris

PROTOZOA XENOPHYOPHOREA : PSAMMINA ZONARIA

51

Both outer surfaces appear smooth apart from protruding xenophyae, but when examined more closely in

oblique light it is seen that they are both slightly undulating, with about 3 mm from one crest to the next. The

crests of the two test sides arc arranged symmetrically. The test edges are rounded. Both test surfaces and the edges

have numerous small openings, < 0.1 mm in diameter, which are probably the passages for pseudopodia, although

some of them may be places where small xenophyae have fallen out. A number of larger holes, 0.5-1 mm in

diameter, to the interior are merely damages. Granellare and stercomare branches are visible in some cases along

the broad end. protruding among the xenophyae; they are not seen along the sides.

Fig. 1 . — A : The holotype of Psammina zonaria sp. nov. — B : A fragment of Psammina zonaria sp. nov. One side is

dissected away to show the interior compartmentalization. Scale in mm.

Xenophyae are planktonic foraminifer shells of all sizes, packed with the smaller ones well cemented as an

infilling around the large ones.

The space between the plates, is 0.5-1 mm wide. The inner sides of the plates are connected by long solid, well

cemented bars of xenophyae running across the whole width of the test and subdividing the interior space into long

narrow, 1-2 mm wide compartments (Fig. 1 B). The bars are 0.5-1 mm wide, and the xenophyae are of the same

kind as in the plates. The bars lack larger openings, but are occasionally crossed by branches of granellare and

stercomare passing through small spaces between the xenophyae.

Granellare : The strings of granellare are sparsely developed and seem mostly to occur in the widest end.

immediately under the test plates. The dichotomously dividing yellow-white branches are 100-120 pm in diameter;

anastomoses are not seen. The plasma contains enormous numbers of granellac. up to 5 pm long. Their size,

form, colour and distribution are as in other xenophyophores, and that the main constituent is BaS04 is confirmed

by EDS analysis (Fig. 2).

Stercomare : The strings of stercomare are very strongly developed and fill the compartments completely

(Fig. 1 B). The dark brown or black branches are 100-200 pm in diameter and anastomose in a tight network with

open spaces about 0.1 mm in diameter. The covering membrane is thin but solid and slightly shiny. Rounded

xanthosomes up to 12 pm in diameter are commonly seen among the masses of stercomata.

DISCUSSION

Remarks on Taxonomy. — Because of the basic test construction, (two well defined plates of xenophyae

connected by pillars or bars) the species is placed in the genus Psammina Haeckel. 1889. Six other species arc

presently referred to the genus. Superficially, a broken specimen of P . zonaria can be mistaken for P. globigerina

52

0. S. TENDAL

Haeckel, 1889, or maybe P . plakina Haeckel. 1889, a poorly known species (Tendal, 1972 : 33), but the pro¬

nounced compartmentalization of the inner test space is a clear difference from these species. Large specimens of

P. globigerina may have irregularly distributed bar-like pillars between the plates, but these do not compart¬

mentalize the interior space. The only other known species with well defined bars is P. sabulosa Gooday &

Tendal, 1988. but here the bars are regularly perforated by large openings, and the xenophyae are well sorted

sandgrains.

Growth. — The compartmentalization reflects a distinct growth pattern showing a striking resemblance to

that ot a number of other xenophyophore species, viz Psammina sabulosa Gooday & Tendal, 1988.

Stannophyllum zonarium Haeckel, 1889. 5. globigerinum Haeckel, 1889, 5. granularium Tendal, 1972

S.fragihs Tendal- 1972 and S. flustraceum Haeckel, 1889. The explanation might be found in the very important

phenomenon of episodic growth recently discovered in the xenophyophore Reticulammina labyrinthica from the

Madeira abyssal plain in the Atlantic (Gooday el. al„ 1993).

P. zonaria has directed growth, the narrow part being the oldest, the wide part the youngest. This can be seen

from the distribution of granellare which are only present in the youngest part of the test. The tubular oldest part

can be considered a juvenile stage; thus, P. zonaria is a new example within the xenophyophores of young speci¬

mens having another body form than the older ones (Tendal, 1972 : 80). The existence of strong morphological

(Inferences between two parts of the same specimen implies the risk that an isolated fragment is attributed a wrong

taxonomic placement, a phenomenon also evident in other xenophyophores (Gooday, 1991 : 210).

n ~JtT&SSammina species have t*6" found on ,he surface in box cores, 0" soft sediment

( L in & THOMAS, 1988); they may initially have started to grow on some small hard object. Further, there are

many photographs from deep-sea bottoms showing supposed Psammina specimens on the sediment surface

PROTOZOA XENOPHYOPHOREA : PSAMMINA ZONARIA

53

sometimes among or maybe on manganese nodules (Tilot, 1992, and unpubl.; Tendal, unpubl.). Accordingly,

the main part of the body of at least some Psammina species protrudes into the water above the bottom. In

P. zonaria all the tubular parts are open at one end, have the same dimensions, and are still filled with stercomare.

This suggests that they do not represent an area that gradually dies off and decomposes; it is more likely that they

were originally fixed to a hard substrate, the brittle test breaking at the weakest point, probably just above the

base. All evidence taken together thus suggests that P. zonaria lives upright on some solid substrate.

General Ecology. — The sampling locality lies close to the summit of a seamount (Richer de Forges &

Menou, 1993). Psammina zonaria was found in a sample taken with a War6n dredge, while four other samples

from the same area, three of them taken with a fish trawl and the last with an epibenthic sledge, yielded no

xenophyophores, perhaps because these fragile organisms were fragmented beyond recognition. It is difficult to

conclude anything on this, but it should be pointed out that xenophyophores are abundant near the summits of

many eastern Pacific seamounts, probably because intensification of currents in these places leads to enhanced flux

of suspended food particles (Levin & Thomas, 1988).

ACKNOWLEDGEMENTS

I thank P. BOUCHET, Museum national d'Histoire naturelle, Paris, and A. CROSNIER, Institut fran^ais de

Recherche scientifique pour le D6veloppement en Cooperation (ORSTOM), for providing the material, for

information, and for help with the French summary; G. Bohrmann, GEOMAR, Kiel, for the EDS analysis;

A. J. GOODAY, IOSDL, Wormley, and F. Riemann, AWI, Bremerhaven, for valuable suggestions improving

the manuscript; G. Brovad and M. E. PETERSEN, both Zoological Museum, University of Copenhagen, for the

photographs and the linguistic correction of the manuscript, respectively.

REFERENCES

Gage, J.D. & TYLER, P.A., 1991. — Deep-Sea Biology. Cambridge University Press, Cambridge, 504 pp.

Gooday, A.J., 1991. - Xenophyophores (Protista, Rhizopoda) in box-core samples from the abyssal northeast Atlantic

Ocean (BIOTRANS area): their taxonomy, morphology, and ecology. J.foramin. Res., 21: 197-212.

Gooday, A.J. & Tendal, O.S., 1988. — New xenophyophores (Protista) from the bathyal and abyssal north-east

Atlantic Ocean. J. nat. Hist., 22 : 413-434.

Gooday, A.J., Bett, B.J. & Pratt. D.N., 1993. — Direct observation of episodic growth in an abyssal xenophyophore

(Protista). Deep-Sea Res., 40 : 2131-2143.

Gooday, A.J. & Tendal, O.S., (in press). — Phylum Granuloreticulosea, class Xenophyophorea. In : Illustrated guide to

the Protozoa. Allen Press, Lawrence, Kansas.

Levin, L.A., 1991. — Interactions between metazoans and large agglutinating protozoans : Implications for the

community structure of deep-sea benthos. Am. Zool., 31 : 886-900.

Levin, L.A. & Thomas, C.L., 1988. —The ecology of xenophyophores (Protista) on eastern Pacific seamounts.

Deep Sea Res., 35 : 2003-2027.

Levin, L.A. & Gooday, A.J., 1992. — Possible roles for xenophyophores in deep-sea carbon cycling. P. 93-104. In :

T. Rowe & V. Pariente (eds). Deep-sea food chains and the global carbon cycle. Kluwer Academic Publishers,

The Netherlands.

Marshall, N.B., 1979. — Developments in deep-sea biology. Blandford Press, Poole, Dorset, 566 pp.

Page, F.C., 1982. —Xenophyophorea. P. 525-526. In : S.P. Parker (ed.), Synopsis and classification of living

organisms. I. McGraw-Hill Book Company, New York, 1166 pp.

Richer de Forges, B. & Menou, J.-L., 1993. — La campagne MUSORSTOM 7 dans la zone 6conomique des ties Wallis et

Futuna. Compte rendu et liste des stations. In : A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM. Volume 10.

Mem. Mus. natn. Hist, nat., 156 : 9-25.

54

O. S. TENDAL

RlE^mNthe''wSlSnLS0'S' GlNGELE\F;X" 1993' Reticulammina anlarclica nov. spec. (Xenophyophora. Protista)

irom the Weddell Sea. and aspects of the nutrition of xenophyophores. Polar Biol., 13 : 543-547.

Tendal, O.S., 1972. — A Monograph of the Xenophyophoria (Rhizopodea, Protozoa). Galathea Rep., 12 : 7-99.

T”-pS5*3K“,sT“' in Ihe »f **■« <Mollu.„,

TTrCHAPM^9^;rHPhyHlr,X?ThrPh0ra- V35’138' ,n ■ L- MaRGULIS. J.O. Corliss, M. Melkon.an &

D.J. Chapman (eds). Handbook of the Protoctista. Jones and Bartlett Publishers. Boston, 914 pp.

TlLOl-, V.. 1992. — La structure des assemblages megabenthiques dune province a nodules polymdtallioues de l'ocean

aci ique tropical Est. IFREMER, Brest (Th5se de doctoral en science de l’Universitd de Bretagne Occidental), 380 pp.

Source : MNHN, Paris

.TATS DES CAMPAGNES MUSORSTOM VOLUME 12 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES C/

Bryozoa : Living and fossil species

of the catenicellid subfamilies

Ditaxiporinae Stach and Vasignyellinae nov.

Dennis P. GORDON

New Zealand Oceanographic Institute

National Institute of Water & Atmospheric Research (NIWA)

P.O. Box 14 901 Kilbirnie

Wellington, New Zealand

&

Giampietro BRAGA

Dipartimento di Geologia, Paleontologia e Geofisica

Universitci di Padova, via Giotto 1

35137 Padova, Italia

ABSTRACT

The discovery of living species of the predominantly Tertiary catenicellid subfamily Ditaxiporinae on the Norfolk

Ridge has necessitated a revision of the subfamily, which is characterised by biseriate multizooidal segments. The type

species of the genera of Ditaxiporinae and of the related family Ditaxiporinidae were examined by scanning electron

microscopy, leading to the recognition of six genera (two new) and 18 species (four new) and the incorporation of the

Ditaxiporinidae into the Ditaxiporinae. The earliest occurring species is Caberoides rockallensis sp. nov. in the late

Paleocene of the North Atlantic. There are only two living species - Bryosartor sulilis gen. et sp. nov. and Plagiopora

recens Gordon, both on the northern Norfolk Ridge. A new monotypic genus, Ahcheethamia, is introduced for Caberoides

corniculatus Cheetham from the British Eocene. With the exception of two species from North America, the subfamily is

clustered in two centres of diversity - northwestern Europe and Australasia, the latter including Caberoides miranda sp.

nov. and Plagiopora alma sp. nov., both newly recorded from the Eocene of New Zealand. Thus a Tethyan distribution of

the subfamily was achieved relatively early in the Paleogene.

Just as in other catenicellids, there seem to have been parallel trends in the Ditaxiporinae in the diversification ol the

frontal shield from a spinocyst to a perforated gymnocyst on the one hand and with cryptocystal elements (derived from

expanded shallow pore-chambers) on the other. A unique development is indicated by the genus Vasignyella. Hitherto

Gordon, D. P. & Braga, G., 1994. — Bryozoa : Living and fossil species of the catenicellid subfamilies

Ditaxiporinae Stach and Vasignyellinae nov. In : A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM, Volume 12.

M£m. Mus. naln. Hist, not., 161 : 55-85. Paris ISBN 2-85653-212-8.

56

D. P. GORDON & G. BRAGA

included in the family Savignyellidae, Vasignyella appears to have been derived from Ditaxiporina or a common ancestor

by reduction to unizooidal segments and the loss of ovicells. A new subfamily of Catenicellidae, Vasignyellinae, is

established for this genus.

RESUME

Bryozoa : Especes vivantes et fossiles des sous-familles catenicellides : Ditaxiporinae Stach

et Vasignyellinae nov.

La decouverte, sur la ride de Norfolk, d'espfeces vivantes appartenant a la sous-famille catenicellide des Ditaxiporinae,

surtout repandue au Tertiaire, a conduit & la revision dc cette sous-famille caractdrisee par des segments h nombreux

zooides bisdrids. Les especes type des genre des Ditaxiporinae et de la famille des Ditaxiporinidae ont dtd examinees au

MEB. Ceci a conduit a reconnaltre six genres (dont deux nouvcaux) et 17 especes (dont trois nouvelles) et ik incorporer les

Ditaxiporinidae dans les Ditaxiporinae. L'espece la plus ancienne est Caberoides rockallensis sp. nov. du Paldocdne

superieur de 1'Atlantique nord. Scules deux especes actuelles existent, Bryosartor sutilis gen. et sp. nov. et Plagiopora

recens Gordon, toutes deux sur la partie nord de la ride de Norfolk. Un nouveau genre monotypique, Ahcheethamia, est cree

pour Caberoides corniculatus Cheetham de l'Eocene d'Angleterre. A l'exception de deux especes trouvees en Amerique du

Nord, la sous-famille sc regroupe suivant deux centres de diversification : le Nord-Ouest de l'Europe et l'Australasie, le

second incluant Caberoides miranda sp. nov. et Plagiopora alma, r6cemment trouvees toutes deux dans l'Eocene de

Nouvelle-Zelande. Ainsi une distribution tethysiennc de la sous-famille a dte realisde relativement tot dans le Palaeogene.

De meme que chez les autres catenicellides, il semble y avoir eu des tendances paralleles chez les Ditaxiporinae dans la

diversification du bouclier frontal, depuis un spinocyste jusqu'S un gymnocyste perfore d'une part et avec des elements

cryptocystiques (ddrivds de chambres porales peu profondes, dilatees) d'autre part. Le genre Vasignyella prdsente un

developpement unique. Indus jusqu'a present dans la famille des Savignyellidae, Vasignyella semble derivd de

Ditaxiporina ou d'un ancetre commun, par reduction a des segments & un seul zooide et la perte d'ovicelles. Une nouvelle

sous-famille de Catenicellidae, celle des Vasignyellinae, est crede pour ce genre.

INTRODUCTION

Among the Bryozoa collected by the Musorstom cruises in New Caledonian waters were some taxa of the

cheilostomate superfamily Catenicelloidea that were either previously known only from the Tertiary of Victoria,

Australia ( Chelidozoum ), or which have their earliest representation there ( Petalosiegus , Petalostegidae, and

Ditaxiporinae) (Gordon & d’Hondt, 1991; Gordon, 1993). The latter taxon, Ditaxiporinae, is a subfamily of

the Catenicellidae, a fairly widely distributed family with its greatest abundance in the Australasian region. Until

recently, when a new species attributed to Plagiopora was discovered on the Norfolk Ridge north of Norfolk Island

(Gordon, 1989a), the Ditaxiporinae was thought to be extinct. Now a new genus, with a major ancestral feature,

has been discovered in the Musorstom collections from New Caledonia. Its discovery provides a much-needed

opportunity to revise and discuss the relationships of the whole subfamily Ditaxiporinae.

The Catenicellidae Busk, 1852 is a very distinctive and well-circumscribed family. The bushy colonial

morphology is characteristic, with branches comprising long chains of uni- to multizooidal segments separated by

chitinous joints. This characteristic morphology fortunately permits the classification of some very disparate

zooidal morphologies in the one family (Banta & Wass, 1979). It is this varied zooidal morphology that is the

most remarkable feature of the family, comprising a range of frontal-shield types that elsewhere in the order

Cheilostomatida might each have significance at the family level. Nevertheless, within the Catenicellidae it is

possible to determine ancestral and derived characters and consequent polarities, permitting the construction of

morphological lineages. Thus, the presence of a field of costal spines is an ancestral character, a lack of spines and

a concomitant extensive gymnocyst a derived character. Similarly, any kind of cryptocystal feature is derived.

Appropriately, the oldest-known catenicellid, from the Maastrichtian (Upper Cretaceous) of Jamaica (Cheetham,

pers. comm, in Banta & Wass, 1979 ; 22) resembles Costaticella, a spinocystal genus, but by the early Eocene

in the Indian Ocean, Catenicella, with an advanced morphology, is already present (Labracherie & SlGAL, 1976,

as Vittaticella).

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

57

The range of zooidal morphologies in the Catenicellidae led to the recognition of different groupings within the

family. Accordingly, when Busk (1852a) introduced the family, with two included genera, Catenicella and

Calpidium , he recognised three groupings within Catenicella - fenestratae, vittatae, and inermes, but not as formal

taxa. He continued to use these groupings (Busk, 1852b, 1884), only changing 'inermes' to 'simplices' in the

former work but abandoning the family name in the latter. In subsequent decades various authors recogised several

new genera within Busk's groupings. Stach (1933, 1934, 1935a) was the first to recognise subfamilies within

the Catenicellidae, basing them primarily on the position of the ovicell within the segment. He named five

subfamilies - Vittaticellinae (Stach, 1933), Scuticellinae (Stach, 1934), and Catenicellinae, Comuticellinae, and

Ditaxiporinae (Stach, 1935a). The distinction between the Vittaticellinae and Catenicellinae was artificial and

based on a misunderstanding, since Vittaticella is a junior synonym of Catenicella (Gordon, 1984), and work by

Wass & Banta (1981) on catenicellid ovicell complexes has shown that the comuticelline grouping may also be

included in the Catenicellinae. Thus Banta & Wass (1979) and Wass & Banta (1981) accepted the distinction

between the Catenicellinae, in which the ovicelled segment does not comprise the terminal segment of a branch,

and the Scuticellinae, in which it does.

The remaining subfamily, Ditaxiporinae, was segregated primarily on the basis of the characteristic biseriate

multizooidal segments, found in the fossil genera Ditaxipora MacGillivray, 1895, and Ditaxiporina Stach, 1935a.

The chief features cited by Stach (1935a) as distinguishing these two genera were the longitudinal band on the

frontal area in the former and the evenly porous shield lacking a band in Ihe latter. Stach (1935b) noted the

occurrence of multizooidal forms in ihe European and North American Paleogene. Because these were then the

oldest-known catenicellids, he regarded them as archaic. [In the event, as mentioned above, the earliest-known

catenicellid, from the Maastrichtian (mentioned above), resembles Costaticella , which makes the Scuticellinae the

earliest-occurring subfamily.] Cheetham (1963) discussed the differences between Ditaxipora and Ditaxiporina ,

concluding that the evenly porous frontal shield and other (minor) features of the latter must preclude a close

relationship. He therefore established a new family, Ditaxiporinidae, for Ditaxiporina and Caberoides Canu. 1908

(Cheetham, 1963). Notwithstanding having segregated the Ditaxiporinidae, he nevertheless concluded "Future

investigations will probably verify the proximity of this family to the basal stock from which the Vittaticellidae

[i.e., Catenicellidae] have sprung. ... However, the Vittaticellid roots should be sought amongst the archaic groups

of Cheilostomata in Lower and Middle Eocene beds, not the more specialized forms of the Upper Eocene and

Oligocene" (Cheetham, 1963 : 489). However, at that time, pre-Miocene scuticellines and catenicellines had not

yet been discovered.

Thus, by 1966, only the three biseriate-multizooidal genera. Ditaxipora, Ditaxiporina, and Caberoides,

segregated in two families, had been recognised. Subsequently, the discovery of a new species of Plagiopora

MacGillivray at 831 m depth northwest of Norfolk Island (Gordon, 1989a) indicated that this multizooidal genus

ought also to be included in the Ditaxiporinae. Previously known only from a single Middle Miocene species, this

genus had been included by MacGillivray (1895) in the now-abandoned (and unrelated) family Lepraliidae.

The recognition of a new, living, genus of Ditaxiporinae in the MUSORSTOM samples from the northern

Norfolk Ridge, plus the discovery of new ditaxiporine species in the New Zealand Eocene, prompted a

reconsideration of all the multizooidal genera, especially based on scanning-electron-microscopic examination of

type species.

LIST OF STATIONS

New Caledonia.

Biocal : station DW 70, 4.9.85, 23°24.70’ S. 167°53.65' E, 965 m : Bryosartor sutilis (+ Bifaxaria modesta,

Diplonotos serratus, Petalostegus scopulus).

Musorstom 4 : station CP 214. 28.9.85, 22°53.80' S, 167°13.9' E, 425-440 m : Bryosartor sutilis.

BIOGEOCAL : station DW 313, 2.5.87, 20°58.95' S, 166°59.04' E, 1640-1600 m : Bryosartor sutilis (+ Bifaxaria

me no rah).

58 D. P. GORDON & G. BRAGA

FlG. 1 a-f. — Bryosarlor sutilis gen. et sp. nov. : a, arrangement of zooids in the multizooidal segment (x 42) ;

b. $ zooid with ovicell (x 73) ; c. autozooid. showing the narrow costal field (x 144) ; d, proximal zooids of a

young segment (x 86) ; e, dorsal view of a developing segment (x 58) ; f, interior view of costae from one side of a

costal field and the irregular suture-line at their tips (x 305). (All from BlOCAL Stn DW 70.)

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

59

SYSTEMATIC ACCOUNT

Suborder ASCOPHORINA Levinsen, 1909

Infraorder CRIBRIOMORPHA Harmer, 1926

Superfamily CATENICELLOIDEA Busk, 1852a

Family CATENICELLIDAE Busk, 1852a

Subfamily DITAXIPORINAE Slach. 1935a

Genus BRYOSARTOK nov.

DIAGNOSIS. — Multizooidal, biseriate. Frontal shield includes a narrow costal field with several pairs of

vestigial costae and a few extracostal windows; the adjacent gymnocyst with large shallow pore-chambers, 1 axial

(1-2 on female zooids), 2 marginal, and another distal to the outer avicularium. Only a single avicularium on

autozooids, at the outer distolateral corner; paired avicularia in female zooids. Ovicell large, with a large exposure

of granular endooecium surrounded by smooth ectooecium. Dorsal zooidal surface smooth, lacking pore-chambers.

Range. — Holocene.

TYPE species. — Bryosartor sutilis sp. nov.

Bryosartor sulilis sp. nov.

Fig. 1 a-f

Material EXAMINED. — New Caledonia. BIOCAL : stn DW 70, 965 m.

Musorstom 4 : stn CP 214, 425-440 m.

Biogeocal : stn DW 313, 1640-1600 m.

Description. — Colony erect, branching, to 31 mm high, the branches biserial with 4-6 zooids per fully

formed segment; maximum segment length 4.74 mm and width 0.66 mm. All joints are single - at bifurcations

the 2 distal zooids are angled away from each other, producing widely separated branches, and there is only a single

zooid at the proximal end of each segment, which buds 2 daughter zooids to generate biseriality. Zooids vary in

length according to position in segment - the shortest zooids (-0.34 mm) occur at the proximal end, the longest

zooids (-0.64 mm) occur immediately distal to female zooids; zooidal width -0.34 mm. The frontal side of each

zooid has several components - centrally, there is a narrow, adaxially curving, costal field of about 11-19 short

flattened spines with a median suture running through the field to the orifice; the distalmost costae are very broad;

outside the costal field are 3-4 gymnocystal foramina. Pore-chambers are large and shallow, with 1 on the axial

side and 3 on the outer, marginal, side of each zooid - the axial one is the largest, extending from just distolateral

to the orifice, past an avicularium (if there is one on that side), to a position adjacent to the middle of the costal

field; laterally, there is a chamber behind the outer distolateral avicularium and two larger chambers proximal to

the avicularium. Zooidal orifice proportionately large, suboval, with a pair of tiny condyles about one-third the

distance from the proximal rim. Avicularia 1-2, 1 at the outer distolateral comer of every zooid. with complete

cross-bar and acute rostrum; a similar, less-acute, avicularium occurs opposite, adjacent to the distalmost costae, in

ovicelled zooids and occasionally in other zooids. Ovicell large, about the size of the maternal zooid, occupying

the complete frontal surface of the distal zooid, the ovicellular surface mostly granular endooecium with 'flaps' of

smooth ectooecium distally and proximally; the orifice of fertile zooids generally a little longer than other orilices.

Dorsal surface of branch segment with grooves marking the boundaries between zooids. especially along the axis.

60

D. P. GORDON & G. BRAGA

otherwise fairly smooth with no ridges. Colony anchored to substratum by a number of rhizoids descending the

proximal parts of the colony; rhizoids issue from the proximal parts of zooids dorsally. Ancestrula small,

0.32 mm long and 0.23 mm wide including the distolatcral corners; resembling later zooids but the proximal half

evenly narrow and claviform, supported by a rhizoid issuing from the proximal end. The postancestrular and some

later segments unizooidal.

TYPES. — Hololype : a colony from BlOCAL Stn DW 70, 965 m, MNHN-Bry 19934.

Paratypes : fragments of mature colonies from the same station, and young attached colonies from Biogeocal

Stn DW 313, 1640-1600 m, MNHN-Bry 19933.

Etymology. — The genus name is partly derived from sartor (Latin), a tailor, and the species name from

sutilis (Latin), sewed together. Both names allude to the narrow costal field which gives the zooidal frontal shield a

stitched appearance.

DISTRIBUTION. — Southern New Caledonia and northern Norfolk Ridge, 425-1640 m.

Remarks. — Bryosartor differs from Ditaxiporina in several important characters, including the presence of

costae and extra-costal foramina, the number and disposition of frontal pore-chambers, and the complete lack of

dorsal pore -chambers. At present, Bryosartor is monospecific.

Genus CABEROIDES Canu. 1908

Diagnosis. — Multizooidal, biseriate. Frontal shield wilh a tiny suboral costal field with a few vestigial

costae; no extracostal windows; adjacent gymnocyst with large shallow pore-chambers - 1-2 axially and

1-2 laterally, with another behind the outer lateral avicularium. Avicularia single or paired. Ovicell large, with a

narrow exposure of endooecium surrounded by a large area of smooth ectooecium. Dorsal zooidal surface with at

least 2 pore-chambers, with one or both narrow and curved.

Range. — Thanetian (Upper Paleocene) to Priabonian (Upper Eocene).

Type SPECIES. — Caberoides canaliculata Canu. 1908.

Congeneric species. — Four other species may be included in the genus. These are :

Catenicella continua Waters, 1891 = Caberoides continua (Waters, 1891) comb, nov., Priabonian, northeast

Italy.

Caberoides grignonensis Canu, 1908, Lutetian, near Paris, France.

Vitaticcllid [j/c], n. gen. Cheetham & HOkansson, 1972 = Caberoides rockallensis sp. nov., Thanetian.

Rockall Plateau, Northeast Atlantic.

Caberoides miranda sp. nov., Ypresian, Chatham Island, New Zealand.

Caberoides canaliculata Canu, 1908

Fig. 2 a-d

Caberoides canaliculata Canu. 1908 : 88. pi. 11 [84, pi. 8], figs 11-12. — Bassler, 1953 : G212, fig. 159.4. —

Cheetham, 1966 : 81.

MATERIAL EXAMINED. — France. Museum national d'Histoire naturelle : Holotype specimen R. 53409, Institut de

PaI6ontologie, Canu Collection, Orglandes, near Paris, Lutetian.

Source MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

61

DESCRIPTION. — Segments up to 2.26 mm long and 0.97 mm wide, with up to 12 zooids per segment.

Zooids alternating, 0.29-0.38 mm long and 0.31-0.39 mm wide. Frontal shield with a central Y-shaped

gymnocystal area flanked by pore-chambers, with a small costal field between the arms of the Y. The pore-

chamber on the axial side extends from near the distolateral comer of the orifice to the proximal third of the zooid;

the marginal pore-chamber extends from the lateral avicularium to the same level; more proximally is a very

narrow pore-chamber that extends partly onto the dorsal surface. This arrangement is apparent in non-ovicelled

zooids (at the proximal end in fertile segments). Orifice with a broad shallow poster. Avicularia single, on the

lateral margin facing laterally, or double, as in ovicelled zooids. The axial avicularia occur in a series along the

Fig. 2 a-d. — Caberoides canaliculaia Canu : a, fertile segment with numerous ovicells (x 62) ; b, enlargement of the

bottom right zooid in fig. a, showing the expanded lateral pore-chambers flanking the Y -shaped gymnocyst with

small costal field (x 142) ; c, dorsal face of a segment, showing the disposition of pore-chambers (x 42) ; d, part of

fig. c, magnified (x 119). [SEMs of Canu's uncoated figured specimens from Orglandes (CaNU, 1908. pi. 8 (pi. 11),

figs 11-12), slide R53409, Institut de Pal6ontologic, Museum national d'Histoire naturelle.]

62

D. P. GORDON & G. BRAGA

centre of the segment, with each obliquely facing the zooid which bears it; both axial and lateral avicularia with

semicircular opcsiae, complete cross-bars, and acute triangular rostra; frequently the lateral avicularium is projected

beyond the rostrum. Ovicell relatively large, obliquely recumbent over much of the frontal shield of the zooid

distal to it; the ectooecium smooth with a narrow axial fenestra (widening proximally) and possibly a few scattered

pores. Dorsal surface of segment with a distinctive arrangement of pore-chambers - a long and narrow, vibraculum-

like, chamber extends from behind each lateral avicularium obliquely down and across the zooidal surface beyond

the midline of the segment: thus the oblique pore-chambers on one side of the segment overlap with those on the

other side. A somewhat semicircular pore-chamber, with several scattered septular pores, occurs in the centre of

the dorsal wall of each zooid.

Distribution. — Middle Lutetian (Middle Eocene), near Paris.

Remarks. — Canu (1908) had some difficulty in interpreting this genus, not being intimately acquainted

with the southern hemisphere catenicellids. He called the axial row of avicularia vibracula, presumably akin to

those in the branch axils of Scrupocellaria, and referred to the oblique dorsal pore-chambers (like catenicellid

'vittac') as "a system of gutters containing vibracular bristles" [our translation]. Nevertheless, he recognized the

obvious ascophorine nature of the genus and, while not then suggesting a family for his new genus, Canu &

Bassler (1929) later suggested the Reteporidae. Cheetham (1963) recognized the catenicellid affinities of

Caberoides and Dilaxiporina , meanwhile establishing a new family. Ditaxiporinidae, for these two genera. Later,

Cheetham (1966) described an apparent new species of Caberoides (C. corniculatus) from the Bartonian (upper

Middle Eocene) of Sussex, England. We have examined this species by scanning electron microscopy and conclude

that it is not congeneric with Caberoides — later in this paper, we introduce a new genus for it ( Ahcheethamia ).

Caberoides grignonensis Canu, 1908 (fig. 3 a-b) is very similar to C. canaliculata, differing in details of the

costal field and dorsal surface and in its longer and narrower segments. Another species that may be included in

Caberoides is Catenicella continua Waters, 1891 (fig. 3 c-f) from the Priabonian (late Eocene) of northeast Italy

(Braga & Barbin, 1988, as Villaiicella). Waters's (1891) attribution of this species to Catenicella was based

primarily on the presence of conspicuous lateral pore-chambers (vittae), like those of Catenicella but larger. When

MacGili.ivray (1895) established his new genus Ditaxipora, he implied that C. continua might be included in it

but this idea was not followed by later authors (e.g.. Waters, 1913; Canu & Bassler, 1917, 1920) until the

work of Stach (1935a, b), who first used the combination Ditaxipora continua. This is surprising, especially in

the case of Waters (1913 : 483), who allied the species with D. internodia when he first described continua

(Waters, 1891). In the event, C. continua must be excluded from Ditaxipora which lacks any trace of a costal

shield and in which the dorsal surface of each zooid is mostly composed of a single large pore-chamber. On the

other hand, C. continua shares the important characters of Caberoides miranda and C. rockallensis (see below), viz,

a small costal field flanked by pore-chambers, laterally facing avicularia, and a vibraculum-like dorsal pore -chamber

and may be included in the genus as a new combination, Caberoides continua. It is interesting that C. continua.

occurring later in the Paleogene (Priabonian) than the other species (Thanetian to Lutetian), has much longer

frontolateral pore-chambers that anticipate those in Ditaxipora. which may have been derived from Caberoides. It is

to be noted that, whereas the supraavicularian pore-chamber is long and vibraculum-like in most of the species, it

is the proximolateral one in C. continua that has this form. When more species become known it may be appro¬

priate to divide Caberoides into subgenera on the basis of this character.

Fig. 3 a-b. — Caberoides grignonensis Canu : a, fertile segment with numerous ovicells (x 39) ; b, part of fig. a showing

the distalmost ovicells. Note the small costal field in the upper left-hand zooid (x 120). (SEM of CaNU's uncoated

figured specimen from Grignon [Canu, 1908, pi. 8 (pi. 11), fig. 15), slide R53358, Institut de Paleontologic,

Musdum national d'Histoire naturelle.J

Fig. 3 c-f. — Caberoides continua (Waters) : c, e, autozooidal frontal shields showing the arrangement of pore-chambers,

median gymnocyst, and small costal field (x 209 ; x 360) ; d, f, dorsal surface of segment and zooids, showing

disposition of pore-chambers (x 47 ; x 128). (From Monteccio di Costozza, Italy, late Eocene ; Braga Collection,

Dipartimento di Geologia, Paleontologia e Geofisica dell'Universita di Padova.)

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

63

Source : MNHN, Paris

64

D. P. GORDON & G. BRAGA

Caberoides miranda sp. nov.

Fig. 4 a-d

MATERIAL EXAMINED. — New Zealand. From New Zealand Geological Survey (NZGS) collection GS 14765 from

the Matanginui Limestone, Ypresian (Lower Eocene), 50 m north of the mouth of Waipapa Creek, Chatham Island

(locality CH/f603), collected by H.J. CAMPBELL and P.A. MAXWELL. March 1981; three other poorly preserved non-type

specimens in a separate well on the same slide.

Description. — Segments incomplete in the present material, the largest attaining 1.20 mm long and

0.43 mm wide, comprising parts of 8 zooids. Zooids small, 0.33-0.43 mm long and ~ 0.25 mm wide, widest in

the distal half, sometimes tapering narrowly in the proximal half, the median line between zooids tracing an

irregular zigzag course along each segment. Zooidal orifice with a pair of small rounded condyles separating a

broad anter from smaller poster - the exact shape of the latter is uncertain owing to poor preservation. The frontal

area of each zooid comprises 3 components - gymnocystal and cryptocystal (of approximately equal area) and

spinocystal. The spinocyst comprises a small suboral field of ~6 tapering costae whose acute tips probably did not

fuse in life. Part, at least, of the proximal rim of the orificial sinus was defined by costal elements. Four

cryptocystal areas, occasionally 5. representing pore-chambers, occur frontally. The largest, with up to 7 septuiar

pores evident, occurs on the axial side of the orifice and costal field. In 1 or 2 zooids in a segment this pore-

chamber is shortened by the occurrence of an avicularium beside the orifice. More typically, avicularia are restricted

to the zooidal margin, where 1 occurs facing laterally; it has a semicircular opesia, complete cross-bar, and

triangular rostrum with no palate. Distal to this avicularium is a second pore-chamber (with ~2-3 pores) which

curves obliquely round and down the dorsal surface. Immediately proximal to the avicularium, and partly overhung

by it, is a third pore-chamber (with ~5 pores), which also curves round onto the dorsal surface. Near the

proximolateral margin is a fourth frontal pore-chamber (with ~5 pores), of which the larger part curves obliquely

upwards onto the dorsal surface towards the proximal end of the supraavicularian pore-chamber. Occasionally, a

small rounded fifth pore-chamber (with 3 pores) is interpolated between the subavicularian and proximolateral

chambers. Dorsally, the median boundary between zooids describes a series of curving zigzags along the segment.

Of the 3 pore-chamber areas that occur on the dorsal side of each zooid, the downcurving supraavicularian one is

notable for its superficial resemblance to a vibracular groove. Ovicell unknown.

Type. — Holotype ; the segment illustrated in figure 4 a-d, from the Matanginui Limestone (Lower Eocene),

Waipapa Creek, Chatham Island, type number BZ 161, in the collection of the Institute of Geological & Nuclear

Sciences, Lower Hutt. New Zealand.

Distribution. — Ypresian (Lower Eocene), Chatham Island, New Zealand.

Etymology. — The specific name is a Latin adjective meaning wonderful, strange, singular.

Remarks. — The Matanginui Limestone is a white, massive, bryozoan-echinoid-foraminiferan-bivalve

packstone (Wood el al. 1989). On Chatham Island it is up to 25 m thick and commonly soft. The occurrence of

the foraminiferan Asierocyclina in the lower part of the limestone on Chatham Island indicates deposition in a

shallow warm sea within the photic zone. The scarcity of shallow-water foraminifers in the middle and upper parts

of the limestone indicate middle- to outer-shelf depths, the likely provenance of C. miranda. The bryozoan

diversity in this limestone is extraordinarily rich, with abundant representation of erect branching, bilamellar,

articulated, and rod-like forms, indicative of relatively fast, nutrient-rich currents.

Fig. 4 a-d. Caberoides miranda sp. nov. : a-c, SEMs of best-preserved specimen, showing disposition of frontal pore-

chambers, gymnocyst, and costal field in relation to the orifice and distolateral avicularium (x 114 ; x 285 ; x 211) ;

d, same, dorsal view (x 120). (Holotype specimen, in collection of NZGS, Lower Hutt, from the Matanginui

Limestone, Chatham Island, New Zealand, Lower Eocene.)

Fig. 4 e- — Caberoides rockatlensis sp. nov., SEM of uncoated specimen (x 106). USNM specimen 172444 from Deep

Sea Drilling Site 117A, Rockall Plateau, North Atlantic.

Source : MNHN , Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

65

Source : MNHN, Paris

66

D. P. GORDON & G. BRAGA

It is obvious from the light-micrograph illustrations of the frontal and dorsal sides of the late Paleocene

"Vitaticellid [s/c], n. gen." of Cheetham and HAkansson (1972) that this species is very similar to C. miranda.

Their two specimens, one comprising two zooids (illustrated in dorsal view) and the other comprising four zooids

(illustrated in frontal view) have many features in common - namely the small costal field flanked by pore-

chambers, a laterally facing avicularium, a proximolateral pore-chamber curving round dorsally, and an identical

disposition of pore-chambers dorsally including the vibraculum-like chamber that originates distal to the

avicularium. Their species differs only in details, including the proportionately larger avicularium and wider rims

of the pore-chambers. We hereby name the species represented by their specimens Caberoides rockallensis sp nov

We designate the larger of the two specimens (USNM slide 172444) (fig. 4, e) as holotype , the smaller (slide

172445) as paratype. Both specimens are held in the collection of the Smithsonian Institution (National Museum

of Natural History), Washington, D.C.

Caberoides rockallensis was taken at Deep Sea Drilling Site 117A on the Rockall Plateau (57°20.17' N.

15 23.97 W, 1038 m) in the North Atlantic Ocean. On the basis of certain other taxa in the assemblage,

Cheetham and HAkansson estimated that the depth, in life, of the assemblage was not more than 60 m.

Genus DITAXIPORA MacGillivray, 1895

Diagnosis. Multizooidal, biseriate. Frontal shield lacking any trace of costae; comprising a T-shaped

median gymnocystal ridge flanked on either side by a wide pore-chamber with conspicuous septular pores in a

single longitudinal series. An additional, small, pore-chamber occurs distal to the avicularium; a similar pore-

chamber on the opposite side of the orifice may be confluent with the lateral pore-chamber. Avicularia single in

autozooids, 1-2 in ovicelled zooids. Ovicell large, with an extensive area of endooecium exposed frontally Dorsal

surface with a single large pore-chamber per zooid.

Range. Lutetian (lower Middle Eocene) to ?Messinian (uppermost Miocene).

Type species. — Calenicella inlernodia Waters, 1881.

Congeneric species. — Three other species may be included in this genus. These are :

Bactridium labiatum Canu, 1910, Bartonian, Biarritz, southwest France.

Ditaxipora luteciana Canu, 1913, Lutetian, near Bordeaux, Southwest France.

Diiaxipora pannonensis Braga in Antolini el al., 1980. Priabonian, northeast Italy.

Ditaxipora inlernodia (Waters, 1881)

Fig. 5 a-c

Calenicella inlernodia Waters, 1881 : 318, pi. 18, figs 78-79. — Waters 1883 • 430

Dl,aZZa JoTTcSsk' Ma^™AY’ l,895 : pL 2l flg' 31- 31a- - Milestone; 1898 : 21; 1904 : 191. - Stack,

^935a . 394; 1935b ; 40, figs 2-3. — Bassler, 1953 : G222, fig. 167, 6 a-b. — Brown, 1958 : 82. — Wass, 1973 ;

Slrophipora (Ditaxipora) internodia - CANU & Bassi.ER, 1929 ; 452, fig. 185 F-G.

Material EXAMINED. — Australia. Royal Melbourne Institute of Technology : specimens of Ditaxipora

inlernodia from Princetown (#507) and Fyansford (#514), Miocene, Victoria, collected by P.E. BOCK.

Description. - Segments up to 2.34 mm long and 0.47 mm wide, comprising up to 10 regularly alternating

zooids; this alternation tends to be maintained even in the most proximal zooids of a segment. Paired joint-holes

at the distal end of a segment imply branch bifurcations. Zooids 0.47-0.55 x 0.22-0.28 mm, near-rectangular with

parallel sides or evenly tapering somewhat towards the proximal end. Orifice a little longer than wide, with a pair

of condyles which separate the broad rounded poster from the anter. Frontal shield with 2 large cryptocystal areas

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

67

(pore-chambcrs), each with a linear row of septular pores along its length, separated by a T-shaped median

gymnocystal ridge. This ridge is composed of 2 longitudinal components with a median suture-line between that

runs into the cross-piece of the T. The lateral pore-chambers may curve narrowly round to the distal side of the

orifice or, more usually, they terminate adjacent to the orifice, especially at the avicularium, leaving the small

distal chambers isolated. A single avicularium per zooid, always on the outer (marginal) side of the orifice, with a

semicircular opesia, complete cross-bar, and acute triangular rostrum. Ovicell large, similar to that in Bryosutor

sutilis, with a granular endooecium bordered by smooth ectooecium, two 'flaps' of which encroach toward the

centre of the granular area. Dorsal surface with a sinuous median interzooidal groove and a single large pore-

chamber per zooid, each with linear septular pores.

Fig. 5 a-c. — Ditaxipora internodia (Waters) : a-b, frontal views of zooids (x 65 ; x 185) ; c, dorsal surface of segment

showing large pore-chambers (x 104). (From Princetown, Victoria, Miocene ; Bock Collection, Royal Melbourne

Institute of Technology.)

Distribution. — Victoria, Australia, Early Oligocene to Late Miocene.

Remarks. — On the basis of the localities cited for this species by Waters (1881, 1883), MacGillivray

(1895), Maplestone (1904), Stach (1935b), and Brown (1958), and the stratigraphic work of COCKBAIN

(1971), Wass (1973) gave the range of Ditaxipora internodia as Early Oligocene (Rupelian) to Late Miocene

(?Tortonian/Messinian).

Both Waters (1881) and Maplestone (1898) noted the occurrence of a foramen in the middle of the

gymnocystal ridge and depicted the cross-piece of the T as more oval, features not apparent in the material

68

D. P. GORDON & G. BRAGA

described by MacGillivray (1895). Maplestone (1898) and Stach (1935b) concluded that this variation is

intraspecific. One of the segments in the material kindly donated to us by Phil Bock (Royal Melbourne Institute

of Technology) does have the median foramen in several zooids. These foramina are predator boreholes. Caienicella

internodia var. angustaia Waters, 1883 merely represents older internodes of this species according to Stach

(1935b).

Ditaxipora pannonensis Braga in Antolini el ai, 1980 (fig. 6 a-c) from the Priabonian (Upper Eocene) of

northern Italy is clearly related. It has a similar T-shaped gymnocystal ridge frontally and large dorsal pore-

chambers dorsally. The ovicell is also similar, with proportionately larger proximal ectooccial 'flaps'. Ditaxipora

labiata (Canu, 1910) from the Bartonian of France is likewise congeneric - the characteristic frontal gymnocystal

ridge and large dorsal pore-chambers are typical of Ditaxipora, not Ditaxiporina , as Labracherie (1968) correctly

pointed out.

Fig. 6 a-c. — Ditaxipora pannonensis Braga in Antolini et at. : a, sterile segment (x 35) ; b, part of fig. a (x 216) ;

zoo‘<ls (x 96). (From Pannone, late Eocene, Braga Collection, Dipartimento di Geologia, Paleontologia

e Geofisica dell'Universita di Padova.)

Genus PLAGIOPORA MacGillivray, 1895

Diagnosis. Multizooidal, biseriate. Zooidal orifice slightly asymmetrical and at an angle to the segmental

axis. Frontal shield lacking costae; comprising an obliquely set narrow gymnocystal band suborally that separates

several broad, shallow, pore-chambers (cryptocystal fields) tiny sparse septular pores; the gymnocystal band with

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPOR1NAE AND VASIGNYELLINAE

69

or without foramina and a suture. An additional 1-2 smaller pore-chambers distolateral to the orifice. Avicularia

paired or single. Ovicell unknown. Dorsal surface with broad pore-chambers confluent with the latcrofrontal

chambers.

Range. — Priabonian (Upper Eocene) to present day.

Type species. — Plagiopora disticha MacGillivray, 1895.

Congeneric species. — Two other species may be included in the genus. These are :

Plagiopora recens Gordon, 1989a, Holocene, near Norfolk Island, southwest Pacific.

Plagiopora alma sp. nov., Priabonian, Alma, near Oamaru, New Zealand.

Plagiopora dislicha MacGillivray, 1895

Fig. 7 a-e

Plagiopora disticha MacGillivray, 1895 : 79, pi. 13, fig. 14 a-c. — Maplestone, 1904 : 206. — Bassler, 1953 : G212.

fig. 160, 3 a-b. — Gordon 1989a : 1330.

MATERIAL EXAMINED. — Australia. Museum of Victoria : holotype slide P27769, from Muddy Creek, Victoria,

Middle Miocene (single internode on slide subsequently inadvertently lost by DPG). Royal Melbourne Institute of

Technology : slides of specimens from Princelown and Balcombe Bay, Victoria, Middle Miocene, collected by

P.E. Bock.

Description. — Segments up to 2.70 mm long and 0.47-0.72 mm wide (depending on avicularian

projections) with up to 10 zooids per segment. Zooids 0.43-0.49 mm long and 0.37-0.45 mm wide (again,

varying according to avicularian length), alternating along the segment with a zigzag interzooidal boundary axially.

Frontal shield comprising large shallow cryptocystal areas (pore-chambers) separated by thin gymnocystal strips.

The pore-chambers arc arranged such that, when seen in frontal view, each zooid has 2 (supraavicularian and lateral-

oral) on its inner, axial side (only 1 if there is no avicularium separating them), and 3 (supraavicularian, lateral-

oral, proximolateral) on its outer, marginal side. The 2 inner ones have a single septular pore; the 3 outer ones

have 2 to several pores clustered on or near the zooidal margin. The thin gymnocystal ridges separating, and

surrounding, the pore-chambers are continuous. The ridge between the inner lateral-oral and outer proximolateral

pore-chambers is widest and somewhat Y-shaped, with 6-7 small foramina distributed along the arms and tail of

the Y; the longer arm that courses obliquely from the orifice to the branch axis has a longitudinal suture flanked

by apparent vestigial costal elements, but the extent of their development varies considerably from zooid to zooid.

Orifice longer than wide, orientated obliquely on the zooid. Avicularia usually paired, occasionally single, with

complete cross-bar and acute triangular rostrum; facing laterally, the avicularian process short or considerably

projecting. Ovicell unknown. Dorsal surface comprising a pair of large pore-chambers per zooid which are

continuous with the outer lateral-oral and proximolateral ones on the frontal side; a few tiny septular pores occur

near the lateral margin in each chamber. Axial interzooidal boundary sinuous dorsally, not zigzag.

Distribution. — Victoria, Australia, Early Oligocene to Middle Miocene (Rupelian to ?Langhian/

Serravallian).

Remarks. — This species has been little reported on. According to the localities given by Maplestone

(1904) and Cockbain's (1971) corrected ages given by Wass (1973), this species is distributed from Early

Oligocene to Middle Miocene. The range of the genus in time, however, was extended to the present day when

Gordon (1989a) discovered in N.Z. Oceanographic Institute samples a new species, P. recens (fig. 8 a-c), at

831 m depth on the western flank of the Norfolk Ridge north of Norfolk Island. Although this was the first living

ditaxiporine species to be discovered, only isolated segments (still with membranes intact) were found, so it was of

some interest to discover several mature and attached colonies of a second living ditaxiporine (Bryosutor sutilis sp.

nov.) in Musorstom samples from the far northern part of the Norfolk Ridge. This biogeographic connection

70

D. P. GORDON & G. BRAGA

between the Norfolk Ridge and the Tertiary of Victoria is paralleled by other catenicelloidean Bryozoa, viz, species

of Peialosiegus and Chelidozoum (Gordon & d'Hondt, 1991). Subsequently, an Upper Eocene species (P. alma

sp. nov., see below) was discovered at Alma, South Island, New Zealand.

Fig. 7 a-e. —Plagiopora disticha MacGillivray : a, c, frontal views of segment and zooids (x 37 ; x 122) ; b, dorsal

view of proximal end of a segment (x 65) ; d. e, magnified views of the median gymnocystal ridge and vestigial costal

elements (x 388 ; x 683). (From Princetown, Victoria ; Bock Collection, Royal Melbourne Institute of Technology.)

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

71

The finding of a suture and apparent vestigial costae in the median gymnocystal ridge in Plagiopora dislicha

(made possible by the remarkable preservation) is significant, for it indicates that Plagiopora was derived from a

form with a costal field, a larger area of gymnocyst with extracostal windows, and smaller pore-chambers.

Plagiopora recens is even more derived but may nevertheless be included in the genus in spite of no trace of

vestigial costae. One significant difference in this species, however, is the occurrence of transverse gymnocystal

ridges on the dorsal surface. Plagiopora alma also lacks any trace of costae or gymnocystal windows, although it is

the earliest species of the genus. In frontal view, it more closely resembles P. recens than P. dislicha. Dorsally, it

is closer in appearance to P. disticha in lacking transverse gymnocystal ridges.

Plagiopora alma sp. nov.

Fig. 8 d-g

MATERIAL EXAMINED. — New Zealand. At Alma, north Otago. New Zealand, from the fine fraction in a

calcareous conglomeratic tuff, Waiareka Volcanic Formation, Runangan age (= Priabonian, Upper Eocene); N.Z. Fossil

Record File No. J41/f88, grid reference J41/453639, collected by D.P. Gordon, March 1993.

Description. — Unique segment 0.98 mm long and 0.32 mm wide, comprising 4 zooids. Zooids small,

O. 39-0.45 mm long and - 0.16 mm wide, widest in the distal half, tapering narrowly in the proximal half if at the

proximal end of the segment. Zooidal orifice somewhat pyriform and asymmetrical, slightly oblique, with the

distal end a little closer to the segmental axis; proximal end with a sinus-like narrowing, with 1 or 2 condyles

evident at the corners of the sinus. Frontal shield comprising shallow cryptocystal sectors (essentially broad

flattened pore -chambers) separated by narrow gymnocystal ridges; these sectors include 1 on each side subjacent to

the orifice and proximal to each lateral-oral avicularium (if there are 2), and a single large area over the proximal

half of the zooid. There are 2 additional sectors distolateral to the orifice above each avicularium. Thus there are

3 such areas along every outer zooidal margin, each of which continues around the margin to the dorsal side.

Septular pores are not evident in the cryptocystal sectors. Avicularia evidently generally paired, either side of the

orifice, each with a complete cross-bar, the inner one facing frontally and directed obliquely towards the orifice, the

outer one facing laterally and directed distally. Where the inner avicularium is lacking, as on the proximal zooid of

a segment, the cryptocystal sectors adjacent to the orifice merge as one. Ovicell unknown. Dorsal side of segment

with a sinuous median groove longitudinally paralleled by the gymnocystal margins adjacent to the cryptocystal

sectors.

Type. — Hololype : the unique segment illustrated in figure 8 d-g. from the Waiareka Volcanic Formation.

Alma, north Otago, New Zealand, type number BZ 162, Institute of Geological & Nuclear Sciences, Lower Hutt,

New Zealand.

DISTRIBUTION. — Runangan (= Priabonian, Upper Eocene), Oamaru, South Island, New Zealand.

Etymology. — The specific name is that of the type locality.

Remarks. — The road cutting through the Waiareka Volcanic Formation has exposed a calcareous tuff packed

with small stones, frequently encrusted by bryozoans, tubicolous polychaetes, and coralline algae. The line fraction

includes isolated segments of buguloidean and cellarioidean bryozoans, rare catenicellid fragments and, at one end

of the exposure, the warm-water foraminiferan Asterocyclina. The overall setting appears to have been the shallow

subtidal in well-oxygenated water of oceanic or near-oceanic salinity.

In its overall morphology, Plagiopora alma is somewhat intermediate between P . disticha and P. lecens.

Gymnocystal foramina are lacking in P. alma, as are vestigial costae, except that the proximal rim of the orificial

sinus gives the appearance of being derived from a suboral pair. While the trontal side of the segment closely

resembles that of P. recens. the dorsal side lacks transverse gymnocystal ridges. Although occurring earlier.

P. alma is less derived in frontal-shield morphology than P. disticha.

72

D. P. GORDON & G. BRAGA

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

73

Genus AHCHEETHAMIA nov.

Diagnosis. — Multizooidal, biseriate. Frontal shield a porous gymnocyst. Proximal rim of orifice composed

of a pair of vestigial costae. Avicularia single, borne laterally, ?or paired. Oviccll evenly porous like the frontal

shield. Dorsal surface with numerous pores.

Range. — Bartonian (Middle Eocene).

Etymology. — The genus is named after the author of the species. Dr Alan H. Cheetham (Smithsonian

Institution) in recognition of his outstanding contributions to bryozoology.

Type species. — Caberoides corniculatus Cheetham, 1966.

Ahcheetharnia corniculata (Cheetham, 1966)

Fig. 9 a-f

Caberoides corniculatus Cheetham, 1966 : 82, figs 60-61.

MATERIAL EXAMINED. — England. The Natural History Museum, London: Holotype specimen D48746, from the

Upper Bracklesham Beds, Fisher Bed 21, Selsey, Sussex, England (National Grid Reference SZ 836936), collected by

D. Cb'RRY and presented November 1961; paratypes D48751, D48756, D48766, D48767. from same locality as

holotype. Smithsonian Institution: 4 non-type specimens, from same locality as holotype.

Description. — Segments up to -2.20 mm long and 0.68 mm wide, with up to 12 zooids per segment.

Zooids 0.32-0.37 mm long and 0.26-0.37 mm wide, the gymnocystal frontal shield coarsely perforated by

numerous pores. Orifice with a pair of small condyles: the concave proximal rim composed of a pair of broad

vestigial costae. On the axial side of each orifice is an umbo, perforated like the frontal shield. On the outer,

lateral, side of the orifice is a frontolaterally facing avicularium, the opesia semicircular, the cross-bar complete,

the rostrum triangular; a process behind the rostrum may project beyond the avicularium. Ovicell recumbent, set

obliquely to the segmental axis, perforated like the frontal shield. Dorsal surface of segment with a median zigzag

suture line, the zooidal basal walls evenly perforated like the frontal shield.

Distribution. — Lower Bartonian (Middle Eocene), Sussex, England.

Remarks. — Our description of the species is based on SEM observations of uncoated specimens (most of

which were generously provided by Dr Paul D. Taylor, The Natural History Museum, London). The perforations

in the dorsal surface of each zooid are very curious. They do not appear to represent septular pores (no pore-

chamber boundary is evident on the lateral margin): rather, they appear to be the same as the frontal gymnocystal

pores, which is somewhat unusual.

Although Ahcheetharnia corniculata segments resemble those of Caberoides in their general form, the species

differs from the type and other species of Caberoides in having a porous frontal gymnocyst. only a suboral pair of

vestigial costae, no frontal pore-chambers, and a perforated ovicell. The porous gymnocyst. in fact, invites

comparison with that of Ditakiporina.

The genus is at present monotypic.

Fig. 8 a-c. — Plagiopora recens Gordon : a, frontal view of a segment with seven zooids (x 39) ; b, dorsal view of a

segment with six zooids (x 42) ; c, orifices and avicularia of two zooids (x 167). (From NZOI Stn G4, western Bank of

Norfolk Ridge north of Norfolk Island, 831 m depth).

Fig. 8 d-g. — Plagiopora alma sp. nov. : d, frontal view of the holotype segment (x 100) ; e, zooidal orifice (x 411) ;

f, dorsal view of the holotype segment (x 100) ; g, close-up of frontal shield (x 217). (From Alma, North Otago, New

Zealand, Priabonian, Institute of Geological & Nuclear Sciences, Lower Hutt.)

74

D. P. GORDON & G. BRAGA

Fig. 9 a-f. — Ahcheethamia corniculata (Cheetham) : a, fertile segment with three ovicelled zooids (x 43) ; b. same,

showing the three ovicells (x 95) ; c, another segment, showing the distolateral avicularia (x 95) ; d, e, dorsal surface

(x 54 ; x 152) ; f, close-up of left-hand ovicelled orifice in fig. b (x 190). (Figs a-c, f, uncoated holotype D48746,

BMNH, London ; d-e, uncoated paratype D48756, BMNH ; from the Upper Bracklesham Beds, Sussex.)

Source : MNHN, Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

75

Genus DITAXIPORINA Stach, 1935a

Diagnosis. — Multizooidal, biseriate. Frontal shield an evenly porous gymnocyst. Proximal rim of orifice

composed of a vestigial pair of costae, each with a lumen pore (pelma). Between each costa and the paired

distolateral avicularia is a tiny pore-chamber. Ovicell smooth, imperforate or with a pair of ectooecial fenestrae.

Dorsal surface of zooids smooth; a tiny pore-chamber may occur subjacent to each avicularium.

Range. — Lutetian (lower Middle Eocene) to Rupclian (Lower Oligocene).

TYPE SPECIES. — Catenicella septentrionalis Waters. 1891.

Ditaxiporina septentrionalis (Waters, 1891)

Fig. 10 a-d, 11 c-d

Catenicella septentrionalis Waters. 1891 : 5, pi. 1, figs 1-8. — Braga, 1963 : 41.

Ditaxiporina septentrionalis - Stach, 1935a : 395; 1935b : 46. — Cheetham, 1963 : 485. — Braga in ANT0UN1 el al„

1980 : 61, fig. 44. — Braga & Barbin, 1988 : 528.

MATERIAL EXAMINED. — Italy. The Manchester Museum. U.K. : slides #7488/140. #7490/140 of Catenicella

septentrionalis. Waters Collection, from Montecchio Maggiore, Upper Eocene. Dipartimento di Geologia dell'Universita

di Padova : specimens from Pannone P'l and Toara, Collezione micropaleontologica. Upper Eocene.

Description. — Segments up to 4.22 mm long and 0.51-0.56 mm wide, with up to 8 zooids per segment.

Zooids alternating, 0.48-0.74 mm long and -0.30 mm wide, relatively slender, the most proximal zooid of a

segment more or less claviform and proximally tapered. The axial furrow between the zooids zigzagging along the

segment. Frontal shield a gymnocyst, almost evenly perforated (fewer centrally) by numerous pores. Orifice with a

broad U-shaped sinusoidal poster, the proximal rim defined by a raised pair of short broad costae, with a thin

suture-line between them and each with a proximal pore. Adjacent to the orifice is a pair of avicularia, the axial

one possibly sometimes aborted, otherwise facing obliquely frontally, the marginal one facing laterally; each

somewhat projecting, each with a semicircular opesia, complete cross-bar. and short triangular rostrum; a tiny

pore-chamber occurs frontally between each avicularium and the adjacent costa. Ovicell subglobular, evidently

smooth, covering approximately half the frontal shield of the distal zooid. Dorsal surface of segment smooth¬

surfaced, the basal wall of each zooid somewhat convex, with only a single tiny pore-chamber behind the lateral

avicularium. The interzooidal furrow traces a sinuous course along the segment between the zooids.

Distribution. — Northeast Italy, Upper Eocene.

Congeneric species. — Three other species may possibly be included in this genus. These are :

[?]Catenicella granulosa Canu, 1908, Lutetian, near Paris, France.

Catenicella subseptentrionalis Canu & Bassler, 1917, Rupelian. southwest Arkansas, USA.

Ditaxiporina bifenestrata Cheetham. 1963, Rupelian, southwest Arkansas, USA.

Remarks. — The genus Ditaxiporina was discussed quite thoroughly by Cheetham (1963), who described a

new species, D. bifenestrata, with bifenestrate ovicells. In all other respects this species closely resembles the type

species. Although it is not impossible that the ovicell was truly bifenestrate, the possibility that the fenestrae

represent eroded predator bore-holes cannot be ignored - selective predation of ovicells has been noted by Gordon

& d'Hondt (1991) in the catenicelloidean genus Petalostegus. Apart from the type species, the ovicell has not

been discovered in any of the other species ascribed to Ditaxiporina, which may be distinguished on the basis of

zooidal characters, such as details of size, distribution of frontal pores, and placement and orientation of the

avicularia.

76

D. P. GORDON & G. BRAGA

The exact status of CateniceUa granulosa is uncertain. It was described as lacking avicularia and, indeed, the

unique (holotype) specimen (fig. 9 e) appears to lack them, but the preservation is so poor that it is impossible to

be certain. The dorsal surface [now inaccessible, since it was glued to the slide after Canu (1908) illustrated it]

was described as smooth and convex. Thanks to the courtesy of Dr Agnbs Rage (Institut de Paldontologic,

Museum national d'Histoire naturelle, Paris) and Dr Paul Taylor (Department of Palaeontology, The Natural

History Museum, London) it was possible to borrow and scan (uncoatcd) the holotype specimen. Because of the

poor state of preservation it is impossible to be certain if it belongs to Ditaxiporina, though the description of a

smooth dorsal surface would support this.

The occurrence of uni-, bi-, tri-, and multizooidal segments in the fossil samples (see Waters, 1891) suggests

the manner in which the segments grew. More significantly, the unizooidal segments are nearly identical to those

of the present-day species Vasignyella otophora (Kirkpatrick, 1890), which has only unizooidal segments. In fact,

the similarity is so great (Gordon & d'Hondt, 1991) that it is inconceivable that Vasignyella is unrelated to

Ditaxiporina. For this reason, Vasignyella is here separated from the Savignyellidae and included in the

Catenicellidae.

Subfamily VASIGNYELLINAE subfam. nov.

Diagnosis. — As for the sole included genus.

Genus VASIGNYELLA Gordon 1989b

Diagnosis. — All segments unizooidal. Zooidal frontal shield an evenly perforated gymnocyst. Proximal rim

of orifice composed of a vestigial pair of costae, lacking lumen pores (pclmata). Between each costa and the

distolateral pair of avicularia is a tiny pore-chamber. A row of small round uniporous pore-chambers occurs along

each margin. Ovicell unknown. Dorsal surface smooth, with only a small pore-chamber behind each avicularium."

Range. — Holocene.

Type species. — Catenaria otophora Kirkpatrick, 1890.

Vasignyella otophora (Kirkpatrick, 1890)

Fig. 11 a-b

Catenaria otophora Kirkpatrick, 1890 : 17, pi. 5, figs 1-lc. — Philipps, 1900 : 441. — Tiiornely 1912 • 139 —

Gravely, 1927 : 89, pi. 11, fig. 6.

Savignyella otophora - HaRMER, 1957 : 763, pi. 51, figs 19-21. — DUMONT, 1981 : 636. — Winston 1986 • 27

Vasignyella otophora - Gordon. 1989b : 453, figs 13-15. — Ryland & Hayward. 1992 : 247, fig. 14a.

Material EXAMINED. — Western Samoa. N.Z. Oceanographic Institute : sparse material from Sa’aga, Upolu

Island, collected by J.E. Morton (University of Auckland).

Description. — Colony erect, comprising branching uniserial chains of unizooidal segments. Zooids

claviform, bent somewhat forward, 0.79-1.30 mm long and 0.24-0.25 mm wide. Frontal shield a porous

Fig. 10 a-d. — Ditaxiporina septentrionalis (Waters) : a, segment of four autozooids (x 79) ; b, close-up of the right-hand

proximal zoo id in fig. a - note the lumen pores in the suboral costae (x 317) ; c, ovicelled zooids (x 88) ; d, dorsal

surface of zooids (x 108). (Figs a-b, Pannone, Italy. Upper Eocene ; Waters Collection. Manchester Museum. Figs c-d,

same locality, Braga Collection, Dipartimento di Geologia, Paleontologia e Geofisica dell'Universita di Padova.)

Fig. 10 e. — Ditaxiporina granulosa (Canu), uncoated unique holotype segment affixed to slide (x 56). (From Orglandes,

near Paris, Middle Eocene ; slide R53352, Institut de Paleontologie, Museum national d'Histoire naturelle, Canu

Collection.)

Source : MNHN. Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

77

Source : MNHN. Paris

78

D. P. GORDON & G. BRAGA

gymnocyst. Orifice with a pair of small condyles delimiting the anter from the broadly V-shaped poster. Proximal

rim of orifice delimited by a pair of short broad costae with a median suture. Lateral-oral avicularia with triangular

rostrum, the cross-bar lacking, facing frontolaterally. Ovicell unknown. A row of ~5 small circular pore-chambers

occurs along each lateral margin, the septulum uniporous. An additional small pore -chamber occurs on each side

between the avicularium and the adjacent costa, and behind the avicularian rostrum. Dorsal surface smooth.

Fig. 1 1 a-b. — Vasignyella otophora (Kirkpatrick) : a, autozooid (x 103) ; b, same, close-up of orificial region -note the

small pore-chambers between the avicularia and suboral costae, and the lack of lumen pores in the latter (x 205).

(From Sa'aga, Upolu Island. Western Samoa.)

Fig. 1 1 c-d. Dilaxiporina septentrionalis (Waters) : c, orificial region of another unizooidal segment ; cf. plate 10, b -

note the lumen pores in the suboral costae (x 299) ; d, young unizooidal segment prior to budding of the additional

zooids (x 149). (From Pannone, Italy, Upper Eocene ; Waters Collection, Manchester Museum.)

Distribution. — Sudanese Red Sea, Indian Ocean, China Sea, Philippines, New Guinea. Great Barrier Reef,

Fiji, Western Samoa, intertidal to 49 m.

Remarks. Although the unizooidal segments of Dilaxiporina septentrionalis appear to lack the small

lateral pore-chambers found in Vasignyella , the arrangement of elements in the orificial region is extraordinarily

similar, including the presence of small pore-chambers between the avicularia and costae and behind the avicularia.

The costae of Vasignyella otophora lack lumen pores, however, and individual zooids are somewhat larger than

those of D. septentrionalis. There is one other difference, which may be significant in view of its otherwise

apparent universality among ditaxiporines - viz, the lack of an avicularian cross-bar in Vasignyella.

Source : MNHN. Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VASIGNYELLINAE

79

At present, Vasignyella is monotypic and has no known fossil record. The absence of small lateral pore-

chambers in Ditaxiporina may indicate that Vasignyella was not derived directly from it but from, a common

predecessor. Significantly, ovicells have never been found in Vasignyella, possibly indicating internal brooding.

Because of this, we propose a separate subfamily, Vasignyellinae, additionally characterised by only unizooidal

segments and the lack of an avicularian cross-bar. In our opinion, Vasignyella has no direct relationship with

Savignyella Levinsen and Halysisis Norman, although we suspect that the Savignyellidae may nevertheless be

derived from an early catenicelloidean stock. We intend to discuss the genera of the Savignyellidae in a future

paper.

SYSTEMATIC AFFINITIES

On the basis of the mixture of shared and derived characters among the genera discussed in this paper, we

conclude that, with the exception of Vasignyella, all the other genera may be allied in the one subfamily,

Ditaxiporinae Stach, 1935a, in which the family Ditaxiporinidae Cheetham, 1963 may be subsumed.

Diversification of the family Catenicellidae (Maastrichtian to present day) from the Cribrilinidae sensu laio

presumably occurred during the Senonian, though the details are unknown. Although some cribrilinids evidently

had articulated colonies, the segments were stick-like, not oligozooidal. Segments of one or a few zooids, and

relatively large ovicells, are shared with another catenicelloidean family, the Petalostegidae, which may be regarded

as a sister group. These characters are thus synapomorphies for these two families. In the subfamily Ditaxiporinae,

ancestral characters (plcsiomorphies) include multiple costal elements, small or linear pore-chambers, and lateral-

oral avicularia. Derived characters (apomorphics) include vestigial suboral costae or no costae, enlarged pore-

chambers (especially dorsally), and gymnocystal ridges. The assignment of morphological polarities is based on

evident trends in other catenicelloideans as well as in hippothooideans (separately derived from cribrilinids), viz, a

reduction in the size of the costal shield and/or the number of costal elements with a concomitant increase in the

area of either gymnocyst or cryptocystal elements (e.g., extracostal windows and/or pore-chambers), with

accompanying specialisations (see Banta & Wass, 1979; Gordon & d'Hondt, 1991). An autapomorphy found

only among the Ditaxiporinae is biseriate multizooidal segments of non-reproductive zooids (in contradistinction

to the occasional fertile multizooidal segments in some species of Catenicella). Although known only from the

Holocene, Bryosartor is in many respects the least-derived ditaxiporine genus. In fact, there are significant

similarities to Costaticella (Scuticellinae) in the multicostate spinocyst and the extracostal foramina. Bryosartor

may be a much older genus than is indicated by its lack of a fossil record, a possibility supported by its occurrence

on the northern Norfolk Ridge where a high proportion of archaic taxa occur (Richer de Forges, 1987, 1990).

Alternatively, Bryosartor may be a more recently derived genus which has retained ancestral features, but whose

immediate predecessors are unknown.

There are several parallels between scuticelline/catenicellinae and ditaxiporine genera. For example, the earliest-

occurring genera of both subfamilies have costal shields, and there are similar trends in the diversification of

frontal-shield morphologies. Strophipora resembles Ditaxipora in the possession of a median gymnocystal band

flanked by broad shallow pore-chambers that effectively make the frontal shield largely cryptocystal. For this

reason, Canu and Bassler (1929) proposed that the latter genus should be a subgenus of the former, a suggestion

that Stach (1934) rightly rejected. Cribricellina, with its perforated gymnocystal shield, parallels Ahcheethamia

and Ditaxiporina.

BIOGEOGRAPHY

The distributions in time and space of the genera of Ditaxiporinae and Vasignyellinae are shown in figures 12

and 13. Apart from the occurrence of two species of Ditaxiporina in southwest Alabama, the combined distribution

of all the taxa is notably Tethyan. For the Ditaxiporinae, there are two main distributional clusters - amphi-

Atlantic/European, and Australasian. Ditaxiporina is the only American (and amphi-Atlantic) genus. Interestingly,

80

D. P. GORDON & G. BRAGA

PALEOGENE

PALEOCENE

EARLY LATE

g

>

2

1

>

2

g

>

Caberoides

EOCENE

EARLY

-c

•0

33

m

c/>

>

2

Ditaxiporina

Ditaxipora

MIDDLE

c

H

g

Ahcheethamia

LATE

>

33

3

OLIGOCENE

EARLY

33

c

■0

m

□

>

2

Ptagiopora

8!

LATE

o

X

>

NEOGENE

MIOCENE

EARLY

03

c

X

g

G)

>

MIDDLE

c n

m

33

2

LATE

3

3

Bryosartor

Vasignyella

0)01 w .

-vj io ^ 2

FIG-2- ~ Stratigraphic distribution of the genera of the catenicellid subfamilies Ditaxiporinae and Vasignyellinae

throughout the Cenozoic [chronometnc scale based on Harland el at. (1990)].

Fig. 13. Biogeographic distribution of the genera of the catenicellid subfamilies Ditaxiporinae and Vasignyellinae.

Source : MNHN. Paris

BRYOZOA : SUBFAMILIES DITAXIPORINAE AND VAS1GNYELI.INAE

81

the earliest-known genus. Caberoides, achieved a potentially Tethyan-wide distribution by the Ypresian, ranging

from the Northeast Atlantic to Chatham Island, New Zealand within 10-Ma. Clearly, the earliest ditaxiporines

must be sought in the Danian or Maastrichtian. Like Caberoides , Diiaxipora is found at either end of the Tethyan

corridor. It is significant that the oldest species of both of these genera are European, indicating that the

Australasian taxa are derivative. Lagaaij and Cook (1973) and Braga (1987) have documented similar

distributions in other cheilostomes, from which Braga concluded that faunal migration was eastwards from the

Mediterranean to the Indo-Pacific. The information presented here supports that scenario for the amphi-Tethyan

ditaxiporine genera.

The Ditaxiporinae of the western area of Tethys, belonging to the Paleogene of Venetia, occur in levels that are

sometimes thick and continuous. Reported in the literature as 'Bryozoa marls' or 'Brendola marls', they are a key

bed in studying the Upper Eocene sequences and are very rich in bryozoans. They outcrop over a wide area,

including the Southern Trentino, the Lessini Mountains, the Berici Mountains, the piedmont area of the Asiago

plateau, and the western Treviso Hills. This area, here called the Venetia shelf, was a structural high (fig. 14)

where shallow-water marly calcareous deposits (nummulitic limestones, fossiliferous marls) were laid down. It

corresponds largely to the resurrected Trento platform, an important Liassic structure drowned and successively

overlain by Upper Jurassic pelagic limestones of the Ammonitico Rosso Formation (BOSELLINI, 1989).

Fig. 14. — Location of the Upper Eocene bryozoan outcrops in the Venetia shelf, an important paleogeographic structure

in the western Tethyan basin (modified from Bosellini 1989). Explanation of symbols:

1 : westward - marly limestones (Scaglia cinerea Formation); eastward - marls and sandy and calcareous shales (flysch

facies)

2 : shallow water deposits (shelf facies)

3 : hemipelagic deposits

4 : main Upper Eocene bryozoan outcrops of Venetia (NE Italy), marked by asterisks (*): 1, Possagno; 2, Brendola-

Toara, Monteccio di Costozza; 3, Val di Lonte; 4, Priabona; 5, Verona; 6, Pannone.

82

D. P. GORDON & G. BRAGA

Paleoecologically, the Venetia shelf was marginally characterised by patch reefs, lagoons, and coastal

embayments, with coralline-algal meadows and sublittoral bryozoan associations. The Bryozoa marls show an

extremely rich diversity, with abundant representation of rigid-erect branching growth forms, indicative of

relatively fast water flows well supplied with particulate food, especially microalgae. There were also articulated,

rod-like, and vagrant colonial morphologies. Thus, in addition to catenicellids, there were also stratigraphically and

paleoecologically significant species like Lacrimula perfecta (Accordi), typically associated with soft seafloor

sediments of carbonate mud or more or less muddy sands.

A significant diversity occurs among the petrofacies of the different marginal deposits of the Venetia shelf. For

example, the residue of the Pannone washed samples (western area) shows extrabasinal clasts (quartz, potassium

feldspar, mica), presumably coming from a source area containing granitic and metamorphic rocks of the

neighbouring northern land. In the eastern area (Possagno), silty clayey or clastic distal sediments prevail,

originating from the eastern platforms or the above-mentioned area. In the centre of the shelf, east of Lake Garda

(fig. 14, inset), the Bryozoa levels are are fairly rich in carbonates and contain numerous nummulites, bivalves,

algae, and coral fragments. The bryofaunas are, however, quite uniform and would not seem to have been limited

by environmental factors.

According to sedimentological characteristics, faunal content, growth-form parameters, and inferences

concerning the species in life, the depth range of the shelf would have been within the photic zone of the inner-

outer interval, not more than 100 m. The sea was warm-temperate or subtropical, with well-oxygenated water of

normal salinity, judging from the high species diversity of the Bryozoa. In summary, we can hypothesize that the

modem, predominantly Australasian, distribution of catenicellids (including the Ditaxiporinae and Vasignyellinae)

may be derived from a wider circumtropical Paleogene distribution (cf. Lagaaij & Cook, 1973; Braga, 1987).

ACKNOWLEDGEMENTS

We wish to extend our special thanks to the following people : Rosemary Swart (Museum of Victoria,

Melbourne), Phil Bock (Royal Melbourne Institute of Technology), Alan CHEETHAM (Smithsonian Institution,

Washington, D.C.), and Agnes Rage (Museum national d'Histoire naturelle, Paris) for the loans of specimens;

and Paul Taylor (The Natural History Museum, London) for technical assistance to D.P. Gordon in the

scanning of uncoated type material. This research was partly funded by the New Zealand Foundation for Research,

Science and Technology (Contract CO 1217) as part of NIWA's Marine Taxonomy Programme. Financial

assistance to DPG was also provided by the Museum national d'Histoire naturelle, Paris (July 1989), and

ORSTOM, Noumea and the British Council (July 1992). G. Braga's research was partly supported by grant

MURST-60% 1993 (G. Piccoli).

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ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULTATS DES C A

Cirripedia Thoracica : Verrucomorpha

of New Caledonia, Indonesia,

Wallis and Futuna Islands

John S. BUCKERIDGE

UNITEC Institute of Technology

Private Bag 92095, Auckland, New Zealand

ABSTRACT

Verrucomorpha from deep sea collections made by several French cruises in New Caledonian area (including

Chesterfield and Loyalty Islands), Wallis Island and Futuna Island waters, and by the French-Indonesian cruise Karubar

in Indonesian waters, over the period 1985-1993, are discussed. The fauna includes 16 species, seven of which are new;

the bathymetric and geographic ranges of many other taxa are extended, making this one of the most diverse

verrucomorph faunas known. The status of species within both Altiverruca and Metaverruca is considered, with six names

being synonomised with Metaverruca recta (Aurivillius), and a key to genera of the Verrucidae is given.

The distribution of the Verrucidae in the region is discussed, with Metaverruca recta being confirmed as the species

with the greatest geographic, bathymetric and stratigraphic distribution of any verrucid.

RESUME

Cirripedia Thoracica : Verrucomorpha de Nouvelle-Caledonie, d'lndonesie et des ties Wallis et

Futuna.

Les VerTucomorphes, principalement bathyaux, recoltes lors de diverses campagnes franfaises au large de la Nouvelle-

Caledonie et des lies Loyaut6, Chesterfield, Wallis et Futuna et lors de la campagne franco-indon6sienne KaRUBAR dans

les eaux indonesiennes, entre 1985-1993, sont etudids ici. Seize especes ont 6t<5 trouvees, parmi lesquelles sept sont

nouvelles pour la science. La faune des Verrucomorphes etudiee ici apparait ainsi comrne l'une des plus diversifiee connue

dans une meme region. Les distributions bathymetrique et geographique de plusieurs especes sont dtendues.

Le statut des especes des genres Altiverruca et Metaverruca est examine. Six especes sont mises en synonymie avec

Metaverruca recta (Aurivillius), et une cl6 des genres de la famille Verrucidae est propos6e.

La repartition des Verrucidae dans la region prospectec est discutee, et il est confirme que Metaverruca recta est

l’espfcce ayant la plus vaste repartition geographique, bathymetrique et stratigraphique de tous les Verrucidae.

Buckeridge, J. S., 1994. — Cirripedia Thoracica : Verrucomorpha of New Caledonia, Indonesia, Wallis and Futuna

Islands. In : A. Crosnier (ed.), Resultats des Campagnes MUSORSTOM, Volume 12. Mem. Mus. natn. Hist. not.. 161 :

87-125. Paris ISBN 2-85653-212-8.

88

J.S. BUCKERIDGE

INTRODUCTION

The deep sea verrucomorph cirripede fauna of Indonesia, New Caledonia and the Wallis and Futuna Islands had

not been collected prior to 1977; although previous work on verrucomorphs (generally shallow water species) in

the Indo-Pacific region has been carried out by Hoek (1907, 1913); Pilsbry (1907, 1912); Broch (1922, 1932);

Gruvel (1907. 1920); Rosell(1981. 1991); Nilsson-Cantell (1929); Foster (1978); Buckeridge (1983)

and Zevina (1987, 1988). This paper records material collected from New Caledonian waters by Biocal (1985),

Musorstom 4 (1985). Chalcal 2 (1986), Biogeocal (1987), Smib 5 (1989), Smib 8 (1993); from the Loyalty

Islands region by Musorstom 6 (1989); from the Chesterfield Islands region by Musorstom 5 (1986) and

Corail 2 (1988); from the volcanoes south of Vanuatu by Volsmar and Gemini (1989); from the Wallis and

Futuna Islands region by Musorstom 7 (1992), and from Indonesian waters by Karubar (1991) expeditions.

Some specimens collected by the Musorstom 3 (1985) expedition in the Philippines and not studied by Rosell

(1991) are also recorded. The collections examined have shown an exceptional diversity of Vcrrucidae : from the

124 sites considered. 16 vcrrucid species are recorded; seven of these are new to science, and the range of many

other taxa is extended. This paper also considers aspects of verrucomorph systcmatics, biogeography and

distribution.

The Verrucidae are characterised by six calcareous plates. The shell is made up of the carina and rostrum, plus a

tergum and scutum, the latter two having moved from the operculum to become "fixed" as part of the shell wall.

The remaining plates, a tergum and scutum (hereafter termed movable plates) make up the operculum. It appears to

be totally random as to whether the "left" or "right" opercular plates move to complete the shell wall (Darwin,

1854), in the specimens studied the ratio being close to 50:50. However, in Neoverruca, right or left sidedness

depended on which side the juveniles settled on, relative to the current (Newman, 1989).

Types : All new taxa are either from New Caledonian or Wallis and Futuna waters; holotypes are held by the

Musdum national d'Histoire naturclle (e.g. MNHN-Ci) in Paris, France. Paratypes are held by the Museum

national d'Histoire naturelle, and when numbers of specimens available permit, by the National Museum of

Natural History, Washington (e.g. USNM) and UNITEC Institute of Technology, New Zealand (e.g. CAX).

In addition to types, figured material collections have also been allocated registration numbers (e.g. MNHN-Ci.

held by the Musdum national d'Histoire naturclle, Paris).

In the lists of material examined the abbreviations of the gears used are : CC = otter trawl (shrimps); CP =

beam trawl; DC = Charcot dredge; DS = Sanders epibenthic dredge; DW = Warren dredge; KG = Usnel Box-Corcr.

LIST OF SPECIES

Genus VERRUCA Schumacher, 1817

Verruca albatrossiana Pilsbry. 1912

Genus ALTIVERRUCA Pilsbry, 1916

Alliverruca cristallina (Gruvel, 1907)

Al liver ruca galapagosa Zevina, 1987

Alliverruca laeviscuta sp. nov.

Alliverruca navicula (Hoek. 1913)

Alliverruca nitida (Hoek, 1883)

Genus CAMERAVERRUCA Pilsbry, 1916

Cameraverruca nodiscuta sp. nov.

Genus BROCHIVERRUCA Zevina, 1993

Brochiverruca dens (Broch. 1932)

Brochiverruca poly striata sp. nov.

Genus META VERRUCA Pilsbry, 1916

Metaverruca defayeae sp. nov.

Metaverruca norfolkensis sp. nov.

Metaverruca paciftca sp. nov.

Metaverruca plicata sp. nov.

Metaverruca recta (Aurivillius, 1898)

Genus ROSTRATOVERRUCA Broch, 1922

Rostratoverruca intexta (Pilsbry, 1912)

Rostratoverruca kruegeri (Broch, 1922)

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

89

SYSTEMATIC ACCOUNT

Order THORACICA Darwin, 1854

Suborder VERRUCOMORPHA Pilsbry, 1916

Diagnosis. — Sessile thoracican cirripedes, with an asymmetrical shell wall comprising a fixed scutum, fixed

tergum, rostrum and carina, closed by a movable scutum and movable tergum (Verrucidae), sometimes including a

rostrolatus and a carinolatus on the movable side (Proverrucidae), or basal whorls of imbricating plates

(Neoverrucidae); basis membranous or calcareous; caudal appendages usually present; mandible generally tridentoid;

crest of bullate labrum usually bearing teeth.

DISCUSSION. — Newman and HESSLER (1989) described Neoverruca brachylepadoformis , a living, abyssal

species that demonstrated links between the Brachylepadomorpha and the Verrucomorpha. The significance of this

find warranted a closer analysis of verrucomorph cirripedes, with a resultant split of the family, such that the

revised Verrucidae comprise only those verrucomorphans with six plates.

Family VERRUCIDAE Darwin. 1854

(amend. Newman & Hessler, 1989)

Diagnosis. — Verrucomorphans with the primary wall (carina, rostrum, fixed scutum, fixed tergum), in

contact with the substratum and without latera of any sort.

Discussion. — In 1916, PILSBRY proposed four "Sections" or subgenera in the genus Verruca, and these were

later added to by Broch (1922), and Zevina (1978), who also elevated the subgenera to generic status. This paper

follows Zevina's revision and includes her 1993 genus : Brochiverruca.

The presence of a myophore on the fixed scutum is generally of considerable value in classification, although

in Brochiverruca, the myophore is absent from B. dens but present in B. polystriata sp. nov. It should also be

noted that the angle of the operculum to the base may vary, particularly in Alliverruca , where the nature of the

substrate appears to have a strong influence. Metaverruca and Verruca however, appear far more consistent, as

I have not seen specimens with the operculum even close to perpendicular to the base. Pilsbry (1916) concluded

that much could be deduced about the habitat of verrucids by the shell shape. His material demonstrated that

shallow water forms were invariably depressed, with broadly spreading walls, whilst erect forms were found in deep

water. The present collection, with depressed forms like Metaverruca recta having been collected from depths in

excess of 2000 metres, does not bear this out.

I am a little uncertain about the status of Spongoverruca Zevina, (1978). Although some verrucids appear to be

host specific, I am not confident that this is a sufficient criterion to warrant generic separation. Further, many

workers, from Pilsbry (1916), consider a fixed scutal myophore to be of importance in defining verrucid genera,

unfortunately in her diagnosis of Spongoverruca, Zevina makes no specific reference io this character (V.

Melnikov, pers. com.).

Morphology of the soft parts can be of considerable help in identifying species, particularly of Metaverruca.

However all new taxa described here possess differences in shell morphology as well, with most species being able

to be identified on shell morphology alone. Most verrucids possess a mandible with a tridentoid cutting edge, the

lower angle being pectinate, hirsute or a mixture of both. The first maxilla generally has one or two prominent

upper spines, a central notch that may be hirsute or spinose, and a lower angle with shorter spines and hairs. Both

the penis and caudal appendages vary considerably between taxa. ranging from less than the length of the basal

segment of the pedicel of cirrus VI. to more than the length of the entire cirrus. The nomenclature used in this

paper is explained in figure 1.

90

J.S. BUCKERIDGE

STRATIGRAPHIC RANGE. — ?Midclle, Upper Crelaceous to Recent. Europe, South America, Australasia.

DISTRIBUTION. — Cosmopolitan in present seas.

Key to genera of the Verrucidae

1. Rostral and carinal apices marginal . 2

— Rostral apex only removed from margin . Rosiraloverruca

— Rostral and carinal apices removed from margin . Brochiverruca

2. Myophore present on fixed scutum . 3

— Myophore absent on fixed scutum . 4

3. Operculum nearly vertical to base . Cameraverruca

— Operculum nearly parallel to base . Metaverruca

4. Operculum nearly vertical to base . 5

— Operculum nearly parallel to base . Verruca

5. Not embedded in sponge . Altiverruca

— Embedded in sponge . Spongoverruca

Genus VERRUCA Schumacher, 1817

Verruca "Section B"- PlLSBRY, 1916 : 23.

Verruca (Verruca) - Foster, 1978 : 68.

Verruca - Zevina, 1978 : 1812. — Newman & Hessler, 1989 : 268.

Diagnosis. — Verrucids with apices of rostrum and carina marginal; fixed scutum without myophore;

operculum parallel to base.

Type. — Lepas stroemia Muller, 1776. North Atlantic, northern Europe, Mediterranean, ?Red Sea, intertidal -

500 metres.

Species. — 24 taxa are presently attributed to this genus : V. stroemia (Muller, 1776), V. laevigata

(Sowerby, 1827), V. prisca Bosquet, 1853, V. pusilla Bosquet, 1857, V. nuciformis Buckeridge, 1983, V. rocana

Steinmann, 1921, V. spengleri Darwin, 1854, V. tasmanica tasmanica Buckeridge, 1983, V. tasmanica chatheca

Buckeridge, 1983, V. cookei Pilsbry, V. withersi Newman & Schram, 1980, V. alba Pilsbry, 1907,

V. alba caribbea Pilsbry, 1916, V. alba barbadensis Pilsbry, 1916, V. calotheca Pilsbry, 1907. V. calotheca

flavidula Pilsbry, 1916, V. calotheca heteropoma Pilsbry, 1916, V. calotheca niasiensis Nilsson-Cantell, 1929,

V. floridiana Pilsbry, 1916, V. xanthia Pilsbry, 1916, V. xanthia insculpta Pilsbry, 1916, V. entobapta Pilsbry,

1916, V. macani Stubbings, 1936, V. scrippsae Zullo, 1964.

Discussion. — The status of Verruca sensu stricto is a little unclear at present. There are forms like Verruca

stroemia and V. laevigata that clearly fall into a distinct group : the shells possess a low profile, are found in

higher energy, shallow water environments, and of necessity have the operculum approximately parallel to the

substrate. Deeper water forms like Verruca albatrossiana Pilsbry are quite different, and although the operculum is

indeed "sub-parallel" to the basis, V. albatrossiana is normally found growing attached to cylindrical objects like

cidaroid spines. As such the base is curved, and in theory, will at some point parallel the operculum. 1 believe

greater consideration of genus Verruca is warranted, and it is likely that the V. albatrossiana- like forms will form a

separate genus. As there is only one " Verruca " species present in this collection, it is considered prudent to await

consideration of a wider range of material before splitting the genus. A further complication exists in the

Source : MNHN, Paris

VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

91

terminology used by Broch (1932), where he uses the term Eu-Verruca for the central group of genus Verruca

sensu lato. In a somewhat ambiguous footnote, Broch (1932 : 45) states that Eu-Verruca "... may serve to bring

confusion about".

Stratigraphic RANGE. — Cretaceous (Australia, New Zealand, Columbia, western Europe), Palaeogene

(New Zealand, Argentina, Chatham Islands).

Distribution. — Pacific, North Atlantic, Mediterranean, Caribbean, ?Red Sea, Indian Ocean, intertidal -

620 metres.

Verruca albatrossiana Pilsbry, 1912

Fig. 1 a-f

Verruca albatrossiana Pilsbry, 1912 : 292.

Verruca ( Eu-Verruca ) albatrossiana - Broch, 1922 : 290, figs 39, 40.

Verruca grex Hoek, 1913 : 142, pi. 11, figs 7-13, pi. 13, figs 11-13.

MATERIAL EXAMINED. — Indonesia. Karubar :stn CP 39, 07o45'43’'S, 132°28'22"E, 466-477 m, 28.10.1991 :

19 specimens (MNHN-Ci 2291), on cidaroid spines. — Stn CC 57, 08°15'48"S, 131°56'38"E, 603-620 m, 31.10.

1991 : 9 specimens, on cidaroid spines.

Diagnosis. — Verruca with rostrum and carina low, strongly developed longitudinally to give the shell an

oblique appearance; fixed scutum considerably larger than fixed tergum, opercular plates displaced carinally.

Description. — Shell white, walls ribbed vertically; obliquely skewed towards the carina; rostrum moderately

low, with an upper, broad triangular portion extending towards the carina; carina long, low, extending beneath

much of the rostrum.

Fixed scutum considerably larger than fixed tergum, with a "folded over" portion on movable scutal margin,

forming a zone confluent with the plane of the operculum; both fixed tergum and scutum with apices concurrent

with those of the movable plates.

Movable scutum a little narrower than the movable tergum, triangular, with five diagonal ribs, primary rib

arced towards tergal margin, strongly rounded, terminating at the basi-tergal angle, first secondary rib immediately

adjacent to primary rib, secondary ribs interlocking with tergum; apex beaked.

Movable tergum quadrilateral, apex slightly curved towards scutum, primary apico-basal rib almost straight,

with four secondary ribs.

Mandible tridentate, lower angle pectinate and divided into two portions; first maxilla with two large upper

spines, a pectinate notch, lower angle with five shorter spines. Crest of labrum with fine, numerous teeth, palps

pointed.

Cirrus I relatively hirsute compared with other cirri, inner ramus shorter; penis a little less than two thirds the

length of cirrus VI; caudal appendages very long. Cirri from MNHN-Ci 2291 possessed the following number of

segments (r-c being rostro-carinal diameter) :

r-c(mm) I II IE IV V VI c.a.

7.0 9,18 7,18 13,19 12,19 18,19 21,22 27

Discussion. — This species is characterised by "the unusual length of the rostrum and fixed tergum"

(Pilsbry, 1912), giving a distinct obliqueness to the shell, with a resultant displacement the opercular plates

carinally. It falls into a group with Verruca grex Hoek, 1913, but is distinguished from V. grex by possessing a

less elevated operculum, and fewer apico-basal ribs on the movable scutum.

Distribution. — Indo-Pacific, 345-620 metres.

92

J.S. BUCKERIDGE

Fig. 1. — Verruca albatrossiana Pilsbry, 1912 : a, complete shell (right side). MNHN-Ci 2291; b, movable tergum (left,

exterior); c, movable scutum (left, exterior); d. mandible; e, first maxilla; f, pedicel of sixth cirrus, with penis and

caudal appendage attached. All material from KaRUBAR, stn CP 39. Figures b and c are at the same scale.

Abbreviations : a.m.s. = articular margin of the movable scutum; a.m.t. = articular (scutal) margin of the movable

tergum; a.r. = apico-basal ridge of scutum (and tergum); a.u.c. = apical umbone of carina; a.u.r. = apical umbone of

rostrum; b.m.s. = basal margin of the movable scutum; b.m.t. = basal margin of the movable tergum; b.s. = basal

segment of the pedicel of cirrus VI; c.a. = caudal appendage; c.m.l. = carinal margin of the movable tergum; f.s. =

fixed scutum; f.t. = fixed tergum; La. = lower angle of the first maxilla; n. = notch of first maxilla; o.m.s. = occludent

margin of the movable scutum; o.m.t. = occludent margin of the movable tergum; p. = penis; t.h. = penis with hirsute

terminus; u.s. = upper spine of the first maxilla; u.i. = upper tooth of the mandible.

Genus ALTIVERRUCA Pilsbry, 1916

Verruca "Section D": Altiverruca Pilsbry, 1916 : 40.

Verruca (Altiverruca) - FOSTER, 1978 : 68.

Altiverruca ■ Zevina, 1978 : 1813.

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

93

Diagnosis. — Verrucids with erect form, bases of plates not inflected; operculum close to vertical; myophore

absent.

Type. — Verruca hoeki Pilsbry, 1907 : 113. West Indies, 907-1060 metres.

Species. — The 34 taxa presently attributed to this genus include : A. gibbosa (Hoek, 1883), A. incerta

(Hoek, 1883), A. obliqua (Hoek, 1883), A. nitida (Hoek, 1883), A. quadrangularis (Hoek, 1883), A. sulcata

(Hoek, 1883), A. crenata (Aurivillius, 1898), A. erecta (Gravel, 1900), A. longicarinata (Gravel, 1900),

A. radiata (Gravel, 1901), A. cristallina (Gravel, 1907), A. darwini (Pilsbry, 1907), A. hoeki (Pilsbry, 1907),

A. mitra (Hoek, 1907), A. plana (Gravel, 1907), A. cassis (Hoek, 1913), A. casula (Hoek, 1913), A. navicula

(Hoek, 1913), A. bicornuta (Pilsbry, 1916), A. rathburniana (Pilsbry, 1916), A. cristallina laevis (Broch, 1922),

A. gibbosa somaliensis (Nilsson-Cantell, 1929), A. aves (Zevina, 1975), A. angustiterga Zevina, 1987,

A. galapagosa Zevina, 1987, A. gira (Zevina, 1987), A. sublima Zevina, 1987, A. sculpturata Zevina, 1987,

A. longa Zevina, 1988, A. tchesunovi Zevina, 1988, A. vitrea Zevina, 1988, A. mollae Zevina, 1990, plus the

new species described here : Altiverruca laeviscuta sp. nov.

DISCUSSION. — Altiverruca is the largest verracid genus known. On the basis of gross shell morphology, the

material examined here suggests at least two groups within the genus. There are those with very laterally

compressed shells (A. galapagosa, A. laeviscuta, A. navicula, and A. nitida)', whilst the remaining species,

A. cristallina possesses a more rounded cross-section. As discussed under Verruca, any subdivision of the genus is

deferred until a wider range of taxa is able to be considered.

In his initial proposal of Altiverruca, Pilsbry stated that "Section D : Altiverruca" was characterised by

species with very short caudal appendages. Since that time, on consideration of other features, further taxa have

been included within the group (e.g. FOSTER, 1978), with some, like A. navicula and A. nitida, having very long

caudal appendages.

Distribution. — Cosmopolitan, 233- 4950 metres.

Altiverruca cristallina (Gruvel, 1907)

Fig. 2 a-h

Verruca cristallina Gruvel, 1907 : pi. 1, figs 9, 10.

Verruca cristallina laevis - BROCH, 1922 : fig. 41 a-d.

Verruca ( Altiverruca ) cristallina - ROSELL, 1989 : 24, PI. 6. figs d-i. — ROSELL, 1991 : 34.

Material EXAMINED. — New Caledonia. Biocal : stn CP 5, 21°16.49'S, 166°43.56'E, 2340 m, 11.08.1985 :

20 specimens (MNHN-Ci 2321), on pumice. — Stn CP 23, 24°45.84'S, 166°20.33'E, 2040 m, 28.08.1985 :

30 specimens, on pumice and bivalves. — Stn CP 26, 20°39.66'S, 166°27.41’E, 1618-1740 m, 28.08.1985 :

3 specimens. — Stn CP 27, 22°05.52'S, 166°26.41'E, 1850-1900 m, 28.08.1985 : 29 specimens, on pebbles. —

Stn KG 28, 23°05.47’S, 166°27.27'E, 1750 m, 28.08.1985 : 3 specimens. — Stn DW 44, 22°47.35'S, 167°14.50'E.

450 m, 30.08.1985 : 1 specimen, on coral. — Stn CP 57, 23°44'S, 166°58'E, 1490-1620 m, 01.09.1985 :

18 specimens, on pebbles. — Stn CP 58, 23°56.52'S, 166°40.55'E, 2660 m, 01.09.1985 : 1 specimen. — Stn DS 59,

23°56.21'S, 166°41 ,10'E, 2650 m, 02.09.1985 : 3 specimens. — Stn DW 66, 24°55.43’S, 168°21.67,E, 510 m,

03.09.1985 : 1 specimen. — Stn KG 71, 22°04.85'S, 167°32.70'E, 2100 m, 04.09.1985 : 1 specimen, on pumice. —

Stn CP 72, 22°10'S, 167°33’E, 2100-2110 m, 04.09.1985 : 7 specimens, on pumice.

Loyalty Islands. MUSORSTOM 6 : stn 438, 20°23.00’S, 166°20.10'E, 780 m, 18.02.1989 : 2 specimens, on

pumice.

Chesterfield Islands. MUSORSTOM 5 : stn DW 337, 19°53.80'S, 158°38.00'E, 412-430 m, 15.10.1986 :

1 specimen. — Stn DC 385, 20°53.60'S, 160°49.40'E, 745-750 m, 22.10.1986 : 5 specimens, on coral.

Wallis and Futuna Islands. MUSORSTOM 7 : stn CP 551, 12°15.3'S, 178°28.1'W, 791-795 m, 11.05.1992 :

7 specimens, on shell debris. — Stn CP 623, 12°34.2'S, 178°15.1'W, 1280-1300 m, 28.05.1992 : 1 specimen, on

pebble.

Diagnosis. — Altiverruca with carina and rostrum interlocking with up to six ribs; movable tergum with four

to five articulating ribs, at least two of which articulate with the scutum; caudal appendages short.

94

J.S. BUCKERIDGE

Description. — Shell white, almost transluscent, less compressed laterally than most Altiverruca , with

regular, sinusoidal growth lines; rostrum and carina interlock with two to three main ribs, but weaker interlocking

occurs below this with another three to four ribs; base weakly calcified.

Fixed tergum and fixed scutum interlocking with a single rib only on larger specimens.

Movable tergum quadrangular, with three diagonal ribs, increasing to four in specimens over 5 mm in length,

first rib strong, others decreasingly weaker away from diagonal.

Movable scutum triangular, relatively wide, with two diagonal ribs, which interlock with the tergum, the rib

closest to the articular margin weak to absent in smaller specimens.

Mandible tridentate, second and third teeth pectinate on the upper surface; first maxilla with two large upper

spines, notch with two small spines, lower angle with six to seven spines.

Labrum with finely toothed crest; palps pointed.

Cirrus I relatively hirsute compared with other cirri, inner ramus shorter, although it may have slighlty more

segments; cirri II-VI with inner ramus shorter; penis short, slightly hirsute terminally; caudal appendages short.

Cirri possessed the following number of segments (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

4.6 9,11 11,13 21,23 26,30 19+.30 32,35 9

4.8 13,12 11,14 19,21 29,33 32,35 32,35 9

+ incomplete ramis

Of the specimens examined, the length of the caudal appendages was close to that of the basal segment of the

pedicel of cirrus VI.

Discussion. — The considerable amount of material collected of this species has shown the differences

between Verruca crisiallina cristallina and Verruca cristallina laevis listed by Broch (1932), as insufficient for

continued separation. The strength of ribbing on the opercular plates is variable, with larger specimens developing

stronger ribs (the smaller specimens described by Broch, about 4 mm in diameter, had weaker ribs). The long

caudal appendages (with up to 25 segments) noted by Gruvel (1907) have not been recorded in either this material

or in that of Broch (1922), perhaps this anomaly could have been used to split the species, but I feel that there is

little point in following this course of action, particularly in light of Gruvel’s rather schematic drawings.

Up to six interlocking ribs between rostrum and carina can be observed in this material; however, all the

specimens described by Broch (1922) showed only three ribs, although the specimens he studied appeared to be

less than 4 mm in rostro-carinal diameter. These extra ribs, and any rib between fixed tergum and scutum, are

considered ontogenetic :

The degree of mineralisation of the base also varies, with extensive calcification only evident in larger

specimens.

Altiverruca allisoni Rao & Newman, 1972 is close to this species. Rao and Newman have distinguished A.

allisoni on the basis of its possessing seven interlocking ribs between rostrum and carina, lacking multiple

interlocking ridges between the suture of the fixed tergum and scutum, and in lacking the small beaded ridges along

the scutal margin of the rostrum. Although only six interlocking ribs were noted between rostrum and carina in

this study, analysis suggests that the first distinguishing feature (and perhaps the second), may be ontogenetic. The

best way to differentiate between the two species is on the basis of the movable scutum, which in A. cristallina

possesses an extra longitudinal rib.

This collection has extended the known depth range for this species from 1600 to 2340 metres.

Distribution. — Tropical Pacific, 233-2340 metres.

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

95

Fig. 2. — Altiverruca cristallina Gruvel, 1907 : a, complete shell (dorsal view), MNHN-Ci 2321; b, complete shell (fixed

tergal-fixed scutal view), MNHN-Ci 2321; c, movable scutum (exterior); d, movable scutum (interior); e, movable

tergum (exterior); f, mandible; g, first maxilla; h, basal portion of sixth cirrus, with penis and caudal appendage

attached. All material from Karubar, stn CP 5.

Altiverruca galapagosa Zevina, 1987

Fig. 3 a-i

Altiverruca galapagosa Zevina, 1987 : 1814, pi. 1 figs a-h.

96

J.S. BUCKERIDGE

MATERIAL EXAMINED. — New Caledonia. Biocal : stn CP 17, 20°34.54’S, 167°24.68’E, 3680 m, 14.08.1985

2 specimens (MNHN-Ci 2322). — Stn DW 46, 22°53.05'S, 167°17.08’E, 570 m, 30.08.1985 : 3 specimens.

Diagnosis. — Altiverruca with carina and rostrum interlocking with two ribs; movable scutum with one t<

two ribs articulating with the movable tergum; movable tergum with strong primary articulating rib plus ;

moderately strong, distinct secondary rib; caudal appendages short.

Description. — Shell white, thin, compressed laterally, with regularly spaced, non-prominent growth lines

rostrum and carina interlock with one rib each; apex of rostrum slightly beaked, apex of carina slightly produced

rostrum with a flat triangular area extending back from the apex to the basi-scutal margin, characterised by a singli

narrow rib. On the figured specimen, the rostrum arches "beneath" the carina. However this is more a function o

the substrate, possibly a glass sponge spicule (removed), than a characteristic of the species.

Fixed scutum and fixed tergum with apices slightly incurved, extending a little beyond the opercular plates

fixed scutum folded over on movable scutal margin, forming a zone confluent with the plane of the operculum

fixed scutum overlapping the Fixed tergum with a single extension.

Movable tergum quadrangular, with a strong arcuate, primary diagonal rib, plus a single, sharply rounded ridgi

running centrally through the scutal side to interlock with the movable scutum.

Movable scutum triangular, with two sharply rounded diagonal ribs, one of which (the more prominent

interlocks at the basi-tergal angle.

Mandible tridentate, lower tooth very small to be almost confluent with lower angle; first maxilla with twi

large upper spines and three small spines in the upper portion of the notch, lower angle with three large spines.

Cirrus I with rami subequal; penis moderately short, thin; caudal appendages very short. Cirri possessed thi

following number of segments (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

3.9 8,9 7,9 14,15 18,22 18,20 14+.13+ 9

6.9 8,9 10,13 22,23 26+,29+ 20+.21 20+.20+ 9

+ incomplete ramis

The length of the caudal appendages was slightly less than that of the basal segment of the pedicel of cirrus V

on both specimens.

Discussion. — A. galapagosa is close to Altiverruca nitida but may be distinguished on the basis of the singli

secondary rib on the movable tergum, the short caudal appendages and the laterally compressed nature of the shell.

Distribution. — New Caledonia, 570 to 3680 metres. Eastern Pacific, 3830-3850 metres.

Altiverruca laeviscuta sp. nov.

Fig. 4 a-h

MATERIAL EXAMINED. — New Caledonia. Musorstom 4 : stn DW 221, 22°46’S, 167°09'E, 535 m, 29.09.1985

1 specimen, on coral.

Type. — Holotype : MNHN-Ci 2326, rostro-carinal diameter : 3.0 mm, height : 4.6 mm, (frorr

Musorstom 4, Stn DW 221).

Diagnosis. — Altiverruca with movable tergum possessing two diagonal ribs; movable scutum flat, apico

basal line defined by growth-line defection; caudal appendages short.

Description. — Shell white, with regularly spaced, non-prominent growth lines; rostrum and carina interloci

with three ribs; apex of rostrum and carina produced.

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

97

Fig. 3. — Altiverruca galapagosa Zevina, 1987 : a, complete shell (dorsal view), MNHN-Ci 2322; b. complete shell

(fixed tergal-fixed scutal view); c, complete shell (rostral view); d, movable tergum (exterior); e, movable scutum

(exterior); f, movable scutum (interior); g, mandible; h, first maxilla; i, basal portion of sixth cirrus, with penis and

caudal appendage attached. All material from BlOCAL, stn CP 17.

Fixed scutum and fixed tergum with apices extending beyond the opercular plates; fixed scutum interlocking

with the rostrum by two ribs, and folded over on its upper margin to form a surface confluent with the plane of the

operculum.

Movable tergum quadrangular, with a strong, sharply rounded primary diagonal rib, plus a wide secondary rib

running centrally through the scutal side.

98

J. S. BUCKERIDGE

Fig. 4. — Altiverruca laeviscuta sp. nov. : a, holotype, complete shell (dorsal view), MNHN-Ci 2326; b, holotype (fixed

tergal-fixed scutal view); c, movable tergum (exterior); d, movable scutum (exterior); e, apical portion of movable

scutum (interior); f, mandible; g, first maxilla; h, basal portion of sixth cirrus, with penis and caudal appendage

attached.

Movable scutum triangular, externally rather flat, apico-basal line essentially defined by growth-line defection,

a further inflection on the tergal margin corresponds with a very weakly defined secondary tergal rib; interior with a

moderately developed depression for the adductor muscle attachment.

Mandible tridentate, lower angle pectinate with six small spines; first maxilla with two large and one small

upper spines, notch with two small spines, lower angle with one large spine and two smaller spines, hirsute.

Cirrus I with rami longer than those of cirrus II; penis short, thin; caudal appendages very short, stout. Cirri of

MNHN-Ci 2326 possessed the following number of segments (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

3.0 10,8 7,10 14,16 19,22 23,25 24,25 7

The length of the caudal appendages was approximately equal to that of the basal segment of the pedicel of

cirrus VI.

Source MNHN, Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

99

DISCUSSION. — Unfortunately only one specimen of this species has been collected, however the nature of the

movable scutum is sufficiently different to warrant specific differentiation.

Fig. 5. — Alliverruca navicula (Hoek, 1913) : a, complete shell (dorsal view), MNHN-Ci 2328; b, movable scutum

(exterior), MNHN-Ci 2327; c, movable tergum (exterior); d, mandible; e, first maxilla; f, basal portion of sixth

cirrus, with penis and caudal appendage attached; g, mandible (variety), MNHN-Ci 2332. Material ; a, from KaRUBAR,

stn CP 89; b-f, from Karubar, stn CP 38; g, from Biocal, stn CP 23.

100

J.S. BUCKERIDGE

A. laeviscuia may be distinguished from most other verrucids by the flat movable scutum. It is similar to

A. nitida but possesses a stronger secondary rib on the movable tergum and shorter caudal appendages; it is

distinguished from A. galapagosa by a longer rostro-carinal suture. The overall shell form, and soft part

morphology, is similar to that of Altiverruca gibbosa (Hoek, 1883), and although A. laeviscuta falls into an

Aliiverruca group with A. gibbosa, but it may be distinguished from that species by the movable scutum (the

absence of clear apico-basal ribbing), and the fixed scutum (an extra interlocking rib on the fixed tergal side on

shells of similar size).

ETYMOLOGY. — Morphologic : laevis = smooth (Latin) + scuta.

Distribution. — New Caledonia, 535 metres.

Aliiverruca navicula (Hoek, 1913)

Fig. 5 a-g, 16 a-b

Verruca navicula Hoek, 1913 : 134, figs 4-6. — NlLSSON-CANTELL, 1927 : 778, figs a-f.

Material EXAMINED. — Indonesia. Karubar : stn CC 21, 05°14’S, 133°00'E, 688-694 m, 25.10.1991 :

1 specimen, on glass sponge. — Stn CP 38, 07°40'S, 132°27'E, 620-666 m, 28.10.1991 : 1 specimen incomplete

(MNHN-Ci 2327). — Stn CP 52, 08°03'S, 131°48'E, 1244-1266 m, 30.10.1991 : 2 specimens, on glass sponge. —

Stn CP 87, 08°47'S, 130°49'E, 1017-1024 m, 05.11.1991 : 1 specimen. — Stn CP 89, 08°39'S, 131°08'E, 1084-

1058 m, 05.11.1991 : 5 specimens (MNHN-Ci 2328), on glass sponge. — Stn CP 91, 08°44'S, 131°05'E, 884-891 m,

05.11.1991 : 21 specimens (MNHN-Ci 2331), on glass sponge.

New Caledonia. Biocal : stn CP 23, 22°46'S, 166°20'E, 2040 m, 28.08.1985 : 8 specimens (MNHN-Ci 2332), on

glass sponge. — Stn CP 27, 23°06S, 166°26'E, 1850-1900 m, 28.08.1985 : 2 specimens, on glass sponge. —

Stn CP 75, 22°18.65'S, 167°23.30'E, 825-860 m, 04.09.1985 : 1 specimen.

Chalcal 2 : stn DW 73, 29°39.90'S, 168°38.10'E, 573 m, 29.10.1986 : 1 specimen, on coral.

Philippines. Musorstom 3 : stn 94, 13°47'S, 120°03'E, 842 m, 01.06.1985 : 1 specimen juvenile. — Stn 106,

13°47.00'S, 120°30.30'E, 668 m, 02.06.1985 : 1 specimen, on scaphopod.

Diagnosis. — Aliiverruca with carina and rostrum interlocking with single rib from each plate; movable

plates large, scutum with four articular ribs, movable tergum with six articular ribs; growth lines very distinct;

caudal appendages long.

Description. — Shell white, large, with regularly spaced, prominent growth lines; rostrum and carina

interlocking with one primary rib each, carina with a distinct furrow, running from the tergal margin to the apex,

this furrow accommodates the rostral rib at the inferior end; apices of rostrum and carina slightly produced,

rounded; rostrum and carina slightly wider than high.

Fixed scutum and fixed tergum lock together by several ribs.

Opercular plates large, characterised by strong growth lines and apico-basal ribs; apices extend beyond the shell

wall; movable tergum quadrangular, with a strong primary diagonal rib, and up to six secondary ribs on the scutal

side; movable scutum triangular, with pointed apex, primary apico-basal ridge prominent, gently arced, up to four

secondary ribs articulating with the tergum.

Mandible tridentate, lower angle divided into two portions, pectinate; first maxilla with two large upper

spines, hirsute notch and seven spines on lower angle.

Labrum with finely toothed crest; palps pointed.

Cirrus I with inner ramus longer than that of cirrus II; inner ramus of cirrus II bluntly terminated; penis long,

probosciform, terminally hirsute; caudal appendages very long. Cirri possessed the following number of segments

(r-c being rostro-carinal diameter) :

Source : MNHN , Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

101

Of the specimens examined, the length of the caudal appendages was a little more than twice that of the basal

segment of the pedicel of cirrus VI.

Discussion. — The number of interlocking ribs between rostrum and carina can be somewhat confusing. In

A. navicula, there is one primary rib, and this is so dominant that other ribs may go unnoticed; on some of the

larger specimens (e.g. > 10 mm diameter), the very base of the suture between carina and rostrum becomes

sinuous, the effect being an extension of one or two very weak apico-basal ridges. If these were stronger, they

would be termed interlocking ribs. The difference between these ridges and ribs is far less clear in A. crisiallina ,

where up to six interlocking ribs have been observed.

Some variation in the mandible has also been observed, with the lower angle having either a distinct spine (fig.

5 d) or being evenly pectinate (fig. 5 g). Of these, fig. 5 d is at variance with NILSSON-CANTELL's original

description and it is likely that the extra spine is a result of damage and subsequent repair.

The minimum depth of this species has been reduced by approximately 400 metres.

Distribution. — Tropical Pacific, 573 to 2745 metres.

Altiverruca nitida (Hoek, 1883)

Fig. 6 a-g

Verruca nitida Hoek, 1883 : 138, figs 6, 7. — Gruvel, 1905 : 177, fig. 194. — Nilsson-Cantell, 1927 : 778.

MATERIAL EXAMINED. — New Caledonia. Biocal : stn CP 23, 22°46'S, 166°20'E, 2040 m, 28.08.1985 :

1 specimen. — Stn CP 29, 23°08'S, 166°40'E, 1100 m, 29.08.1985 : 7 specimens, on pebbles. — Stn CP 30, 23°09'S,

166°41'E, 1140 m, 29.08.1985 : 7 specimens, on gastropods and pebbles. — Stn DW 36, 23°09’S, 1670ll'E, 650-

680 m, 29.08.1985 : 1 specimen, on coral.

Chesterfield Islands. Musorstom 5 : stn CP 323, 21°18.52’S, 157°57.62'E, 970 m, 14.10.1986. 2 specimens,

(MNHN-Ci 2329), on coral. — Stn CP 324, 21°15.01S, 157°51.33'E, 970 m. 14.10.1986 : 3 specimens, on coral.

Wallis and Futuna Islands. MUSORSTOM 7: stn CP 567, 11°47.0'S, 178°27.3'W, 1010-1020 m, 20.05.1992 :

1 specimen, on pebble. — Stn CP 621, 12°35.0’S, 178°11.5'W. 1280-1300 m, 28.05.1992 : 14 specimens, on glass

sponge. — Stn CP 622, 12°34.5'S, 178°10.9’W, 1280-1300 m, 28.05.1992 : 18 specimens, on glass sponge. —

Stn CP 623, 12°34.2'S, 178°15.1'W, 1280-1300 m, 28.05.1992 : 2 specimens, on glass sponge. — Stn DW 635,

13°49.0'S, 179°56.0'E, 700-715 m, 30.05.1992 : 1 specimen, on coral.

Diagnosis. — Altiverruca with carina and rostrum interlocking with two ribs; movable scutum with two ribs

articulating with one strong diagonal rib of the movable tergum; caudal appendages long.

Description. — Shell white, with regularly spaced, non prominent growth lines; rostrum and carina interlock

with two main ribs; apices of rostrum and carina slightly produced.

Fixed scutum and fixed tergum with apices extending well beyond the opercular plates; interlocking with a

single rib; fixed scutum "folded over" on its upper margin to form a surface confluent with the plane of the

operculum.

Movable tergum quadrangular, with a strong primary diagonal rib, plus a low, broadly rounded, weak ridge on

the scutal side; upper margin shorter than basal margin, but with a rounded extension that locks in behind the

movable scutum.

Movable scutum triangular, with two diagonal ribs, the more prominent interlocking with the rostrum at the

basi-tergal angle.

Mandible tridentate, lower angle pectinate with seven small spines; first maxilla with two large upper spines,

notch hirsute, lower angle hirsute, with three spines.

Cirrus I with inner ramus longer than that of cirrus II; penis long, probosciform, terminally hirsute; caudal

appendages very long. Cirri possessed the following number of segments (r-c being rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

8.3 12,21 10,24 24,28 28,29 30,32 30,36 35

Source : MNHN , Paris

102

J.S. BUCKERIDGE

Of the specimens examined, the length of the caudal appendages was approximately five times that of the basal

segment of the pedicel of cirrus VI.

DISCUSSION. — A. nitida is related to A. navicula, and can be included within a group of Altiverruca

characterised by few interlocking ribs between rostrum and carina, opercular plates each with a prominent

apicobasal ridge and long caudal appendages; as such juveniles of both species may appear similar. The bathyal

range of this species has been extended by approximately 1000 metres.

Distribution. — Tropical western Pacific, 650 to 2040 metres.

Fig. 6. — Altiverruca nitida (Hoek, 1883) : a, complete shell (dorsal view), MNHN-Ci 2329; b, articulated movable

scutum and tergum (interior); c, movable scutum (exterior); d, movable tergum (exterior); e, mandible; f, first

maxilla; g, basal portion of sixth cirrus, with penis and caudal appendage attached. All material from MUSORSTOM 5,

stn CP 323.

Source MNHN, Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

103

Genus CAMERAVERRUCA Pilsbry, 1916

Verruca "Section C": Cameraverruca Pilsbry, 1916 : 39.

Verruca (Cameraverruca) - FOSTER, 1978 : 68.

Cameraverruca - Zevina, 1978 : 1813.

Diagnosis. — Verrucids with shell moderately erect; apical cavities of fixed tergum and rostrum partitioned

off, forming a recess in the general cavity; myophore present on movable scutum.

Type. — Verruca euglypta Pilsbry, 1908 : 108. Florida, 805 metres.

Species. — 3 species are presently attributed to this genus: the type, Verruca euglypta Pilsbry, 1908,

Cameraverruca radiata (Gruvel, 1900), Canary Islands, and Cameraverruca nodiscuta sp. nov.. New Caledonia.

Discussion. — Amongst the features used by Pilsbry (1916) to define Cameraverruca were the erectness of

the shell and the presence of a myophore on the fixed scutum. In C. euglypta , the myophore was described as

"vertical, partition-like". The terminology "vertical" is at first instance confusing, fortunately however, the interior

of the fixed scutum is figured in Pilsbry (1916, pi. 3, fig. 2a) where, although the myophore is shown to be

vertically depending, it is aligned horizontally, and is thus similar to other myophores in other Cameraverruca and

Metaverruca. The term "erect" may also lead to confusion : C. nodiscuta sp. nov. and C. radiata are not as erect as

most Altiverruca species, but are certainly more erect than those of Metaverruca and Verruca.

I have also had an opportunity to examine Gruvel's types of V. radiata and confirm that this possesses both

an "erect" shell and a myophore on the fixed scutum.

Stratigraphic range. — Recent.

Distribution. — Atlantic and western Pacific, 380 - 912 metres.

Cameraverruca nodiscuta sp. nov.

Fig. 7 a-j

Material EXAMINED. — Chesterfield Islands. MUSORSTOM 5 : stn DW 300, 22°48.27'S, 159°23.94'E, 450 m,

11.10. 1986: 1 specimen, on coral.

Loyalty Islands. MUSORSTOM 6 : stn DW 397, 20°47.35'S, 167°05.17'E, 380 m, 12.02.1989 : 2 specimens.

Types. — Holotype : MNHN-Ci 2315, r-c diameter : 4.5 mm, height : 6.6 mm, (from Musorstom 5,

Stn DW 300).

Paratypes : MNHN-Ci 2316-2317, (from MUSORSTOM 6. Stn DW 397).

Diagnosis. — Cameraverruca with carina and rostrum interlocking with up to six ribs; movable scutum with

three articulating ribs, the uppermost extended, node-like; fixed scutum with semicircular, upwardly concave

myophore; caudal appendages short.

Description. — Shell off-white to palest pink, with regularly spaced, non prominent growth lines; carina and

rostrum interlocking with up to six ribs, apex of carina and rostrum slightly produced; rostrum with apical cavity

weakly partitioned off; base of plates thickened, slightly inflected; carina and fixed tergum interlocking with three

ribs; rostrum and fixed scutum interlocking with four ribs; basis calcareous.

Fixed scutum and fixed tergum with apices extending beyond the opercular plates, both folded over on their

opercular margins to form a zone confluent with the plane of the operculum; fixed scutum with semicircular,

upwardly concave myophore; fixed tergum with apical cavity clearly partitioned off, forming a recess in the general

cavity.

104

J.S. BUCKERIDGE

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

105

Movable scutum rather narrow, triangular, with a moderately strong, arcuate, apico-basal ridge interlocking at

the basi-tergal angle with the movable tergum, two further ribs articulate with the movable tergum, the uppermost

of which is extended and node-like, to slot in between the second and third ribs on the tergum, interior with

moderatly deep adductor muscle pit.

Movable tergum with a strong apico-basal rib, plus a moderately strong and distinct secondary rib; scutal side

further characterised by weak longitudinal striae; interior smooth except for a narrow zones of growth lines on

upper and fixed tergal margins.

Mandible tridentate, lower angle pectinate; first maxilla with two strong upper spines, three smaller spines in

the notch and four spines on an hirsute lower angle.

Cirrus I with anterior ramus longer, cirrus II with posterior ramus longer; penis thin, probosciform, about

twice the length of the basal segment of the pedicel of cirrus VI; caudal appendages short, slightly less than that of

the basal segment of the pedicel of cirrus VI. Cirri possessed the following number of segments (r-c being rostro-

carinal diameter) :

r-c (mm) I 0 III IV V VI c.a.

4.5 12,9 9,12 15,16 15,20 21,22 21,21 8

Discussion. — C. nodiscuta may be distinguished from other Cameraverruca by the narrow, nodose scutum.

It extends the geographic range of the genus from the Atlantic to the Pacific Ocean.

Etymology. — Morphologic : from nodus = node (Latin) + scuta.

Distribution. — New Caledonia, 380 to 450 metres.

Genus BROCHIVERRUCA Zevina, 1993

Diagnosis. — Verrucids with apex (umbo) of both carina and rostrum removed from the margin of the plates;

fixed scutum with or without a myophore.

Type. — Brochiverruca margulisae Zevina. 1993 : 10. Mozambique Channel, Indian Ocean, 935-950 metres.

SPECIES. — Three species are here attributed to this genus, the type, Brochiverruca margulisae Zevina,

Brochiverruca dens (Broch) and Brochiverruca polystriata sp. nov.. New Caledonia.

Discussion. — broch (1932) included B. dens within Rostratoverruca, on the basis that it was most closely

allied to taxa within that group. The inclusion however was tentative, and Broch suggested that future

investigations may warrant separation of the species from Rostratoverruca. Certainly the nature of the rostrum and

carina and the cubic, "tooth-like" appearance make Brochiverruca one of the most easily distinguished verrucid

genera. It is considered likely that future workers may wish to further divide Brochiverruca. particularly on the

basis of the presence of a myophore on the fixed scutum of the new species. At this juncture, I consider the

placement of these species within Brochiverruca, based on the similarity of overall shell morphology, to be a

tentative solution to the problem.

Stratigraphic range. — Recent.

Distribution. — Western Pacific and Western Indian Ocean, 348-950 metres.

Brochiverruca dens (Broch. 1932)

Verruca (Rostratoverruca? ) dens Broch, 1932 : 47, fig. 17 a-g.

Material EXAMINED. — Indonesia. Karubar : stn CP 69, 08°42'S, 131°53'E, 356-368 m, 02.11.1991 :

3 specimens without opercula and soft parts, on coral.

106

J.S. BUCKERIDGE

Diagnosis. — Shell white, finely sculptured with delicate longitudinal striae crossed by fine, distinct, growth

lines; rostrum and carina each with the apex separated from the upper margin by one third the distance from the

basal margin; fixed scutum without a myophore.

Description. — Shell white, tooth-like, exterior very finely sculptured with delicate longitudinal striae

crossed by distinct, but not prominent growth lines.

Rostrum and carina each with the apex separated from the upper margin by one third the distance from the basal

margin; the margin between both plates is characterised by up to 13 fine, non-prominent, articulating ribs. Fixed

scutum without a myophore. Soft parts and operculum absent from this collection.

In his original description, Broch noted that this species possessed a broad, trapezoid movable tergum, with

up to 12 delicate ribs articulating with the movable scutum; the movable scutum is about half the width of the

tergum, internally with a shallow adductor muscle scar. Of the soft parts, it is significant to note that the labrum

is smooth, and the palps short and narrow; B. dens is characterised by a relatively "stout penis, a little longer than

the caudal appendages", which themselves are long, having 25 segments (about half as long as the sixth cirrus).

Discussion. — This species often has a surface texture closely approximating that of the substrate. A large

specimen from Karubar, Stn CP 69, with a rostro-carinal diameter of 6.8 mm, attached to a smooth surfaced

stylasterid coral, showed little evidence of the normally characteristic fine sculpture. The specimen however, was

represented by a shell wall only, and may have been subsequently modified by the coral and/or the physical

environment.

Distribution. — Western Pacific. 348-368 metres.

Brochiverruca polystriata sp. nov.

Fig. 8 a-j

Material EXAMINED. — New Caledonia. Biocal : stn DW 51, 23°05.27’S, 167°44.95'E, 700 m, 31.08.1985 :

1 specimen with opercula and soft parts absent, on Ellanospammia. — Stn DW 66, 24°55.43'S, 168°21.67'E, 510 m

03.09.1985 : 1 specimen juvenile.

Chalcal 2 : stn DW 77, 23°38.35'S, 167°42.68'E. 435 m. 30.10.1986 : 13 specimens, most without opercula and

soft parts, on coral.

Types. — Holotype : MNHN-Ci 2292, rostro-carinal diameter : 4.3 mm, height : 4.5 mm, (from Chalcal 2

Stn DW 77).

Paraiypes ; MNHN-Ci 2293, (from Biocal, Stn DW 66), MNHN-Ci 2294-2297 (from Chalcal 2.

Stn DW 77); UNITEC Cat. No. CAX 101, (from Chalcal 2, Stn DW 77); USNM (from Chalcal 2

Stn DW 77).

Diagnosis. — Shell finely sculptured with delicate longitudinal striae crossed by growth lines to give a finely

nodose surface texture; rostrum and carina each with the apex separated from the upper margin by slightly less than

a third the distance from the basal margin; fixed scutum with myophore.

Description. Shell white, tooth-like, exterior finely sculptured with numerous delicate longitudinal striae

crossed by distinct growth lines with a resultant nodose effect.

Rostrum and carina each with the apex separated from the upper margin by slightly less than one third the

distance from the basal margin; the margin between both plates is characterised by up to 21 fine, non-prominent,

articulating ribs.

Fixed scutum protruded along rostral margin to form a covered portion on the upper surface adjacent to the

movable scutum; articulating with the fixed tergum with up to seven delicate ribs, and with the rostrum with up

to 13 ribs; with a clear lunulitiform myophore on inner surface (Fig. 80; fixed tergum and carina articulating with

up to seven ribs.

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

107

Fig. 8. — Brochiverruca polystriala sp. nov. : a, holotype, complete shell (dorsal view), MNHN-Ci 2292; b, holotype,

complete shell (fixed tergal-fixed scutal view); c, articulated movable scutum and tergum (interior); d, movable tergum

(exterior); e, movable scutum (exterior); f, shell with opercula removed, showing pendant myophore on fixed scutum;

g, complete shell (fixed tergal-fixed scutal view); h. mandible; i, first maxilla; j, basal portion of sixth cirrus, with

penis and caudal appendage attached. Material : a-e, from Chalcal 2, stn DW 77; f-g, from BlOCAL, stn DW 51;

h-j, from Biocal, stn DW 66.

108

J.S. BUCKERIDGE

Movable tergum of low relief, quadrangular lo kile shaped, with up lo seven narrow, delicate ribs articulating

with the movable scutum.

Movable scutum quadrangular, less than half the width of the tergum. with a weak apico-basal ridge and five to

six very weak ribs articulating with the movable tergum: interior with shallow adductor muscle depression

positioned immediately below the apical growth lines.

Mandible tridenlate, lower angle pectinate; first maxilla with two larger spines above the notch and four spines

on the lower part; penis long and relatively stout basally, at least twice the length of the caudal appendages,

terminally hirsute. Cirri with subequal rami, with the following number of segments (r-c being rostro-carinal

diameter) :

r-c (mm)

I II III IV V VI c.a.

8,6 6,7 10,13 14,16 17,19 19,20 9

11.11 8,10 12,13 15,17 15,19 20,20 8

2.6

4.3

Of the three specimens examined, the length of the caudal appendages was about twice that of the basal

segment of the pedicel of cirrus VI.

Living material from Chalcal 2, Stn 77 included shells with a pale pink colouration, but this disappeared

when the darkish soft parts were removed.

DISCUSSION. — When this material was first encountered, I identified it tentatively as Verruca

(Rostraloverruca? ) dens Broch; the nature of the apices of the rostrum and carina being considered rather diagnostic.

The presence of a myophore on the fixed scutum, plus the surface texture, however marks this form as distinct. In

his original description of V. dens, Broch did not observe a myophore. The possibility that the myophore may

have been lost, or reduced during growth, was considered, and in this instance further specimens of V. dens were

studied, (MNHN-Ci 2154 ; from La Reunion). These specimens did not possess a myophore on the fixed scutum.

Further differences in the soft parts (see below), strengthened this as a distinct species.

Clearly this species is most closely related to B. dens, but may be distinguished by a slightly coarser external

texture, the scutal myophore, the significantly shorter caudal appendages and the longer penis (It has been noted in

Barnes (1992) however, that the length of the penis in some species may be related to the breeding condition).

Etymology. — Morphologic : poly + striatus = many + striae (Latin).

Distribution. — New Caledonia. 435-700 metres.

Genus META VERRUCA Pilsbry, 1916

Verruca "Section A": Metaverruca Pilsbry, 1916 : 21.

Verruca ( Metaverruca ) - FOSTER, 1978 : 68.

Metaverruca - Zevina, 1978 : 1812.

Diagnosis. — Verrucids with apices of carina and rostrum marginal; fixed scutum with myophore; operculum

parallel to base; base inflexed, thickened.

TYPE. — Verruca coraliophila Pilsbry, 1916 : 21. Between Bahamas and Cape Fear. 506-794 metres.

Species. Eight species arc presently attributed to this genus, this includes a revised Metaverruca recta

(within which Verruca capsula Hoek, 1913; Verruca coraliophila Pilsbry, 1916; Verruca halotheca Pilsbry, 1907;

Verruca linearis Gruvel, 1900; Verruca magna Gruvel, 1901; Verruca sculpta Aurivillius, 1898, are synonymised).

Metaverruca corrugata Broch. 1922, Metaverruca lepista Zevina. 1987, Metaverruca seriola Zevina. 1987. plus the

tour new species described here : Metaverruca defayeae sp. nov.. Metaverruca notfolkensis sp. nov.. Metaverruca

pacifica sp. nov., and Metaverruca plicata sp. nov.

Stratigraphic range. — Lower Miocene (New Zealand) to Recent.

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

109

Distribution. — Cosmopolitan, 167- 4100 metres.

Metaverruca defayeae sp. nov.

Fig. 9 a-g

MATERIAL EXAMINED. — Volcanoes south of Vanuatu, VoLSMAR : stn DW 55, 20°59.20'S, 170°01.90’E,

710 m, 5.07.1989 : 1 specimen.

Loyalty Islands. Musorstom 6 : stn DW 468, 21°05.86'S, 167°32.98'E, 600 m, 22.02.1989 : 1 specimen. —

Stn DW 480, 21°08.50'S, 167°55.98'E, 380 m, 22.02.1989 : 1 specimen, on pebble. — Stn DW 486, 20°2 1.40'S,

167°47.65'E, 370 m, 23.02.1989 : 1 specimen, on gastropod.

Wallis and Futuna Islands. MUSORSTOM 7 : stn DW 537, 12°30'S, 176°41'W, 325-400 m, 16.05.1992 :

1 specimen, on pebble. — Stn DW 540, 12°26.7'S, 177°28'W, 600 m, 17.05.1992 : 1 specimen, on pebble. —

Stn DW 541, 12°27'S, 177°28'W, 500-505 m, 17.05.1992 : 1 specimen, on pebble. — Stn DW 547, 12°26'S,

177°26'W, 455 m, 17.05.1992 : 1 specimen, on pebble. — Stn DW 586, 13°H’S, 176°13'W, 510-600 m, 22.05.1992 :

I specimen, on pebble. — Stn DW 594, 12°31'S, 174°20'W, 510-600 m, 24.05.1992 : 1 specimen, on pebble. —

Stn DW 604, 13°21'S, 176°08'W, 415-420 m, 26.05.1992 : 1 specimen, on coral.

Fig. 9. — Metaverruca defayeae sp. nov. : a, holotype, complete shell (dorsal view), MNHN-Ci 2298; b, holotype,

complete shell (fixed tergal-fixed scutal view); c, movable scutum (interior); d, movable tergum (interior);

e, mandible; f, first maxilla; g, basal portion of sixth cirrus, with penis and caudal appendage attached. Material :

a-d, from Volsmar, stn DW 55; e-g, from Musorstom 7, stn DW 537.

Types. — Holotype : MNHN-Ci 2298, rostro-carinal diameter ; 10.4 mm, height : 3.4 mm, (from VOLSMAR,

Stn DW 55).

110

J.S. BUCKFRIDGE

Paraiypes : MNHN-Ci 2299. (from MUSORSTOM 7. Stn DW 537); MNHN-Ci 2300. (from MUSORSTOM 7.

Stn DW 540); UNITEC Cat. No. CAX 102. (from MUSORSTOM 7, Stn DW 541); USNM, (from MUSORSTOM 7

Stn DW 547).

DIAGNOSIS. — A large Metaverruca with movable tergum and scutum having four articular ribs; exterior

smooth with closely spaced growth lines; mandible quinquedentate, caudal appendages short, less the length of the

basal segment of the pedicel of cirrus VI.

Description. — Large sized Metaverruca with shell white, low conic, sides steep; operculum sub-parallel to

base, orifice D-shaped; exterior generally smooth, with relatively closely spaced, concentric growth lines; rostrum

and carina articulating with up to five ribs; basis membranous.

Fixed scutum with a well formed myophore for adductor muscle attachment; basal margin of shell thickened

and inflected on larger specimens; fixed tergum a little less than the width of fixed scutum, with basal margin

thickened and inflected on larger specimens.

Movable tergum quadrangular, with a well defined apico-basal rib, and three secondary ribs, interlocking with

the movable scutum.

Movable scutum triangular, relatively narrow, with three closely spaced ribs interlocking with the tergum and a

further adjacent apico-basal rib terminating at the rostro-carinal suture; internally with a weak depression for

adductor muscle attachment; both opercular plates together form a slightly convex rostro-carinal hinge.

Mandible quinquedentate. last tooth very small, lower angle hirsute, with one main spine; first maxilla with

two large upper spines, notch with four small spines, lower angle hirsute, with five spines.

Cirri particularly fine, possessing the following number of segments (r-c being the rostro-carinal diameter) :

The caudal appendages are very short and thin, with length a little less than that of the basal segment of the

pedicel of cirrus VI; penis vestigial.

The soft parts of larger specimens possessed a distinct pink coloration, particularly in the vicinity of the

mouthparts.

Discussion. — M. defayeae is very close to M. recta, differing essentially in the number of ribs present on the

opercular plates. However, in all specimens studied, the spacing of the growth lines was also found to be distinct,

being arranged closer in M. defayeae. At first it appeared that these two factors may fall within the morphological

range of M. recta, with growth line spacing being a function of growth rate (and thus environment). However this

is less likely to be the case as both species have been recorded from the same station. The soft parts also assist in

distinguishing this species, particularly in respect of the mandible, which has five teeth, (although the fifth is

rather small), this character has not been encountered in any other verrucids studied in the collections. Further,

unlike M. recta , the rami of cirrus II are either longer or equal to those of cirrus I.

Etymology. Named for Dr Danielle Defaye, Museum national d'Histoire naturelle, Paris, for her

assistance during this research project.

Distribution. — Western Pacific, 370-710 metres.

Metaverruca norfolkensis sp. nov.

Fig. 10 a-k

MATERIAL EXAMINED. — New Caledonia. Biocal : stn CP 29, 23°08'S, 166°40'E, 1100 m, 29.08.1985 ■

1 specimen, on pumice.

Smib 5 : stn DW 93, 22°20.00'S, 168°42.30'E, 255 m, 11.09.1989 : 1 specimen.

Source : MNHN. Paris

VF.RRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

111

Types. — Holotype : MNHN-Ci 2301, rostro-carinal diameter : 4.1 mm, height : 1.8 mm, (from Biocal,

Stn CP 29).

Paratype : MNHN-Ci 2302, (from Smib 5. Stn DW 93).

Diagnosis. — A medium size Metaverruca with movable tergum and scutum having four articular ribs;

exterior smooth with moderately spaced growth lines; caudal appendages short, less the length of the basal

segment of the pedicel of cirrus VI.

Fig. 10. — Metaverruca norfolkensis sp. nov. : a, holotype, rostrum and carina, MNHN-Ci 2301; b, movable tergum

(exterior); c, movable tergum (interior); d, movable scutum (interior); e, movable scutum (exterior); f, fixed scutum

showing myophore (interior); g, fixed scutum (exterior); h, fixed tergum (exterior); i, mandible; j, first maxilla;

k, basal portion of sixth cirrus, with penis and caudal appendage attached. All material from MNHN-Ci 2301,

Biocal, stn CP 29.

112

J.S. BUCKERIDGE

Description. — Medium sized Metaverruca with shell while, low conic, sides steep; operculum sub-parallel

to base, orifice D-shaped; exterior generally smooth, with relatively closely spaced concentric growth lines;

rostrum and carina articulating with up to three ribs; basis membranous.

Fixed scutum quadrangular, with a well formed myophore for adductor muscle attachment; basal margin of

shell thickened and inflected on larger specimens; fixed tergum a little narrower than the fixed scutum.

Movable tergum quadrangular, with a well defined apico-basal rib. and three secondary ribs, interlocking with

the movable scutum; articular margin moderately concave; movable scutum triangular, with three ribs interlocking

with the tergum and a further adjacent curved, apico-basal rib terminating at the rostro-carinal suture; occludent

portion lacking ribbing; internally with a moderately strong depression for adductor muscle attachment; both

opercular plates together form an almost straight rostro-carinal hinge.

Mandible tridentate, lower angle pectinate; first maxilla with two large upper spines, notch with four small

spines, lower angle hirsute, with four large spines.

Cirri possess the following number of segments (r-c being the rostro-carinal diameter) ;

r-c (mm) I II HI IV V VI c.a.

4- l 12,9 7,8 1 1,13 20,21 23,24 24,27 7

5- 0 13.1 1 9,1 1 14,15 20,22 22,24 22,25 9

The caudal appendages are very short and thin, with length a little less than that of the basal segment of the

pedicel of cirrus VI; penis thin, short, length a little less than the caudal appendages.

DISCUSSION. — Metaverruca norfolkensis is very close to M. recta, differing essentially in the number of ribs

present on the opercular plates. It may be distinguished from M. defayeae by the mandible, shorter caudal

appendages, more widely spaced growth ridges, a broader scutum and a more quadrangular tergum.

Etymology. — Geographic : Norfolk Ridge, southwest Pacific Ocean.

Distribution. — New Caledonia. 255-1100 metres.

Metaverruca pacifica sp. nov.

Fig. 1 1 a-g, 16 c-d

Material EXAMINED. — New Caledonia. Musorstom 4 : stn CP 198, 18°49.40'S, 163°18.80'E, 585 m

20.09.1985 : 12 specimens, on bivalves.

Loyalty Islands Musorstom 6 : stn CP 438. 20°23.00'S, 166°20.10'E, 780 m, 18.02.1989 : 1 specimen, on

m’ 540 m’ 21 02 ,989 : 3 specimens, on gastropod and Coronuta. -

j C, 7 com ' E' 575 m' 21 02 1989 : 1 specimen, on Coronula. — Stn DW 483, 21°19.80'S,

lo/ 4 /.oU b, 6UU m, 23.02.1989 : 1 specimen, on brachiopod.

Chesterfield Islands. Musorstom 5 : stn DW 341, 19°45.90'S, 158°43.37’E, 620-630 m. 16 10 1986 •

2 Admens, on ^‘"oderm. _ Stn DW 342, 19°43.50'S. 158M7.72E. 660 m, 16.10.1986 : 4 specimens, on bivalve!

Sin' nMW63iq' S j !i8ifi°cE'^5'700 m' l9'101986 : 125+ specimens, on shell detritus inch Nautilus. —

e. rr ,, ’ -cfcfo'o-e' 675 m' 1910'1986 : 29 specimens, on spatangoids and annelid tubes. —

1Q,^ACRn'R5^S89osVfnmS'ass807o'OOE' 7'° m’ 1910'1986 : 50+ specimens, on Nautilus, etc. — Stn CC 367,

19 36.80 S, 158 53.20 E, 855-830 m, 19.10.1986 : 35 specimens (MNHN-Ci 2330), on pebbles

Corail 2 : stn CP 17, 20°48.14’S, 160°57.14'E, 500 m, 21.07.1988 : 1 specimen.

Types. — Holotype : MNHN-Ci 2303, rostro-carinal diameter : 6.5 mm, height ; 3.9 mm (from

Musorstom 5, Stn CP 363).

Paratypes : MNHN-Ci 2304, (from Corail 2, Stn CP 17), MNHN-Ci 2305, (from Musorstom 4,

Stn CP 198), MNHN-Ci 2306-2309, (from Musorstom 5, Stn CP 363); UNITEC Cat. Nos. CAX 103-105.

(from Musorstom 5, Stn CP 363); USNM. (from Musorstom 5, Stn CP 363).

Diagnosis. — Metaverruca with movable tergum and scutum having four articular ribs; exterior generally

smooth with widely spaced growth lines; caudal appendages more than twice the length of the basal segment of the

pedicel of cirrus VI.

Source : MNHN. Paris

VF.RRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

113

Fig. 11. — Metaverruca pacifica sp. nov. : a, holotype, complete shell (dorsal view), MNHN-Ci 2303; b. articulated

movable scutum and tergum (interior); c, movable scutum (exterior); d, movable tergum (exterior); e. mandible;

f. first maxilla; g, basal portion of sixth cirrus, with penis and caudal appendage attached. Material : a, g. from

Musorstom 5. stn CP 363; b-d, from Corail 2, stn CP 17; e-f, from MUSORSTOM 4, stn CP 198.

Description. — Medium sized Metaverruca with shell white, low conic, sides steep; operculum sub-parallel to

base. D-shaped; exterior generally smooth, with well spaced concentric growth lines; rostrum and carina

articulating with up to five ribs. Rostrum with a distinctly ribbed zone arcing toward the base of the movable

scutum. In large specimens, this zone can have up to four separate ribs, and can therefore be relatively diagnostic

for the species; basis membranous.

Fixed scutum with a well formed myophore for adductor muscle attachment; basal margin of shell thickened

and inflected on larger specimens; fixed tergum a little over half the width of fixed scutum, on larger specimens

basal margin thickened and inflected.

Movable tergum quadrangular, with three large and one or more narrow articular ridges, movable scutum

triangular, with corresponding ridges externally and a moderate pit for adductor muscle attachment internally; both

plates together form a straight rostro-carinal hinge.

Mandible tridentate, lower angle pectinate and divided into two portions; first maxilla with two prominent

upper spines, a notch without spines and with four spines on a hirsute lower angle; penis moderately short.

114

J.S. BUCKERIDGE

approximately two and a half times the length of the basal segment of the pedicel of cirrus VI. Cirrus I with

anterior ramus particularly hirsute. Cirri possessed the following number of segments (r-c being the rostro-carinal

diameter) :

Of the specimens examined, the length of the caudal appendages was consistently more than twice that of the

basal segment of the pedicel of cirrus VI.

Discussion. — M. pacifica is characterised by the presence of four prominent articular ribs on the opercular

plates, and the long caudal appendages: all except small specimens have a rostrum with a distinct ribbed zone

arcing back to the base of the movable scutum, clearly differentiating this species from M. recta, M. norfolkensis

and M. defayeae. On rare occasions, excessive development of this zone behind the apex of the rostrum may cause

the rostrum to move slightly in from the plate margin, showing similarity to Rostratoverruca. It is important to

note that of the more than 100 specimens studied only one possessed an apex clearly removed from the margin,

and this only by a small distance; further, the fixed scutum on this species has a myophore.

Etymology. — Geographic : Pacific Ocean.

Distribution. — New Caledonia. 500-830 metres.

Metaverruca plicata sp. nov.

Fig. 12 a-i

,<i^lAJERIAL EXAMINED- — Loyalty Islands. MUSORSTOM 6 : stn DW 405, 20°29.75'S, 166°41 00'E 520 m

14.02.1989 : 1 specimen. — Stn DW 468, 21°05.86'S, 167°32.98'E, 600 m, 21.02.1989 : 1 specimen.

Types, — Holotype : MNHN-Ci 2310, rostro-carinal diameter : 8.9 mm. height : 3.6 mm, (from Musorstom

6. Stn DW 405).

P aratype : MNHN-Ci 2311, (from Musorstom 6, Stn DW 468).

DIAGNOSIS. — A medium to large Metaverruca with strong external ribbing; movable tergum and scutum

having three articular ribs; movable scutum moderately narrow; caudal appendages long, about two and a half

times the length of the basal segment of the pedicel of cirrus VI.

description. — Medium to large sized Metaverruca with shell white, low conic, sides steep; operculum sub¬

parallel to base, orifice D-shaped; exterior characterised by strong, sharp vertical ribbing, with relatively closely

spaced concentric growth lines, secondary ribs develop in the interstices of the primary ribs as the shell diameter

increases, thus maintaining regularity of spacing; rostrum and carina articulating with up to six ribs- basis

membranous.

Fixed scutum quadrangular, with a well formed myophore for adductor muscle attachment, articulating with the

fixed tergum with one rib and with rostrum with four ribs; basal margin of shell thickened and inflected on larger

specimens; fixed tergum of similar width to the fixed scutum, articulating with the carina with four ribs.

Movable tergum quadrangular, with a well defined apico-basal rib, and two secondary ribs, interlocking with

the movable scutum: movable scutum triangular, relatively narrow, with two ribs interlocking with the tergum

and a further adjacent curved, apico-basal rib terminating at the rostro-carinal suture; occludent portion with fine

longitudinal striae crossing growth lines; internally with a moderately weak depression for adductor muscle

attachment; both opercular plates together form an almost straight rostro-carinal hinge.

Source MNHN. Paris

VF.RRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

115

Mandible tridentate, lower angle pectinate, divided into two parts; first maxilla with a prominent upper spine

and three smaller adjacent spines, lower angle with four long and three shorter spines.

The holotype cirri possess the following number of segments (r-c being the rostro-carinal diameter) :

r-c (mm) I II III IV V VI c.a.

4.1 12,16 10,13 15,17 23,25 28,32 31,31 27

The caudal appendages are rather long, almost 3 times that of the basal segment of the pedicel of cirrus VI;

penis vestigial.

Fig. 12. — Metaverruca plicata sp. nov. : a, holotype, complete shell (dorsal view), MNHN-Ci 2310; b, holotype,

portion of base (ventral view); c, articulated movable scutum and tergum (interior); d, movable tergum (exterior);

e, movable scutum (exterior); f, holotype, complete shell (fixed tergal-fixed scutal view); g, mandible; h, first

maxilla; i, basal portion of sixth cirrus, with penis and caudal appendage attached. All material from MusORSTOM 6,

stn DW 405.

Discussion. — M. plicata is very distinct species, showing greatest similarity to Metaverruca corrugata

(Broch, 1932), from which it differs in having significantly longer caudal appendages, a narrower movable scutum.

116

J.S. BUCKERIDGE

a grealer number of articulaling ribs on ihe wall plates and a lower shell profile. Like most Metavenuca, the rami

of cirrus I possess more segments than those of cirrus II.

Etymology. — Morphologic : in allusion to strong external ribbing, plicatus = ribbed or folded (Latin).

Distribution. — Loyalty Islands. 520-600 metres.

Metaverruca recta ( Auri villius, 1898)

Fig. 13 a-f

Verruca recta Aurivillius, 1898 : 195.

Verruca sculpta Aurivillius, 1898 : 197.

Verruca linearis Gruvel, 1900 : 243; 1902 ; 107, pi. 5 figs 1 1, 12 .

Verruca magna Gruvel, 1901 : 261; 1092 : 109. pi. 5 figs 1. 2 .

Verruca haiotheca Pilsbry, 1907 ; 188, pi. 12 figs 9, 10.

Verruca capsula Hock, 1913 : 130, pi. 12 figs 1-3, pi. 13 figs 1-4.

Verruca coraliophiia Pilsbry, 1916 : 21, pi. 1 figs 1-5.

Verruca cookei - Rosell. 1989 ; 299, pi. 11 figs r.s.u.v; 1991 : 33. Non Pilsbry, 1927.

Material EXAMINED. — New Caledonia. Biocal : stn DW 8, 20°34.35'S, 166°53.90'E 435 m P 08 1985 •

1 specimen, on pebble. — Stn CP 23. 22°46'S. 166°20'E, 2040 m, 28.08.1985 : 27 specimens (MNHN-Ci 2312) on

pebbles and bivalve. — Stn CP 26. 20°40'S, 166°27’E, 1618-1740 m, 28.08.1985 ; 2 specimens (MNHN-Ci 2313)’. —

Stn CP 27, 22°06S, 166°27E, 1850-1900 m, 28.08.1985 ; 20 specimens, on pebbles. — Stn CP 29, 23°08'S, 166°40'E

1100 m, 29.08.1985 : 6 specimens, on pumice. — Stn CP 30, 23°09'S, 166°41'E, 1140 m, 29.08.1985 : 6 specimens’

on pumice. — Stn CP 52, 23°06'S, 167’47’E, 540-600 m, 31.08.1985 : 1 specimen, on coral. — Stn CP 57. 23°44'S,

1 66°58'E, 1490-1620 m, 01.09.1985 ; 17 specimens, on pumice. — Stn CP 62, 24°19'S 167'49’E 1395-1410 m

02.09.1985 : 1 specimen, on pumice. — Stn CP 72, 22°10'S, 167°33'E, 2100-2110 m, 04.09.1985 : 3 specimens on

bivalve and pumice. — Stn DW 81, 24°29.31’S, 166°46.56'E, 430-470 m, 31.08.1985 : 1 specimen.

Chalcal 2 : stn DW 72, 24°54.50S, 168°22.30'E, 527 m, 29.10.1986 : 1 specimen, on bivalve

Biogeocai. : stn CP 272, 21°00'S, 166°57'E, 1615-1710 m, 20.04.1987 : 1 specimen

Smtb 5 : stn DW 86, 22°19.80'S, 168°42.80'E, 320 m. 11.09.1989 : 1 specimen, on coral

Smib 8 : stn DW 146. 24°55.20’S, 168°21.70'E, 514-522 m, 27.01.1993 : 1 specimen, on sponge.

Chesterfield Islands. Musorstom 5 : stn DW 299. 22°47.70'S, 159°23.70'E, 360-390 m 11 10 1986 •

4 seamens, on Stenohelia. - Stn CP 323, 21°18.52'S, 157‘57.62'E. 970 m, 14.10.1986 : 4 specimens, on pumice!

coo " ^ 3-4- “ 3 01 S, 157 51.33 E, 970 m, 14.10.1986 : 2 specimens, on pumice. — Stn DW 335, 20°03.24'S,

,°™E' ,3'5 m- 15101986 : 4 specimens, on coral. — Stn DC 345, 19°39.70'S, 158°32.40’E, ’ 305-3 10 m!

16. 1U. 1986 : 1 specimen, on pebble.

Loyalty Islands. Musorstom 6 : stn DW 413, 20°40.10'S. 167°03.50'E, 463 m, 15.02.1989 : 6 specimens, on

7^" ™ ,4’7o,;(ll4E80'S- 167°03-65'E’ 283 m. 16.02.1989 : 11 specimens, on brach.opods. -

StnDW 439, -0 46.40S, 167 17.40 E, 288 m, 19.02.1989 : 1 specimen. — Stn DW 452, 21°00.30’S, 167°25.50'E

300 m, 20.02.1989 : 6 specimens, on pebbles. — Stn DW 456, 21°00.71’S, 167°26.35'E 240 in ^0 02~1989 ’

8 specimens, on spatangoids. — Stn DW 457, 21°00.42'S. 167°28.71'E, 353 m, 20.02.1989 : 1 specimen on pebble’

~ Sln ^ 472- 21°08.60'S, 1 67°54.70'E, 300 m, 22.02.1989 : 1 specimen. — Stn DW 480, 21°08.50'S. 167°55 98'E

, 0 m- 22.02.1989 : 1 specimen, on pebble. — Stn CP 481, 21°21.85'S, 167°50.30’E, 300 m, 23 0’’ 1989 •’

1 specimen. - Stn DW 482, 21°21.50'S. 167°46.80'E, 375 m. 23.02.1989 : 2 specimens, on coral.

Wallis and Futuna Islands. Musorstom 7 : stn DW 502, 14°19.8'S, 178°06.5'W, 516-535 m 11 05 1992 •

4 specimens on pebbles. ~ Stn DW 8 511 . I4«14.0’S. 17S°11.5'W, 400-450 m, 12.05.1992 : 6 specimens, on pebbles!

,7 " n’W53W ann '6 S’ 580-600 m, 16.05.1992 : 1 specimen, on pebble. — Stn DW 537, 12°30.0'S,

l-rns looo o25'400 m’ 16-05-1992 : 1 specimen, on pebble. — Stn DW 540, 12°26.7'S, 177°28.4'W, 600m,

17.05. 1992 : 2 specimens, on pebbles. - Stn DW 541. 12°26.7'S, 177°28.0’W, 500-505 m. 17.05.1992 : 1 specimen,

“ m ™ id 'SStfS; “-810 >8'051992 = 1 - S,„ CP 551,

77°2738SW vLinnW' ' !,8-05 1.992 : 59 specimens, on shell fragments. - Stn CP 552, 12°15.7'S,

m’ 1805-1992 : 2 specimens, on shell fragments. — Stn DW 555, 11047.5’S. 178°19 2’W 540-

D42 m, 19;05 .1992 : 1 specimen, on shell. — Stn DW 557, 11°48.1'S, 178°18.2'W. 600-608 m, 19 05 1992 • ■>

specimens (MNHN-Ci 2314), on pebbles. — Stn DW 558, 11°49.9'S, 178°18.9'W, 635 m, 19.05.1992 : 1 specimen on

TXtrT 17?°£lw S7'702 m' l9051"2 1 3 specimens’ 0,1 Pebbles- - Stn CP 562,

178o?7 iV7fois' imn775‘on7nsm,'oio'05', ,992 : 53 sPecimens’ on echinoderms and shell. - Stn CP 564. 11°46.1'S,

1/8 ^ 1.4 W, 1015-10-0 m, 20.05.1992 : 15 specimens, on pebbles. — Stn CP 565, 11°47.4'S, 178°25.3'W, 900 m,

Source : MNHN, Paris

VERRUCOMORPHA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

117

20.05.1992 : 7 specimens, on shells. — Sin DW 586. 13°10.7'S, 176°13.1'W, 510-600 m. 22.05.1992 : 1 specimen, on

coral. — Stn DW 597, 12°31.4'S, 174°18.6'W, 469-475 m. 24.05.1992 : 1 specimen, on coral. — Sin DW 598,

12°30.5'S, 174°18.6'W, 702-708 m, 24.05.1992 : 2 specimens, on pebbles. — Stn DW 618, 14°21.7'S, 178°00.5'W.

420-435 m, 27.05.1992 : 1 specimen, on spalangoid. — Sin CP 623, 12°34.2'S, 178°15.1'W, 1280-1300 m,

28.05.1992 : 1 specimen, on scalpellid. — Stn DW 636, 13°39.4'S, 179°55.5'E, 650-700 m. 30.05.1992 : 1 specimen,

on coral.

Philippines. MUSORSTOM 3 : sin CP 106, 13°47.00'S, 120°30.30’E, 668 m. 02.06.1985 : 3 specimens, on

bivalve.

Fig. 13. — Metaverruca recta (Aurivillius, 1898) : a, complete shell (dorsal view), MNHN-Ci 2312; b, complete shell,

larger specimen, showing development of interlocking ribs between rostrum and carina (dorsal view), MNHN-Ci

2313; c, mandible; d, first maxilla; e, basal portion of sixth cirrus, with caudal appendage attached; f, articulated

movable scutum and tergum, showing repair to damaged scutum and resultant development of what appears to be a

fourth rib on the tergum (exterior), MNHN-Ci 2314. Material : a, c-e from BlOCAL, stn CP 23; b from BlOCAL,

stn CP 26; f from MUSORSTOM 7. stn DW 557.

118

J.S. BUCKERIDGE

Diagnosis. —Metaverruca with movable tergum and scutum having three articular ribs; exterior generally

smooth with widely spaced growth lines; caudal appendages short.

Description. — A large verrucid, shell white, low conic, sides steep; operculum sub-parallel to base,

D-shaped; exterior generally smooth, with fine, well spaced concentric growth lines; rostrum and carina

articulating with up to seven ribs.

Fixed scutum quadrangular, with a well formed myophore for adductor muscle attachment; basal margin of

shell thickened and inflected on larger specimens; fixed tergum with umbo extending fractionally beyond that of

movable tergum.

Movable tergum quadrangular, of low relief, with two prominent and one narrow articular ribs; movable

scutum triangular of low relief, with an even narrower third rib, internally opercular plates are of low relief, plates

together form a straight rostro-carinal hinge.

Mandible tridentate, lower angle pectinate, divided into two parts, first maxilla with one prominent and two

larger spines above the notch and four on the lower angle; penis short, a little less than the length of the caudal

appendages. Cirri possessed the following number of segments (r-c being rostro-carinal diameter) :

r-c (mm) 1 II HI IV V VI c a

5-2 H.IO 7,14 22.23 22,27 31,31 31,32 8

>4-9 11,12 9,21 32,34 36,40 43,41 40,43 8

01 the specimens examined, the length of the caudal appendages was consistently a little less than that of the

basal segment of the pedicel of cirrus VI.

Discussion. — Numerous workers have recognised the need to synonomise a number of Metaverruca species;

Gruvel (1920), Niusson-Cantell (1929), Buckeridge (1983), with Southward and Southward (1958),

noting that recta has priority.

M. recta is one of the largest verrucids, with specimens over 14.5 mm rostro-carinal diameter being measured

from this collection (Biocal. Stn CP 27). The three articular ribs between movable scutum and tergum, the

generally inornate exterior, the lixed scutum myophore, and the short caudal appendages distinguish this form from

all other verrucid species. (Even specimens of less than 2.0 mm rostro-carinal diameter possessed the three articular

nbs on the movable opercular plates).

The three articular ribs on the opercular plates is an important characteristic for recognising this species; one

specimen lrom Musorstom 7, Stn DW 557, however, possessed four ribs, and in this instance appeared very

similar to M. defayeae , a closer examination showed that the extra rib was a result of damage (mechanical?), with

the movable scutum being broken (fig. 13f). The animal was able to repair the damage, but in the process

developed the further rib.

Distribution. — Cosmopolitan, 160-2110 metres. Miocene (New Zealand, Buckeridge, 1983).

Genus ROSTRATOVERRUCA Broch, 1922

Verruca "Sectio Rostrato-verruca" Broch, 1922 : 298.

Verruca ( Rostratoverruca ) - Foster. 1978 : 68.

Roslratoverruca - Zevina, 1978 : 1813.

Diagnosis. - Verrucids with apex of rostrum removed from margin; fixed scutum without myophore-

operculum sub-parallel to base.

TYPE. — Verruca nexa Darwin, 1854 : 522. West Indies, 60 metres.

SPECIES. — With the placement of Verruca dens within Brochiverruca , the following 8 species are now

attributed to this genus : Rostratoverruca conchula (Hoek, 1913), Timor; Rostratoverruca intexta (Pilsbry. 1912),

Source : MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

119

Indian Ocean - Western Pacific; Rostratoverruca koehleri (Gruvel. 1907), Bay of Bengal; Rosiratoverruca kruegeri

(Broch, 1922), Western Pacific; Rostratoverruca murrayi (Stubbings, 1936), Zanzibar; Rostratoverruca nexa

(Darwin, 1854), West Indies; Rostratoverruca sewelli (Stubbings.1936), Zanzibar; Rostratoverruca malevichi

Zevina, 1988, South Pacific.

Discussion. — As many of the specimens studied in this collection were attached to cidaroid spines, the angle

between operculum and base is often difficult to ascertain. The most important criterion for recognising this genus

being the displaced rostral apex.

Stratigraphic range. — Recent.

Distribution. — West Indies and Indo-Wcst Pacific, 60-3250 metres.

Rostratoverruca intexta (Pilsbry, 1912)

Fig. 14 a-f

Verruca intexta Pilsbry, 1912 : 292; 1916 : 47. — Nilsson-Cantell, 1927 : 774; 1929 : 468, fig. 3. — STUBBINGS,

1940: 389.

Verruca conchula Hoek, 1913 : 146, figs 14-15.

Verruca ( Rostratoverruca ) intexta - Rosell, 1989 : 26, pi. 7 f-g; 1991 : 33.

MATERIAL EXAMINED. — Indonesia. Karubar : stn CP 9. 05°23'S, 132°29'E, 361-389 m, 23.10.1991 :

25 specimens, (MNHN-Ci 2318-2319), on glass sponge. — Sin CC 10, 05°2rS. 132°29’E, 329-389 m, 23.10.1991 :

4 specimens, on flat surface. — Stn CP 12, 05°25'S, 132°37'E, 412-434 m, 23.10.1991 : 10 specimens, on glass

sponge. — Stn CP 35, 06°07'S, 132°44'E, 390-502 m, 27.10.1991 : 2 specimens, on glass sponge. — Stn CC 40,

07°46'S, 132°31'E, 442-468 m, 28.10.1991 : 4 specimens, on glass sponge. — Stn CC 56, 08°16'S, 131°59'E, 552-

549 m, 31.10.1991 : 2 specimens, on glass sponge. — Stn CP 59, 08°20'S, 132°11'E, 405-399 m, 31.10.1991 :

1 specimen. — Stn CP 69, 08°42'S, 131°53’E, 356-367 m, 2.11.1991 : 7 specimens, on coral, Chirona, Scalpeltum. —

Stn CP 71, 08°38'S, 131°44'E, 477-480 m, 2.11.1991 : 1 specimen. — Stn CP 86, 09°25’S, 131°13'E, 226-222 m.

4.11.1991 : 3 specimens, on gastropod.

Philippines. MUSORSTOM 3 : stn CP 106, 13°47.0'S, 120°30.3'E, 640-668 m, 2.06.1985 : 1 specimen.

New Caledonia. Biocal : stn CP 67, 24°55’S, 168°22'E, 500-510 m. 3.09.1985 : 1 specimen, on coral.

Chalcal 2 : stn DW 74, 20°04.36'S, 168°38.38'E, 650 m, 3.09.1985 : 1 specimen, on hydrozoan.

Loyalty Islands. MUSORSTOM 6 : stn DW 449, 20°54.40'S, 167°17.75’E, 300 m, 20.02.1985 : 3 specimens, on

gorgonian.

Diagnosis. — Rostratoverruca with prominent, rounded ribbing; rostrum patelliform, with apex removed

some considerable distance from the scutal border; movable tergum and scutum each with three articular ribs;

operculum sub-parallel to base.

Description. — Shell cream-white, walls with prominent, rounded ribbing; rostrum patelliform, with apex

central in large specimens, apex distinctly beaked; carina with an elongated, straight, "prow-like" upper rib

articulating with the rostrum, apex produced, to extend well out beyond the basi-tergal angle of the tergum; base

membranous.

Fixed scutum broader than fixed tergum, articulating with up to three ribs in larger specimens; both plates with

incurved beaks, and with apices extending beyond those of the movable plates.

Movable scutum only a little narrower than the movable tergum. triangular, with three curved, diagonal ribs on

the tergal side, the upper two secondary ribs interlocking with the tergum. the primary (apico-basal) rib

interlocking with both tergum and rostrum at the basi-tergal angle; three further ribs occur on the occludent

portion, interlocking basally with the rostrum; apex beaked; interior with a rounded moderately well developed

adductor muscle depression.

Movable tergum quadrilateral, apex incurved towards scutum, primary apico-basal rib curved, interlocking at

the basi-scutal angle; two secondary ribs articulate with the movable scutum.

120

J. S. BUCKERIDGE

FlGn R°Slr.alOVeZUCa in,ex,a (Pilsbry' 1912) : a- complete shell (dorsal view), MNHN-Ci o318 b articulated

rZ (CX,Cr'0r): d' m0Vable '«*“*« (exSor ybeX^S;

’ g- bafdl porllon of sixth cirrus- w»h penis and caudal appendage attached; h, articulated movable

MNIfli-Cid2319 “aII material JLctT ^ reSUl,8nt dCVe'0pn,ent °f 3 TOn-ribbed «*>«- (--or).

SDincs^notch li^umT' pCClhvd{Q' divided int0 lwo P**; ^ maxilla with two prominent upper

spines, notch hirsute, lower angle hirsute, with six or more spines.

Drobd^fnliTi^M8 I' SUbTI: Ca,Ud,a‘ appendages Ion8cr than the pedicel of c.mis VI; penis very long,

“ 3 aPPenda8eS' ^ P0SSCSSed ^ f°U°'ri"* n“mber °f s^me"ls <™

r-c (mm)

5.4

I II

9,13 9.14

III IV

18,19 21,22

V VI

23,25 23,26

c.a.

20

Source MNHN. Paris

VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

121

Discussion. — This species is distinguished from Rostratoverruca nexa by the patelliform rostrum, the strong

ribbing, the number of ribs on the movable tergum and scutum, and the prominent prow-like upper rib on the

carina. It is similar to Rostratoverruca kruegeri (Broch, 1922) but possesses a much broader movable tergum and

scutum than that species, and unlike R. kruegeri , has no strong ribbing on the occludcnt side of the apico-basal rib

of the tergum.

The large amount of material available for observation provided an opportunity to gain an appreciation of

morphological variation. One specimen in particular (fig. 14h). appears to demonstrate the effect of pathogenesis

on both the movable scutum and rostrum, with characteristically strong ribbing present in early growth, but

abruptly absent subsequently.

Distribution. — Indo-Pacific, 194-1002 metres.

Rostratoverruca kruegeri (Broch, 1922)

Fig. 15 a-f

Verruca Kriigeri Broch. 1922 : 295, figs 43-44.

Verruca (Roslraioverruca) kriigeri - BROCH, 1932 : 46.

Material EXAMINED.— Indonesia. Karubar : stn CP 46, 08°01'S, I32°51'E, 271-273 m, 29.10.1991 :

28 specimens (MNHN-Ci 2320). on cidaroid spines. — Sin CP 85, 09°22’S, 131°14'E, 245-240 m, 4.11.1991 :

85 specimens, on cidaroid spines.

Philippines. Musorstom 3 : stn CP 139, 11°52.9'S, 120°14.7'E, 240-267 m, 6.06.1985 : 200+ specimens, on

cidaroid spines.

Diagnosis. — Rostratoverruca with prominent, rounded ribbing on rostrum and carina; rostrum patelliform,

with apex removed some considerable distance from the scutal border; fixed scutum and tergum with weak

longitudinal ribbing; movable tergum and scutum each with four articular ribs.

Description. — Shell white, rostrum and carina with prominent, rounded ribbing; rostrum patelliform, with

apex central in large specimens, apex slightly incurved; carina with a moderately elongated, "prow-like" upper rib

articulating with the rostrum, but with superior margin ridged, due to fine secondary ribbing; apex produced, to

extend out beyond the basi-tergal angle of the tergum; carina and rostrum generally interlocking with three ribs;

base membranous.

Fixed scutum broader than fixed tergum, articulating with up to three ribs in larger specimens; longitudinal

ribbing weak, especially on smaller specimens; both plates with apices slightly incurved, that of fixed tergum

extending beyond that of the movable tergum.

Movable scutum only a little narrower than the movable tergum, triangular, with four curved, diagonal ribs on

the tergal side, the upper three secondary ribs interlocking with the tergum, the primary (apico-basal) rib

interlocking with both tergum and rostrum at the basi-tergal angle; three further ribs occur on the occludent

portion, interlocking basally with the rostrum; apex incurved; interior with a rounded, moderately well developed

adductor muscle depression.

Movable tergum quadrilateral, apex incurved towards scutum, primary apico-basal rib curved, interlocking at

the basi-scutal angle; three secondary ribs articulate with the movable scutum; scutal margin wilh sulcus

immediately below upper rib; three further ribs occur on the occludent portion, interlocking with the tergum.

Mandible tridentate, lower angle pectinate; first maxilla with prominent upper spine, notch poorly

differentiated, with three small spines, lower angle hirsute, with four spines.

Cirrus I and cirrus II subcqual; caudal appendages about half the length of cirrus VI; penis very long,

probosciform, slightly longer than cirrus VI. Cirri possessed the following number of segments (r-c being rostro-

carinal diameter) ;

r-c (mm) 1 II III IV V VI c.a.

5.6 8,13 9.13 14,16 19,21 19,23 20,24 19

Source MNHN. Paris

122

J.S. BUCKERIDGE

0.2 mm

kr!‘egT (Br0C^' !,922) : C°mplete Shdl (d0rsal view)- MNHN-Cl 2320; b, movable tergum

aP'C P°rtl0n °f m°VaWe SCU*Um (in,eri°r): *• mandible; f- maxilla.

Discussion. — This species is distinguished from Rostratoverruca intexia (Pilsbry, 1912) by the narrower

rihofh P 3teS' ‘1C numb^r 0f nbs on the movable tergum and scutum, and the less prominent prow-like upper

,S Sim r( l° Rostratoverruca koehleri (Gruvel, 1907), but possesses a much more prominent

apex on the fixed tergum, and greater ribbing on the occludent portion of the movable tergum.

Distribution. — Western Pacific, 233-290 metres.

DISCUSSION

ihe mST 1hIONT IhC material StUdied W3S collected from dePlhs ranging between 223 and 3680 metres. On

Ind c™ s wi.f^ hme^Ci!0nin8 may 56 determined for the region. but it is likely that water temperatures

dismhTnn Of H 3 r8 °n S distribu,ion- Thc fo,lowing provides a broad appreciation of the

entSonmems efrUC 8ener3’ ™ that A"iverruca' in Particular, is characteristic of slope and abyssal

Depth range (metres) Mean (metres) No. of Stns

412-3680 i 407 t 55

240-2110 788 90

466-620 542 /

356-700 520 6

380-450 415

222-668 408 18

+only 2 specimens of Altiverruca were recovered from depths less than 510 metres.

Genus

Altiverruca

Metaverruca

Verruca

Brochiverruca

Canieraverruca

R ostraloverruca

Source MNHN. Paris

VERRUCOMORPIIA OF NEW CALEDONIA, INDONESIA AND WALLIS AND FUTUNA ISLANDS

123

Fig. 16. — Scanning Electron Microscope views. : a , Alliverruca navicula (Hoek, 1913), complete shell. MNHN-Ci

2331; (dorsal view); b. Alliverruca navicula (Hoek, 1913), detail of apico-basal rib on movable tergum (exterior);

c, Metaverruca pacifica sp. nov., complete shell (dorsi-frontal view), MNHN-Ci 2330; d, Metaverruca pacifica sp.

nov., complete shell (dorsal view), MNHN-Ci 2330. Material : a-b, from Karubar, stn CP 91; c-d, from

Musorstom 5, stn CP 367.

124

J.S. BUCKERIDGE

Metaverruca recta was found to be the most abundant species in the region, being recorded from 55% of all

stations collected; it was found associated with other vcrrucids at only 16 stations (24%). This collection also

confirms Metaverruca recta as having the widest geographic, bathymetric and stratigraphic distribution of any

verrucid.

Endemism. — With the exception of Metaverruca defayeae (Loyalty, Wallis and Futuna Islands) all the new

species here are presently restricted to the New Caledonian region. Whilst this is likely to be a function of

incomplete collecting in other areas, it is worth noting that Metaverruca , presently represented by eight species

world wide, has five of these species present in New Caledonian waters. The occurrence of the now cosmopolitan

Metaverruca recta (also the earliest metaverrucid) from the Miocene of New Zealand suggests that the southwest

Pacific is either a centre of reliction (Newman, 1991), or throughout the Neogenc was a centre for Metaverruca

speciation.

Substrate. — Verrucids tend to be rather opportunistic in the choice of substrate, the most common being

the abundant benthic debris, (generally small pebbles), found at many stations, although broken shell material has

also been utilised. Some species appear to be host specific, in particular Altiverruca navicula, which was found

attached to glass sponges in almost 80% of specimens, Brochiverruca polystriata and Brochiverruca dens, both

found attached to coral in all cases, and Rostratoverruca kruegeri, always found attached to cidaroid spines.

ACKNOWLEDGEMENTS

The author wishes to thank Alain CROSNIER. ORSTOM (Institut Fran?ais de Recherche Scientifique pour lc

Devcloppement en Cooperation), for the invitation to work on the Musorstom collections, for providing access

to the material, and lor providing a stimulating working environment. Joseph Poupin, SMCB (Service Mixtc de

Controle Biologique des Armees) and Danielle Defaye (Museum national d’Histoire naturelle, Paris), provided

valuable assistance during the research project. This project was supported by a grant from the Mus6um national

d'Histoire naturelle.

REFERENCES

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Bull. Soc. zool. France. 23 : 189-199.

BrOCH, H.. 1922 —Papers from Dr. Th. Mortensens Pacific Expedition 1914-16. No. 10: Studies on Pacific Cirripeds

Videns. Medd. dansk. naturh. Foren.. 73 : 215-358.

BrOCH, H 1932 — Papers from Dr. Th. Mortensens Pacific Expedition 1914-16. No. 56: Indomalayan Cirripedia.

Vidensk. Medd. dansk. naturh. Foren., 91 : 1-146.

barnes, M., 1992. — The reproductive periods and condition of the penis in several species of common cirripedes

Oceanography and Marine Biology Annual Review, 30 : 483-525.

BUCKERIDGE, J.S 1983. — Fossil barnacles (Cirripedia: Thoracica) of New Zealand and Australia. Bull, paleonl NZ

geol. Surv. , 50 : 1-151. r

Darwim, C.H. 1854. — A monograph on the sub-class Cirripedia, with figures of all species. The Balanidae, the

Verrucidae. 684 pp. + pis 1-30. Ray Society, London.

Darwin, C.H., 1855. — A monograph on the fossil Balanidae and Verrucidae of Great Britain. 44 pp + pis 1-2

1 alaeontological Society Monograph, London (1854).

F°STEr, B.A^1978. - The marine fauna of New Zealand: Barnacles (Cirripedia: Thoracica). Mem. N.Z. oceanogr. Inst.,

GRRViiI''A^'’ 19°°‘ T, SUf lgS e^p6ces nouvelles appartenant au genre Verruca provenant de la campagne du Talisman.

Bull. Mus. natn. Hist, nat., Paris, 6 : 242-244.

GR7SALoA'’ 190L — D'agnoses de quelques especes nouvelles de Cirrhipedes. Bull. Mus. natn. Hist, nat., Paris, 6 :

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VERRUCOMORPHA OF NEW CALEDONIA. INDONESIA AND WALLIS AND FUTUNA ISLANDS

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GRUVEL, A., 1902. — Cirrhipedes. In : Expeditions Scientifiques du Travailleur etdu Talisman pendant les annties 1880-

1883. Masson et Cic, Paris. 178 pp.

GRUVEL, A., 1905. — Monographic des Cirrhipedes ou Thecostracds. Masson et Cie, Paris. 472 pp.

GRUVEL. A., 1907. — Cirrhipedes opercules de l’lndian Museum de Calcutta. Mem. Asial. Soc. Bengal, 2 : 1-10.

GRUVEL, A.. 1920. — Cirrhipedes provenant des campagnes scientifiques de SAS le Prince de Monaco. Result. Camp,

scient. Prince Albert I, 53 : 1-88.

HOEK, P.P.C., 1883. — Report on the Cirripedia collected by H.M.S. Challenger. Rep. scient. Results Voy. Challenger,

Zool., 8 (25) : 1-169.

HOEK. P.P.C., 1907. — Cirripedia. In : Resultats voyage S.Y. Belgica. 1897-1899. Rapp. Sci. Zool. Anvers : 3-9.

HOEK, P.P.C., 1913. — The Cirripedia of the Siboga-Expedition. Cirripedia, Sessilia. Siboga Exped., monogr. 31b :

129-275.

Newman, W.A., 1989. — Juvenile ontogeny and metamorphosis in the most primitive living sessile barnacle.

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Newman, W.A. & HESSLER, R.R., 1989. — A new abyssal hydrothermal verrucomorphan (Cirripedia: Thoracica): The

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790.

Nilsson-Cantell. C.A., 1929. — Cirripedien des genus Verruca der Deutsche Tifsee Expedition auf dem Dampfer

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PlLSBRY, H.A., 1916. — The Sessile barnacles (Cirripedia) collected in the collections of the U.S. National Museum:

Including a Monograph of the American species. Bull. U. S. natn. Mus., 93 : 1-366.

Rao, L.M.V. & Newman. W.A.. 1972. — Thoracic Cirripedia from Guyots of the Mid-Pacific Mountains. Trans. S. Diego

Soc. nat. Hist., 17 (6) : 69-94.

Rosell, N.C., 1981. — Crustacea : Cirripedia. In : Resultats des campagnes MUSORSTOM. I — Philippines (18-28 Mars

1976). Vol. 1 (12). Mem. ORSTOM, 91 : 277-307.

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Resultats des Campagnes MUSORSTOM. Volume 9. Mem. Mus. natn. Hist, nat., (A) 152 : 9-61.

Schumacher, H.C.F., 1817. — Essai d'un nouveau systeme des habitations des vers testaces. 287 pp + 22 pis.

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Southward, A.J. & Southward, E.C., 1958. — On the occurrence and behaviour of two little known barnacles

Hexelasma hirsutum and Verruca recta, from the continental slope. J. mar. biol. Ass. U.K., 37 : 633-647.

STUBBINGS, H.G., 1936. — Cirripedia. Rep. scient. John Murray Exped. 1933-34, 4 (1) : 1-70.

Stubbings, H.G., 1940. — Cirripedia, additional part. Rep. scient. John Murray Exped. 1933-34, 1 (3) : 383-399.

Zevina, G.B., 1978. — Deep-sea Verrucontorpha (Cirripedia, Thoracica) of the Pacific. Zool. Zh., 66 : 1812-1821 (in

Russian).

Zevina, G.B., 1987. — Abyssal Cirripedia Verrucomorpha (Thoracica) of the Atlantic and Indian Ocean. Zool. Zh., 66 :

1304-1313 (in Russian).

Zevina, G.B., 1988. — Deep-sea Verrucomorpha (Cirripedia, Thoracica) of the Pacific: 2. The South Pacific. Zool. Zh.,

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Russian).

Source : MNHN. Paris

TS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULTATS DES CA

Crustacea Amphipoda : Lysianassoids

from the tropical western South Pacific Ocean

James K. LOWRY & Helen E. STODDART

Australian Museum

P.O. Box A285, Sydney South

NSW 2000, Australia

ABSTRACT

There are currently 20 lysianassoid amphipod species known front the tropical western South Pacific Ocean. We

report on 32 species from the area, including one new genus ( Coriolisa ) and 19 new species ( Arislias Ihio, A. uokonia,

Bathyamaryllis ouvea, Clepidecrella tropicalis, Coriolisa novacaledonia, Cyphocaris bellona, Hippomedon vao,

Kerguelenia koutoumo, K. lifou, Lepidepecreella sarcelle, Onesimoides abyssalis, Socarnes rurutu, S. tiendi, S. luscarora,

Socarnopsis honiara, S. tandai, Trischizostoma richeri, Tryphosella ama and T. oupi). This brings the total species

known from the area to 46.

RESUME

Crustacea Amphipoda : Lysianassoides du Sud-Ouest Pacifique tropical.

II est actuellement admis que 20 espfeces d'amphipodes lysianassoides sont connues du Sud-Ouest Pacifique tropical.

Nous traitons ici de 32 espfeces trouv6es dans cette region, comprenant un genre nouveau ( Coriolisa ) et 19 esp£ces

nouvelles ( Arislias ihio, A. uokonia, Bathyamaryllis ouvea, Clepidecrella tropicalis, Coriolisa novacaledonia,

Cyphocaris bellona, Hippomedon vao, Kerguelenia koutoumo, K. lifou, Lepidepecreella sarcelle, Onesimoides abyssalis,

Socarnes rurutu, S. tiendi, S. tuscarora, Socarnopsis honiara, S. tandai, Trischizostoma richeri, Tryphosella ama et

l ■ oupi). Ainsi le nombre total d'especes connues de la region est porte & 46.

INTRODUCTION

Lysianassoid amphipods have never been considered as an important part of tropical ecosystems. The large

work of Schellenberg (1938) on the amphipods of the tropical Pacific Ocean reported only one lysianassoid

Lowry, J. K. & Stoddart, H. E., 1994. — Crustacea Amphipoda : Lysianassoids from the tropical western South

Pacific Ocean. In : A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM, Volume 12. Mem. Mus. rial. Hist, nat.,

161 : 127-223. Paris ISBN 2-85653-212-8.

128

J. K. LOWRY & H. E. STODDART

species,. the comprehensive study of Barnard (1970) on about 120 species of Hawaiian amphipods reported only

one lysianassoid species and the monograph of Myers (1985) on the amphipods of Fiji reported only two

lysianassoid species from a total of 85 amphipod species. PiRLOrs (1933. 1936) large accounts on amphipods

noQi n.d°neSian v?tersurePor,cd 16 lysianassoid species. In the world monograph of Barnard & Karaman

2 of '^a^amS P°dS C°nSidered 10 be bipolar cold wa,er submergents and not mentioned in the

Stod^rt 21“ (19f6H oT;r'Cd 44 SPCdeS °f lysianassoids from the wcs,em Indian Ocean and Lowry &

NTODDART (1993) recorded 22 lysianassoid species from Indonesian waters and 12 from the Philippines

significantly increasing the known diversity of both areas.

Or e^n'rfnr 'y8'3"3580*1? amPhiP°ds are currently known from the tropical western South Pacific

Ocean (for this study the area from the equator south to the Tropic of Capricorn and from the east coast of

Australia to approximately 180°E). Haswell (1879) described Ichnopus tenuicornis (as Glycerina ) the first

ysianassoid amphipod known from the Great Barrier Reef. Australia. No other lysianassoids were reported from

redesTribeT/ , °WRY &|IODDAV,0(io90' '"2) described Wandin Sriffini and Ichnopus capricornus and

redesc nbed /. tenmeorms ^TTBBING (1888) described Cyclocans tahitensis from Tahitian waters and Onesimoides

carmatus from the Coral Sea. Schellenberg (1938) described one lysianassoid species, Aristias tropicalis from

he Bismarck Sea. Birstein & Vinogradov (I960), in their massive study of the pelagic gammaridean

C lySia”aSSOid ""*• from “H- ocean S“e„ of

these species (Bal^allisoma schellenberg, Birstein & Vinogradov, 1958. Cyphocaris anonyx Boeck 1871

ScheneSri92?b and P 1916' C rtchardi Chcvreux- 1905a- Ic^opus pelagicus

Schellenberg. 1926b and Paracyphocans brevicornis Birstein & Vinogradov, 1955) occurred in pelagic waters of

Sew ‘““a ? ‘74°W an<i SCTe" *“*■ «*■»> *- oufsiS «f= area Z,

ew Caledonia Intes (1978) reported Eurythenes gryllus (Lichtenstein. 1822), Ledoyer (1984) described

,ZZ TCUU r LOWRY & ST0DDART <1992) described Ichnopus annasona and /. Llpatun Repelin

978)- in bis ecological study of the pelagic amphipods of the west and central Pacific, reported five species (also

L?d°20% alon!.R17o‘E) anT^RADh°p i'^ *7° Vari°US ,0CaIitieS nor,h-easl of New Caledonia (between 5°N

J ,, .S ’ ' & 1?( ,E) and French Po'ynesia (along the equator between 135°W and 155°W and between Tahiti

descnbed ,hree species : “ - -

N 'V!!i‘SHPaPer rKep0V 0n 32 ‘ysianassoid species from Musorstom collections made in the waters around

1:^777 is'7t °cwai,,s and Fuiuna- and fr°m colons made

tropical ^ 10,31 nUmbCr °f 'ySianaSSOid SFedes now k"™" to® the

fai,;‘iSnf0bvi0,US thal,,he $Pecies list reflec« collecting effort and is no. a true representation of the lysianassoid

ng r^^^r Pa,dflC 3rea- ^ inS,3nCe fiVC °f thC Cighl *** from Austral ale

taken amon^ae There PS' ™ PC‘agiC PrCdat°rS lakcn in planklon ,ows and one secies was

taken among algae. There are no representatives of soft bottom dwellers (such as the pachvnids) no snccies

ofTe^^^welltaRBers6^ "° eCl0paras"cs <such 35 Trischizostoma) and no species

tne wood dwelling genus Onesimoides , so prevalent in other areas. The collection from New Caledonia is mnrP

™ar:f lySianaSSOid life-styles- A,Ihou8h scavenging lysianassoid genelasuch as < OrchomlZa

S,ePhonyx- TryphoseUa and Waldeckia are represented, there has been no trapping in New Caledonian

L b8Vefa m°rC ad^ate eSt,ma'e °f 11,6 diversi,y- InverIebrate associates are

r XPK1 T ”7 Wan<1'"id taXOn iS “ «>wlagoo„al wafers Z some

Source : MNHN. Pari ;

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

129

as norlhern Australia or south-east Asia. However, it is becoming clear that lysianassoid amphipods are a diverse

part of tropical ecosystems.

METHODS

Reports of the Musorstom cruises which made these collections (including maps and station data) can be

found in RICHER DE Forges (1990, 1993). In 1991 one of us (JKL) visited the Service Mixte Controle Biologiquc

dcs Armdes (SMCB) in Tahiti and participated in a cruise on the R.V. Marara to the Austral Isles. Baited traps

were set down to 900 m off each island in the group and in some cases in shallow water inside the lagoons.

Plankton samples were collected off several islands using 1 m nets towed for 6 hours between 50 and 100 m depth.

Shallow water samples of algae and sediment were taken by free diving at each island. A full report of this cruise

is given in Poupin (1991).

Taxonomic descriptions have been generated from the taxonomic data base program DELTA (Dallwitz &

Paine, 1986). All new species names except Clepidecrella tropicalis. Onesimoides abyssalis and Trischizostoma

richeri should be treated as nouns in apposition. Coded setal types on the mandibular palp follow the scheme

presented by Lowry & Stoddart (1993). The codes used in the description of the spine-teeth on the outer plate

of maxilla 1 are explained in Lowry & Stoddart (1992, 1993). Individual spine-teeth are labelled on the

figures. Material is lodged in the Musdum national d'Histoire naturelle, Paris (MNHN), the Australian Museum,

Sydney (AM) and the Pusat Penelitian dan Pcngembangan Oseanologi, Djakarta (PPPO).

The following abbreviations are used on the plates : A, antenna; E. epistome and upper lip; EP, epimeron; G,

gnathopod; H, head; MD. mandible; MDP, mandibular palp; MP, maxilliped; MPIP, maxilliped inner plate;

MPOP , maxilliped outer plate; MPP. maxilliped palp; MX, maxilla; MX1IP, maxilla 1 inner plate;

MXIOP, maxilla 1 outer plate; MX1P, maxilla 1 palp; P, peraeopod: ST, spine-tooth; T. telson; U, uropod;

UR, urosome; I, left; r, right; lat. lateral.

LIST OF RECORDED SPECIES

Marquesas Islands

Orchomenella abyssorum (Stebbing. 1888).

Tuamotu Archipelago

Euryihenes cf. gryllus (Lichtenstein, 1822).

Society Islands

Cyclocaris tahitensis Stebbing, 1888

Parambasia rub Myers, 1985 (reported by MYERS, 1989).

Austral Isles

Cyclocaris tahitensis Stebbing, 1888.

Eurythenes cf. gryllus (Lichtenstein, 1822).

Ichnopus annasona Lowry & Stoddart, 1992.

Ichnopus pelagicus Schellenberg, 1926b (reported by Lowry & Stoddart. 1992).

Orchomenella gerulicorbis (Shulenberger & Barnard, 1976).

Parambasia acuticaudata Lcdoyer, 1984.

Socarnes rurutu sp. nov.

Waldeckia sp. 1 .

130

J. K. LOWRY & H. E. STODDART

Cook Islands

Parambasia nui Myers, 1985 (reported by MYERS, 1990).

Tonga

P arawaldedda mua Myers, 1986.

Wallis and Futuna Islands

Eurythenes cf. gryllus (Lichtenstein, 1822).

Onesimoides carinatus Stebbing, 1888.

Socarnes tuscarora sp. nov.

Fiji

Parambasia nui Myers, 1985.

Parawaldeckia lowryi Myers, 1985.

New Caledonia and the Loyalty Islands

Arisiias thio sp. nov.

Arislias uokonia sp. nov.

Bathyamaryllis ouvea sp. nov.

Clepidecrella iropicalis sp. nov.

Coriolisa novacaledonia sp. nov.

Cyphocaris bellona sp. nov.

Eurythenes cf. gryllus (Lichtenstein, 1822) (reported by Intes 1978)

Figorella tasmanica Lowry, 1984.

Hippomedon vao sp. nov.

Ichnopus annasona Lowry & Stoddart, 1992.

Ichnopus malpatun Lowry & Stoddart’, 1992.

Kerguelenia koutoumo sp. nov.

Kerguelenia lifou sp. nov.

Lepidepecreella sarcelle sp. nov.

Onesimoides abyssalis sp. nov.

Orchomenella distinctus Birstein & Vinogradov, 1960

P arambasia acuticaudata Ledoyer, 1984.

Procyphocaris indurata K.H. Barnard. 1925.

Socarnes tiendi sp. nov.

Stephonyx sp.

Tri sc hizo stoma richeri sp. nov.

Tryphosella ama sp. nov.

Tryphosella oupi sp. nov.

Chesterfield Islands

Cyphocaris bellona sp. nov.

Waldeckia sp. 2.

Coral Sea Basin

Onesimoides carinatus Stebbing, 1888.

Great Barrier Reef, Australia

Ichnopus capricornus Lowry & Stoddart, 1992

Ichnopus tenuicornis (Haswell, 1879) (reported by Lowry & Stoddart, 1992)

Wandtn grtffint Lowry & Stoddart, 1990. '

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

131

Solomon Islands

Socarnopsis Honiara sp. nov.

Socarnopsis tandai sp. nov.

Bismarck Sea

Arislias tropicalis Schellenberg, 1938.

Central and Western South Pacific pelagics

Baihycallisoma schellenbergi Birstein & Vinogradov, 1958 (recorded by Birstein & Vinogradov, 1960).

Cyphocaris anonyx Boeck, 1871 (recorded by Birstein & Vinogradov, 1960, and Rf.pelin, 1978).

Cyphocaris challenged Stebbing, 1888 (recorded by Birstein & Vinogradov, 1960, and Repelin, 1978).

Cyphocaris faurei K.H. Barnard, 1916 (recorded by Birstein & Vinogradov, 1960, and Repelin, 1978).

Cyphocaris richardi Chevreux, 1905a (recorded by BIRSTEIN & Vinogradov, 1960, and REPELIN, 1978).

Ichnopus pelagicus Schellenberg, 1926b [recorded by Birstein & Vinogradov, 1960 (as Socarnes longicornis ),

and Repelin, 1978].

Paracyphocaris brevicornis Birstein & Vinogradov, 1955 (recorded by Birstein & Vinogradov, 1960).

SYSTEMAT1CS

Genus ARISTIAS Boeck, 1871

Aristias thio sp. nov.

Figs 1-3

MATERIAL EXAMINED. — New Caledonia. Calsub : stn PL 13, 21°26'S, 166°22.7'E, east of Thio, 1567-1807 m,

4 March 1989 : 1 2, 14.4 mm, with non-setose oostegites (MNHN-Am 4485).

Fig. 1. — Aristias thio sp. nov., holotype female, 14.4 mm (MNHN-Am 4485), off Thio, New Caledonia.

132

J. K. LOWRY & H. E. STODDART

Types. — The unique specimen is the holotype.

Type Locality. — East of Thio, New Caledonia. 21°26’S, 166°22.7'E. 1567 to 1807 m.

Diagnosis. — Eyes apparently absent. Antenna 1 : accessory flagellum 3-articulate. Mandible : incisors

asymmetrical, left with minutely serrate margin: left lacinia mobilis a short, smooth peg. Maxilla 1 : outer plate

with 7 spine-teeth. 5 in outer row and 2 in inner row; inner plate with 6 plumose setae along inner margin.

Gnathopod 1 : parachelate; coxa vestigial. Peraeopods 5 and 6 : coxae strongly lobate posteriorly. Peraeopods 3 to

7 : propodus without distal spurs. Epimeron 3 : posterovcnlral comer narrowly rounded. Uropod 3 : outer ramus

with short article 2. Telson moderately cleft (49%).

Description. — Based on holotype female. 14.4 mm; male not known. Head : exposed, deeper than long;

lateral cephalic lobe large, broadly rounded; rostrum absent; eyes apparently absent. Antenna I : short, 0.14 times

Fig. 2. — Aristias thio sp. nov., holotype female, 14.4 mm (MNHN-Am 4485), off Thio, New Caledonia. Scale

represent 0.1 mm.

Source MNHN. Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

133

body; peduncular article 1 short, length 1.1 limes breadth; peduncular article 2 long, 0.6 times article 1. without

anterodistal projection; peduncular article 3 long, 0.37 times article 1; accessory flagellum medium length.

0.39 times primary flagellum, 3-articulate, article 1 long, 2.3 times article 2; flagellum 6-articulatc, with weak

2-field callynophore in female without setae or spines. Antenna 2 : slightly longer than antenna 1; weakly

geniculate between peduncular articles 3-4. article 3 short, 0.43 times article 4; flagellum well developed,

6-articulatc.

Mouthpart bundle : subquadrate. Epislome and upper lip : fused, with central notch. Mandible : incisors asym¬

metrical, large with straight margins, left with minutely serrate margin; left lacinia mobilis present, a short

smooth peg; without accessory spines or intermediate setae; molar a reduced setose flap; mandibular palp attached

midway; article 1 short, length 1 times breadth; article 2 slender, length 2.6 times breadth, 1.2 times article 3,

with 2 submarginal posterodistal A2-setae, without D2-setae; article 3 slender, distally truncate, long, length

4.2 times breadth, without A3-setae, with 4 proximal D3-setae and 3 apical E3-setae. Maxilla I : inner plate

tapering distally, at least half of inner margin setose, with 5-6 plumose setae; outer plate broad with 7 spine-teeth

in two rows; outer row with ST1 to ST3 large, stout, multicuspidate, ST4 large, slender multicuspidate, ST5-ST6

absent, ST7 large, broad. 9-cuspidate; inner row with STA large, slender, 4-cuspidate, STB-STC absent, STD

short, broad, 4-cuspidate; palp large, 2-articulate, with 2 long terminal spines, without subterminal setae, flag

spine present, distomedial margin serrate. Maxilla 2 : inner plate broad, outer plate narrow, inner plate 1 times

length outer plate. Maxilliped : inner plate large, subrectangular, with 1 apical nodular spine, oblique setal row

reduced, with 6 plumose setae; outer plate small, subovate, with 2 apical simple setae, without apical spines,

medial spines present, small, submarginal setae absent; palp large, 4-articulate; article 2 very broad, length 1 times

breadth, 1.1 times article 3; article 3 short, broad, length 1.5 times breadth; dactylus well developed, with

1 subterminal seta, unguis present.

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : parachelate; coxa vestigial; basis long, slender, length

2.5 times breadth, anterior margin smooth, without setae; ischium short, length 1 times breadth; merus, posterior

margin with group of long simple setae and patch of short setae; carpus wedge-shaped, produced dorsally, short,

length 1.5 times breadth and 1 times propodus, with patch of very fine setae near posterior margin and long simple

setae along posterior margin; propodus large, subtriangular, length 1.8 times breadth, tapering distally, posterior

margin serrate, convex, with 5 spines, without denticulate patch near posterior margin, palm slightly acute, with

straight, minutely serrate margin, posterodistal comer with 1 medial spine; dactylus simple, with serrate posterior

margin. Gnathopod 2 : minutely subchelate; coxa large, subequal in size to coxa 3; ischium long, length

2.1 times breadth; carpus long, length 3.4 times breadth, posterior margin straight; propodus subrectangular.

long, length 2.6 times breadth, palm transverse, with straight, serrate margin, posterodistal corner with 1 medial

and 1 lateral spines; dactylus not reaching corner of palm, posterior margin smooth.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus without minutely denticulate surface,

posterodistal spur, setae or spines; dactylus short, slender. Peraeopod 4 : coxa deeper than wide, with weak

posteroventral lobe, anterior margin slightly rounded, posterior margin slightly sloping anteriorly; merus weakly

expanded anteriorly; propodus without minutely denticulate surface, posterodistal spur, setae or spines; dactylus

short, slender. Peraeopod 5 ; coxa bilobate, posterior lobe strongly produced vcntrally; basis weakly expanded, pos¬

terior margin straight; merus not expanded posteriorly; propodus without minutely denticulate surface, anterodistal

spur, setae or spines; dactylus short, slender. Peraeopod 6 : coxa small, strongly lobate posteriorly; basis weakly

expanded, with slightly rounded smooth margin, without anteroventral lobe; merus not expanded posteriorly;

propodus without minutely denticulate surface or anterodistal spur, posterior margin without spines; dactylus

short, slender. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly rounded, smooth, posteroventral

comer rounded, posteroventral margin straight; merus with anterior and posterior margins subparallcl; propodus

with minutely denticulate surface, without anterodistal spur or spines; dactylus short, slender.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 6, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 1 : anteroventral comer rounded. Epimeron 3 : posteroventral

comer narrowly rounded. Urosomites : dorsally smooth. Uropod 1 : peduncle with 2 dorsolateral and 1 apicolatcral

spines; rami subequal in length, outer ramus with 1 lateral and 3 medial spines; inner ramus with 2 lateral spines.

Uropod 2 : with 1 apicolatcral and 1 apicomedial spines, without spines along distal margin; rami subequal in

length, outer ramus without spines; inner ramus with 1 lateral spine, without constriction or proximal flange.

134

J. K. LOWRY & H. E. STODDART

Fig. 3. — Aristias thio sp. nov., holotype female, 14.4 mm (MNHN-Am 4485), off Thio, New Caledonia. Scales

C/ropod 3 : peduncle short, length 1.4 times breadth, without dorsolateral flange, without dorsal spines,

midlateral spines or setae or distoventral spines; rami lanceolate, inner ramus reduced, about 0.9 times

outer ramus, outer ramus 2-articulate, article 2 short, rami without spines. Telson : shorter than broad, length

0.85 times breadth, moderately cleft (49%), without dorsal spines or dorsal setae, distal margins rounded, without

marginal pemcillate setae, with 2 simple marginal setae on each lobe, without marginal spines.

Source MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

135

Etymology. — Named for the town of Thio, near the type locality.

REMARKS. — Aristias thio has no eyes, only 3 articles on the accessory flagellum of antenna 1, no distal

spurs on the peraeopods and the inner ramus of uropod 3 is nearly as long as the outer. Aristias coriolis Lowry &

Stoddart, 1993, also has these character states, but it differs from A. thio in having more spine-teeth on the outer

plate of maxilla 1. apically acute posteroventral lobes on the coxae of peraeopods 5 and 6 and a more deeply cleft

telson. One other species, A. expers J.L. Barnard, 1967, shares all of these character states, except the inner ramus

of uropod 3, which just reaches the end of article 1. It differs further from A. thio in having a differently shaped

posterior margin on the propodus of gnathopod 1, a differently shaped basis on pcraeopod 7 and a slightly less cleft

telson.

Distribution. — Aristias thio is known only from south-eastern New Caledonia in 1800 m depth.

Aristias uokonia sp. nov.

Figs 4-6

MATERIAL EXAMINED. — New Caledonia. BlOCAL : stn DW 33, 23°09.71’S, 167°10.27'E to 23°10.80'S,

167°10.45'E, south of the Isle of Pines, 675-680 m, 29 August 1985 : 1 9, 16.2 mm, with setose oostegites (MNHN-Am

4430).

TYPES. — The unique specimen is the holotype.

TYPE LOCALITY. — South of the Isle of Pines, New Caledonia, 23°09.71'S, 167°10.27’E to 23°10.80’ S.

167°10.45'E, 675 to 680 m.

Fig. 4. — Aristias uokonia sp. nov., holotype female, 16.2 mm (MNHN-Am 4430), south of the Isle of Pines,

New Caledonia.

Diagnosis. — Eyes long, narrow, subsigmoid. Antenna 1 : accessory flagellum 6-articulate. Mandible :

incisors symmetrical, with straight, smooth margins; left lacinia mobilis a small spine. Maxilla 1 : outer plate

136

J. K. LOWRY & II. E. STODDART

wilh 13 spine-teelh, 1 1 in outer row and 2 in inner row; inner plate with 10 plumose setae along inner margin.

Gnathopod 1 : parachelate; coxa vestigial. Peraeopods 5 and 6 : coxa 5, posterior lobe weakly produced ventrally;

coxa 6 strongly lobate posteriorly. Peraeopods 3 to 7 : propodus with distal spurs. Epimeron 3 : posteroventral

comer narrowly rounded. Uropod 3 : inner ramus shorter than article 1 of outer ramus; outer ramus with short

article 2. Telson deeply cleft (72%).

Description. — Based on holotype female. 16.2 mm; male not known. Head : exposed, deeper than long;

lateral cephalic lobe large, narrowly rounded; rostrum absent; eyes long, narrow, subsigmoid. brown in alcohol.

FIV- — Arislias uokonia sp. nov., hololype female, 16.2 mm (MNHN-Am 4430), south of the Isle of Pines,

New Caledonia. Scales for A1 and A2 represent 0.5 mm, remainder represent 0.2 mm.

Source . MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

137

Antenna 1 : medium length, 0.22 times body; peduncular article 1 short, length 1.2 times breadth; peduncular

article 2 short. 0.25 times article 1, without anterodistal projection; peduncular article 3 long, 0.25 times article 1;

accessory flagellum, medium length, 0.46 times primary flagellum, 6-articulate, article 1 long, 3.4 times article 2;

flagellum 1 1 -articulate, with strong 2-field callynophorc with 11 large midmedial spines. Antenna 2 : slightly

longer than antenna 1; weakly geniculate between peduncular articles 3-4, article 3 short, 0.35 times article 4;

flagellum well developed, 13-articulate.

Mouthpart bundle: subquadrate. Epistome and upper lip : fused, with central notch. Mandible : incisors

symmetrical, large, with straight margins; left lacinia mobilis present, a small spine; without accessory spines or

intermediate setae; molar a small, smooth flap with setose margins; mandibular palp attached midway; article 1

short, length 1 times breadth; article 2 slender, length 3.3 times breadth, 1.2 times article 3, with 13 submarginal

posterodistal A2-setae, without D2-setae; article 3 falcate, long, length 3.5 times breadth, without A3-setae, with

19 D3-setae along most of posterior margin and 3 apical E3-setae. Maxilla 1 : inner plate tapering distally, inner

margin fully setose, 10 plumose setae; outer plate extremely broad with 13 spine-teeth in two rows; outer row

with ST1 to ST3 large, slender, multicuspidate, 6 spine-teeth from ST4 to ST7 all large, slender, multicuspidate;

inner row with STA large, slender, multicuspidate, STB-STC absent, STD long, slender, multicuspidate; palp

large, 2-articulate, with 2 short terminal spines, without subterminal setae, flag spine present on distolateral

corner, distomedial and distolateral margins serrate. Maxilla 2 : inner plate broad, outer plate narrow, inner plate

1 times length outer plate. Maxilliped : inner plate small, subovate, with 2 apical nodular spines, oblique setal

row strong with 6 plumose setae; outer plate medium size, subrectangular, with 6 apical simple setae, without

apical spines, medial spines present, small, submarginal setae absent; palp large, 4-articulate; article 2 very broad,

length 1.4 times breadth, 1.6 times article 3; article 3 short, broad, length 1.6 times breadth; dactylus well

developed, with 2 subterminal setae, unguis absent.

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : parachelate; coxa vestigial; basis long, slender, length

3.8 times breadth, anterior margin smooth, with simple setae; ischium short, length 1.1 times breadth; merus,

posterior margin with group of long simple setae and patch of short setae; carpus subrectangular, short, length

1.6 times breadth, longer than (1.2 times) propodus, with patch of very fine setae near posterior margin and long

simple setae along posterior margin; propodus large, subtriangular, length 1.6 times breadth, tapering distally,

posterior margin serrate, subtly sinusoidal, with 4 spines, without denticulate patch near posterior margin, palm

transverse, margin convex, serrate, posterodistal comer with 1 medial spine; dactylus simple, without subterminal

teeth or spines. Gnathopod 2 : minutely subchelate; coxa large, subequal in size to coxa 3; ischium long, length

3.1 times breadth; carpus long, length 3.7 times breadth, posterior margin straight; propodus subrectangular, long,

length 2.6 times breadth, palm transverse, with concave, minutely serrate margin, posterodistal corner without

spines; dactylus over-reaching corner of palm, posterior margin smooth.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus without minutely denticulate surface,

with small posterodistal spur, with 3 setae and 1 distal spine along posterior margin; dactylus short, stocky.

Peraeopod 4 : coxa deeper than wide, with weak postcrovcntral lobe, anterior margin slightly rounded, posterior

margin slightly sloping anteriorly; merus weakly expanded anteriorly; propodus without minutely denticulate

surface, with small posterodistal spur, with 2 setae and 1 distal spine along posterior margin; dactylus short,

stocky. Peraeopod 5 : coxa bilobate, posterior lobe slightly produced ventrally; basis expanded with posterior

margin smooth; merus slightly expanded posteriorly; propodus without minutely denticulate surface, with small

anterodistal spur, with 1 distal spine along anterior margin; dactylus short, stocky. Peraeopod 6 : coxa large,

strongly lobate posteriorly; basis expanded posteriorly with smooth posterior margin, without anteroventral lobe;

merus slightly expanded posteriorly; propodus without minutely denticulate surface, with small anterodistal spur,

with 1 distal spine along anterior margin; dactylus short, stocky. Peraeopod 7 : basis expanded posteriorly,

posterior margin slightly rounded, minutely crcnale, postero ventral corner rounded, posteroventral margin rounded;

merus slightly expanded proximally with slightly rounded posterior margin with 5 spines; propodus without

minutely denticulate surface, with small anterodistal spur, with 3 spines and 1 distal spine along anterior margin

and 3 setae along posterior margin; dactylus short, stocky.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 6, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 1 : anteroventral corner narrowly rounded. Epimeron 3 :

posteroventral corner narrowly rounded. Urosomites : dorsally smooth. Uropod 1 : peduncle with 7 dorsolateral.

138

J. K. LOWRY & H. E. STODDART

Fig. 6. — Arislias uokonia sp. nov., holotype female, 16.2 mm (MNHN-Am 4430), south of the Isle of Pines,

New Caledonia. Scales represent 0.5 mm.

5 dorsomedial and 1 apicomedial spines; rami subequal in length, outer ramus with 3 lateral spines; inner ramus

with 4 medial and 5 lateral spines. Uropod 2 : with 5 dorsolateral, 1 apicolateral, 2 dorsomedial and 1 apicomedial

spines, without spines along distal margin; rami subequal in length, outer ramus with 3 lateral spines in weak

Source . MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

139

acclivities; inner ramus with 3 medial and 2 lateral spines, inner ramus without constriction or proximal flange.

Uropod 3 : peduncle short, length 1.5 times breadth, without dorsolateral flange, with 1 apicomedial spine,

without midlateral spines or setae, with 1 distoventral spine; rami lanceolate, inner ramus reduced, about

0.77 times outer ramus, outer ramus 2-articulate, article 2 short, rami without spines. Telson : longer than broad,

length 1.1 times breadth, deeply cleft (72%), without dorsal spines or dorsal setae, distal margins truncated,

without marginal penicillate setae, with 1 simple marginal seta and 1 marginal spine on each lobe.

Etymology. — Named for Point Uokonia, Isle of Pines, near the type locality.

REMARKS. — This is the first record of midmedial spines on the face of the callynophore in lysianassoid

amphipods. As far as we know these spines distinguish A. uokonia from all other aristiids. Arisiias uokonia has

the inner ramus of uropod 3 reduced. Other species with this character state include the Pacific and Indian Ocean

species A. adrogans J.L. Barnard, 1964a, A. expers J.L. Barnard, 1967, A. stenopodus Ledoyer, 1986, and the

North Atlantic species A. tumidus (Krpyer, 1846). Arisiias adrogans and A. stenopodus differ from does

A. uokonia in having imperfectly developed eyes or none at all. Neither A. adrogans nor A. expers has the telson

as deeply cleft as A. uokonia , both have differently shaped posterior margins on the propodus of gnathopod 1 and

A. expers has no spurs on the peraeopods. Arisiias stenopodus has a deeply cleft telson. but it is much wider than

long and the bases of peraeopods 5 to 7 are not as well developed. Arisiias tumidus differs from A. uokonia in

having a differently shaped eye, fewer spine-teeth on the outer plate of maxilla 1 and a better developed

posteroventral lobe on coxa 5.

Distribution. — Aristias uokonia is known from south of the Isle of Pines. New Caledonia, in 680 m depth.

Genus BATHY AMARYLLIS Pirlot, 1933

Bathyamaryllis ouvea sp. nov.

Figs 7-10

MATERIAL EXAMINED. — New Caledonia. Biocal : stn DW 44, 22°47.30’S, 167°14.30'E to 22°47.35’S,

167°14.50’E, south of the Isle of Pines, 440-450 m, 30 August 1985 : I 2. 7.8 mm, ovigerous. 7 eggs (MNHN-Am

4775); 1 6,1.1 mm (MNHN-Am 4776); 28 specimens (MNHN-Am 4382); 25 specimens (AM P42124). — Stn DW 46,

22°53.05'S, 1 67° 17.08'E to 22°53.27'S, 167°17.41'E, south of the Isle of Pines, 570-610 m, 30 August 1985 :

55 specimens (MNHN-Am 4441). — Stn DW 51. 23°05.27'S, 167°44.95'E. south of the Isle of Pines, 680-700 m,

31 August 1985 : 6 specimens (MNHN-Am 4442). — Stn DW 77, 22°15.32'S, 167°15.40'E. north-west of the Isle of

Pines, 440 m, 5 September 1985 : 1 6. 8.0 mm, 1 6 , immature (MNHN-Am 4793). — Stn DW 104, 21°30.62'S,

166°21.26'E to 21°30.95'S, 166°21.55'E, north-east of Thio, 375-450 m, 8 September 1985 : 1 specimen (MNHN-Am

4463).

Types. — The female specimen (from MNHN-Am 4775) is the holotype. the other specimens are paratypes.

Type Locality. — South of the Isle of Pines. New Caledonia, 22°47.30'S, 167°14.30'E to 22°47.35'S,

167°14.50'E, 440 to 450 m.

Diagnosis. — Antenna 1 : peduncular article 2. 0.5 times as long as article 1. Antenna 2 : peduncular article 4

and 5, each 3 times as long as wide. Uropod 2 : outer ramus about 0.7 times as long as inner ramus; inner ramus

with 5 medial spines and 6,1 lateral spines.

Description. — Based on holotype female 7.8 mm (MNHN-Am 4775); paratype male, 8.0 mm (MNHN-Am

4793). Head ; exposed, much deeper than long, extending below insertion of antenna 2 with indentation at level ol

insertion; lateral cephalic lobe moderate, rounded; rostrum moderate; eyes long, oval (very faint in alcohol).

Antenna 1 : elongate, 0.38 times body; peduncular article 1 long, length 2.2 times breadth, ball-shaped

proximally, without dorsal crest, with large midmedial tooth, without posterodistal tooth, without anterodistal

projection; peduncular article 2 long, 0.5 times article 1. without anterodistal projection; peduncular article 3 long.

140

J. K. LOWRY & H. E. STODDART

0.25 times article 1; accessory flagellum short. 0.25 times primary flagellum, 5-articulate (male 7), article 1 short,

1.5 times article 2 (male long, 2.5 times article 2); flagellum 22-articulate (male 21 -articulate), without

callynophore in female (strong 2-field in male); calceoli absent in female and male. Antenna 2 : slightly longer

than antenna 1; peduncle without brush setae in female (weak in male), female weakly geniculate between

peduncular articles 3-4, article 3 short, 0.29 times article 4; flagellum well developed, 23-articuIate (male at least

13, damaged); calceoli absent in female and male.

Fig. 7. — Bathyamaryllis ouvea sp. nov., paratype male, 7.7 mm (MNHN-Am 4776), south of the Isle of Pines,

New Caledonia.

Mouthpart bundle : subquadrate. Epistome and upper lip : fused, with central bulge. Mandible : incisors

symmetrical, small, with slightly convex margins; left lacinia mobilis present, a stemmed distolatcrally cusped

blade; accessory spine row with large distal setal tuft, left and right rows each with 10 long, slender, serrate spines,

with "whip-like" intermediate setae; molar a small, smooth flap with setose margins; mandibular palp attached

midway; article 1 short, length 1.2 times breadth; article 2 slender, length 4.7 times breadth, 1.6 times article 3,

without A2-setae in female (male 14 - 16), without D2-setae; article 3 tapering distally, short, length 2.8 times

breadth, without proximal A3-setae in female (male 1), with 4 distal D3-setae (male 11-12) and 2 apical E3-setae.

Maxilla I : inner plate broad with 2 plumose apical setae; outer plate broad with 1 1 spine-teeth in

6/5 arrangement; outer row with ST1 to ST3 large, stout, weakly cuspidate, ST4 large, stout. 4-cuspidate, ST5

large, stout, 5-cuspidate. ST6 large, stout, 9-cuspidatc, ST7 contiguous with ST6, large, slender, 17-cuspidate

medially: inner row with STA large, slightly displaced from STB, 3-cuspidate, STB large, broad, 4-cuspidate, STC

large, broad, 5-cuspidate, STD broad, smaller than STC. 4-cuspidate; palp absent. Maxilla 2 ; inner and outer

plates narrow, inner plate bulging proximomedially. inner plate 1 times length outer plate. Maxilliped : inner

plate large, subrectangular, with 3 apical vestigial nodular spines, oblique setal row reduced with 7 plumose setae;

outer plate medium size, subovatc. without subapical notch, without apical setae or spines, without medial spines,

submarginal setae vestigial; palp large, 4-articulatc; article 2 slender, length 2.5 times breadth, 1.4 times article 3;

article 3 long, slender, length 2.4 times breadth; dactylus reduced, with 1 terminal and 1 subterminal seta, unguis

absent.

P eraeoriites : 1 to 7 dorsally smooth; peraeonitc 6 without sternal hook. Gnathopod 1 : simple; coxa vestigial;

basis very long, slender, length 6.4 times breadth, anterior margin smooth, with simple setae; ischium long,

length 1.7 times breadth; merus, posterior margin lined with long simple setae; carpus subrectangular, long.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

141

length 2.6 times breadth and 1 times propodus, without denticulate patch near posterodistal margin; propodus

large, subrectangular, length 2.6 times breadth, margins slightly converging distally, posterior margin serrate,

straight, with 4 spines and 4 groups of setae, without disto-medial setae, without denticulate patch near posterior

margin, palm absent; dactylus simple, with serrate posterior margin. Gnathopod 2 : subchelate; coxa reduced,

partially covered by coxa 3; ischium long, length 3.2 times breadth; carpus very long, length 6.2 times breadth,

posterior margin straight; propodus subrectangular, long, length 2.6 times breadth, palm acute, with straight,

serrate margin, posterodistal corner with 1 medial spine; dactylus reaching comer of palm, posterior margin serrate.

Fig. 8. — Bathyamaryllis ouvea sp. nov., holotype female, 7.8 mm (MNHN-Am 4775); paralype immature male, 7.7 mm

(MNHN-AM 4776); south of the Isle of Pines, New Caledonia. Scales represent 0.1 mm.

142

J. K. LOWRY & H. E. STODDART

FlGNew Caledonia”1 Scales^ forln Z ,n°V ’ female ’ 7‘8 mm (MNHN-Am 4775), south of the Isle of P.nes,

iNew uaieaoma. Scales tor Ul-3 represent 0.2 mm, remainder represent 0.5 mm.

Source : MNHN. Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

143

Peraeopod 3 : coxa large; merus not expanded anteriorly; merus-carpus without brush of simple setae in female

(dense brush present in male); propodus with 6 spines and 2 distal spines along posterior margin; daetylus short,

slender. Peraeopod 4 : coxa deeper than wide, with large posteroventral lobe, anterior and posterior margins

obtusely angled; merus not expanded anteriorly; merus-carpus without brush of simple setae in female (dense brush

present in male); propodus with 6 spines and 2 distal spines along posterior margin; daetylus short, slender.

Peraeopod 5 : coxa bilobate, posterior lobe produced ventrally; basis expanded with posterior margin smooth;

merus slightly expanded posteriorly; propodus with 10 spines and 2 distal spines along anterior margin; daetylus

short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis, anterior margin rounded proximally,

straight distally, basis expanded posteriorly with minutely crenate posterior margin, without anteroventral lobe;

merus slightly expanded and rounded posteroproximally, straight posterodistally with 5 setae; propodus with

10 spines and 2 distal spines along anterior margin; daetylus short, slender. Peraeopod 7 : basis expanded

posteriorly, posterior margin almost straight, minutely crenate, posteroventral comer subquadrate, posteroventral

margin straight; merus slightly expanded, convex posterior margin with 6 spines; propodus with 1 1 spines and

2 distal spines along anterior margin and 5 setae along posterior margin; daetylus short, slender.

Oostegiles : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 7, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron I : anteroventral corner rounded. Epimeron 3 : posteroventral

corner notched. Urosomites : dorsally smooth. Uropod 1 : peduncle with 12 dorsolateral, 1 apicolateral,

2 dorsomedial and 1 apicomedial spines; rami subequal in length, outer ramus with 9 lateral spines; inner ramus

with 5 medial and 6 lateral spines. Uropod 2 : peduncle with 2 dorsolateral, 1 apicolateral and 1 apicomedial

spines; outer ramus 0.68 times as long as inner ramus, outer ramus with 7 lateral spines, inner ramus with

5 medial and 6,1 lateral spines; inner ramus with weak constriction. Uropod 3 : peduncle well developed, short,

length 1.6 times breadth, without dorsolateral flange, with 5 dorsomedial and 1 apicomedial spines, without

midlateral spines or setae, without distovcntral spines; rami lanceolate, subequal in length, outer ramus

2-articulate, article 2 short, article 1 with 7 lateral and 4 medial spines; inner ramus with 4 medial and 4 lateral

spines, plumose setae absent in female (present in male). Telson : slightly longer than broad, length 1.1 times

breadth, slightly to moderately cleft (29%), distal margins rounded, with 1 marginal penicillate seta and 1-2 simple

setae on each lobe, without marginal spines.

Fig. 10. — Bathyamary llis ouvea sp. nov., paratype male, 8.0 mm (MNHN-Am 4793), north-west of the Isle of Pines,

New Caledonia. Scales represent 0.2 mm.

144

J. K. LOWRY & H. E. STODDART

Etymology. — Named for the island of Ouv6a in the Loyally Islands.

Remarks. — Immalure males in ihesc samples differ from adulls in several ways : although the proximal

articles of the antenna 1 flagellum have fused, aesthetascs have not developed; there arc no setal brushes on the

merus-carpus of peraeopods 3 and 4; and plumose setae have not developed on the rami of uropod 3.

Bathy amaryllis ouvea and B. perezii Pirlot. 1933, are very closely related. The most obvious difference between

them is that B. perezii has much more slender first and second antennae in which peduncular article 2 of antenna 1

is 0.9 times as long as article 1 and peduncular articles 4 and 5 of antenna 2 are each 6 times as long as wide and

the apparent lack of spines on the medial margin of the inner ramus of uropod 2. More subtle differences in

B. perezii include the maxillipedal palp which is more slender, the palm of gnathopod 2 which is transverse and

the posteroventral corner of the basis of peracopod 7 which is more rounded.

Distribution. — Bathy amaryllis ouvea is known from south of the Isle of Pines and the strait between

New Caledonia and the Loyalty Islands, in 375 to 610 m depth.

Genus CLEPIDECRELLA J.L. Barnard. 1962

Clepidecrella J.L. Barnard. 1962 : 24; 1969 : 338. — Barnard & Karaman, 1991 : 476.

Diagnosis. — Mandible : incisor and lacinia mobilis present, accessory spine row absent, molar vestigial.

Maxilla 1 : inner plate reduced, without apical setae; outer plate with reduced number of spine-teeth. Maxilliped :

inner and outer plates reduced in size and spination. Gnathopod 1 : linear, simple or weakly subchelate.

Peraeopod 4 : coxa with very large posteroventral lobe. Peraeopod 5 : basis slightly to moderately expanded.

Uropod 3 : inner ramus reduced or absent.

Type Species. — Clepidecrella cabinda J.L. Barnard, 1962, by original designation.

Species Composition. — Clepidecrella contains two species : C. cabinda J.L. Barnard, 1962. and

C. tropicalis sp. nov.

Remarks. — Clepidecrella has most of the characters of Kerguelenia, particularly the reduced spine-tooth

arrangement of maxilla 1; reduced inner and outer plates on the maxilliped; simple, linear gnathopod 1; very large

posteroventral lobe on coxa 4; linear to moderately expanded basis on peraeopod 5; and the reduced rami of

uropod 3. It differs in the mandible which has a vestigial incisor and a vestigial molar. Keeping them separate,

based on the mandible, indicates that gnathopod 1 and uropod 3 are homoplastic characters. Although this is not a

satisfactory arrangement we are leaving the genera separate until a more complete phylogenetic assessment is

available.

distribution. — Clepidecrella is known from the tropical western South Pacific Ocean and the South

Atlantic Ocean, in 450 to 5000 m depth.

Clepidecrella tropicalis sp. nov.

Figs 11-13

Material EXAMINED. — New Caledonia. Biocal : stn DW 44, 22°47.30'S. 167°14.30'E to 22°47.35'S,

™-H. south of 'he Isle of Pines, 440-450 m, 30 August 1985 : 1 9 , 5.0 mm, ovigerous. 2 eggs (MNHN-Am

Types. — The unique specimen is the holotype.

Source : MNHN, Pans

I.YSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

145

Type Locality. — South of the Isle of Pines, New Caledonia, 22°47.30'S. 167°14.30’E to 22°47.35'S,

167°14.50'E, 440 to 450 m.

Diagnosis. — Antenna 1 : flagellum 13-articulate. Antenna 2 : flagellum 9-articulate. Mandible : palp article

2 with 6 distal A2-setae. Gnathopod 1 : simple with linear carpus and propodus. Peraeopod 5 : basis moderately

expanded with small posteroventral lobe. Epimeron 3 : posteroventral corner narrowly rounded. Uropod 3 : rami

absent. Telson : entire, posterior margin emarginate.

Fig. 11. — Clepidecrella tropicalis sp. nov., hololype female, 5.0 mm (MNHN-Am 4385), south of the Isle of Pines,

New Caledonia.

Description. — Holotype female, 5.0 mm; male not known. Head : exposed, slightly longer than deep;

lateral cephalic lobe large, narrow, subacute; rostrum small; eyes oval. Antenna 1 : medium length, 0.2 times

body; peduncular article 1 short, length 1.3 times breadth, with dorsal crest, without tooth on distomedial margin,

without anterodistal projection; peduncular article 2 short. 0.13 times article 1, without anterodistal projection;

peduncular article 3 short, 0.13 times article 1; accessory flagellum short, 0.33 times primary flagellum.

3-articuIate, article 1 long. 1 times article 2; flagellum 13-articulate, without callynophore. Antenna 2 : subequal

in length to antenna 1: peduncle without brush setae, female weakly geniculate between peduncular articles 3-4,

article 3 long, 0.75 times article 4; flagellum well developed, 9-articulate, without thick setal brush.

Mouthpart bundle : subquadrate. Epistome and upper lip : unknown. Mandible : incisors vestigial; left lacinia

mobilis present, a small spine; molar vestigial; mandibular palp attached distally; article 1 short, length 1.3 limes

breadth; article 2 slender, length 6.8 times breadth, 2.3 times article 3, with 6 distal A2-setae. without D2-setae;

article 3 slender, distally truncate, long, length 4 times breadth, without proximal A3-setae or D3-setae. with

2 apical E3-setae. Maxilla 1 : inner plate narrow, without apical setae; outer plate small, narrow, with 6 spine-

teeth in modified 7/4 arrangement; outer row wilh ST1 to ST3 small, stout, weakly cuspidate, ST4 small, stout,

3-cuspidate, ST5 absent, ST6 small, stout, 2-cuspidate, ST7 small, broad, 3- to 4-cuspidate distally; inner row

with STA-STD absent; palp large, 2-articulate, with 3 short terminal spines, without subterminal setae, flag spine

present on distolateral comer, distomedial margin smooth. Maxilla 2 : inner plate narrow, outer plate broader.

Maxilliped : inner plate small, subrectangular. with 2 apical nodular spines, oblique setal row absent; outer plate

vestigial, subovate, without apical setae, apical spines or apical teeth, with 2 large distomedial spines,

submarginal setae long, simple; palp large. 4-articulate; article 2 broad, length 1.6 times breadth, 0.81 times

146

J. K. LOWRY & H. E. STODDART

article 3; article 3 long, slender, length 2.5 times breadth; dactylus well developed, with 1 terminal and

1 subterminal setae, unguis absent.

F|Gm12' —C‘ePidecrel‘a 'ropicalis sp. nov„ holotype female, 5.0 mm (MNHN-Am 4385), south of the Isle of Pine

New Caledonia. Scales represent 0.1 mm.

Source MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

147

148

J. K. LOWRY & H. E. STODDART

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : simple; coxa large, as long as coxa 2, anterior margin

slightly convex, anteroventral corner rounded, posterior margin straight; basis long, slender, length 4.5 times

breadth, anterior margin smooth, with simple setae; ischium long, length 2.9 times breadth; merus. posterior

margin without setae; carpus subrectangular, very long, length 4.6 times breadth, longer than (1.9 times)

propodus, without denticulate patch near postcrodistal margin; propodus large, subtriangular, length 3.1 times

breadth, tapering distally. posterior margin smooth, straight, with setae, without disto-medial setae, without

denticulate patch near posterior margin, palm absent; dactylus simple, with large plumose seta. Gnathopod 2 :

minutely chelate; coxa large, subequal in size to coxa 3; ischium long, length 3.6 times breadth; carpus long,

length 3.5 times breadth, posterior margin straight; propodus subrectangular, long, length 2.4 times breadth, palm

obtuse, with straight, serrate margin, postcrodistal comer without spines; dactylus reaching corner of palm,

posterior margin serrate.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus with 5 setae and 1 distal locking spine

along posterior margin; dactylus long, slender. Peraeopod 4 : coxa deeper than wide, with very large posterovcntral

lobe, anterior margin slightly rounded, posterior margin slightly sloping anteriorly; merus weakly expanded

anteriorly; propodus with 5 setae and 1 distal spine along posterior margin; dactylus long, slender. Peraeopod 5 :

coxa bilobate, posterior lobe slightly produced ventrally; basis moderately expanded posteriorly with posteroventral

lobe about as long as merus; merus expanded with rounded posterior margin; propodus with 4 spines and 2 distal

locking spines along anterior margin; dactylus long, slender. Peraeopod 6 : coxa small, not lobatc posteriorly;

basis, anterior margin rounded, basis expanded posteriorly with minutely crenate posterior margin, without

anteroventral lobe; merus expanded with rounded posteroproximal shoulder and straight posterior margin; propodus

with 4 spines and 2 distal locking spines along anterior margin; dactylus long, slender. Peraeopod 7 : basis

expanded posteriorly, posterior margin broadly rounded, posteroventral margin rounded; merus expanded, convex

posterior margin with 3 spines; propodus with 3 spines and 2 distal locking spines along anterior margin and no

spines along posterior margin; dactylus long, slender.

Oostegites : on peraeopod 5 only. Gills : from gnathopod 2 to peraeopod 6, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 3 : posteroventral corner narrowly rounded. Urosomites : dorsally

smooth. Uropod 1 : peduncle with 10 dorsolateral. 1 apicolateral, 1 dorsomedial and 1 apicomedial spines; rami

subequal in length, outer ramus with 2 dorsal spines; inner ramus with 1 dorsal spine. Uropod 2 : peduncle with

large dorsolateral flange, with 1 apicomedial spine, without spines along distal margin; outer ramus slightly

longer than inner ramus, rami without spines; inner ramus without constriction. Uropod 3 : peduncle well

developed, short, without dorsolateral flange, without dorsal spines, with 1 distal simple seta, without distoventral

spines; rami absent. Telson : shorter than broad, length 0.75 times breadth, entire, emarginate, without dorsal

setae, marginal penicillate setae or marginal spines, with 6 simple marginal setae.

Etymology. — The name reflects the lirst species of Clepidecrella reported from a tropical area.

Remarks. Clepidecrella tropicalis and C. cabinda J.L. Barnard, 1962, the only other species in the genus,

differ in the following ways : C. cabinda has no eyes; the flagellum of antenna 1 has only 5 articles, and of

antenna 2, only 4; mandibular palp article 2 has only 1 distal A2-seta; gnathopod 1 is weakly subchclate and the

carpus is not long and linear; the posteroventral comer of epimeron 3 is subquadrate; the peduncle of uropod 1 is

less spinose; uropod 3 is biramous; and the telson is cleft.

Distribution. — South of the Isle of Pines, New Caledonia, in 400 to 450 m depth.

Genus CORIOLISA gen. nov.

Diagnosis. — Antenna 1 with callynophore in male and female. Mandible : incisor distally placed; lacinia

mobilis present; lamina dentata weakly developed; accessory spine row on a raised, setose ridge, spines nodular;

molar, if present, a small setose flap. Maxilla 1 : outer plate with small apical spine-teeth; palp vestigial.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

149

Maxilliped with 3- or 4-articulatc palp, articles 1 and 2 may be partially fused. Gnathopod 1 : palm may be defined

by complex spine.

Type Species. — Coriolisa novacaledonia sp. nov.

SPECIES Composition. — Coriolisa contains two species : C. novacaledonia and C. sculptidentata (Ren in

Ren & Huang, 1991).

ETYMOLOGY. — Named for the R.V. Coriolis , the ship from which many of the tropical western South

Pacific species have been collected.

Remarks. — Lowry (1984) remarked on the similarity between Prachynella and Drummondia. Drummondia

differs in having a dorsally placed incisor, a well developed lamina dentata on the mandible and a 4-articulate

maxillipedal palp. The new species, Coriolisa novacaledonia , described below, obscures these differences. It has a

weakly developed lamina dentata ( Prachynella ), the spine-teeth on the outer plate of maxilla 1 are located apically

{Prachynella), it has a vestigial 1 -articulate palp on maxilla 1 ( Prachynella ) and it has a maxillipedal palp in which

articles 1 and 2 are partially fused leaving a 3-articulate maxillipedal palp ( Drummondia ). Finally, the spines of the

accessory spine row are modified into nodules sitting on a raised setose ridge, a character shared by Coriolisa

novacaledonia sp. nov. and C. sculptidentata (Ren in Ren & Huang, 1991). In fact these two species appear to be

closely related, but C. sculptidentata has a molar, a unique character among the pachynid group. We have seen

evidence of a vestigial lamina dentata in other species of Prachynella.

Distribution. — Coriolisa is known from the tropical western South Pacific Ocean and the western Southern

Ocean, in 400 to 1600 m depth.

Coriolisa novacaledonia sp. nov.

Figs 14-15

Material EXAMINED. — Lovaltv Islands. BiogeocaL : stn CP 317, 20°48.12'S, 166°53.16'E. west of Lifou,

1620-1630 m, 1 May 1987 : 1 (5,6.4 mm (MNHN-Am 4400).

Types. — The unique specimen is the holotypc.

Type Locality. — West of Lifou, Loyalty Islands. 20°48.12'S, 166°14.53.16'E, 1620 to 1630 m.

Diagnosis. — Mandible : accessory spine row, left with 3. right with 4 short nodular spines on a raised setose

ridge. Maxilla 1 : 4 short, stout, sculptured spine-teeth on inner row of outer plate. Gnathopod 1 : palm obtuse

with straight, serrate margin. Pcracopod 5 : posterior margin of basis with 2 large teeth. Uropod 3 : inner ramus

reduced, 0.7 times outer ramus.

Description. — Based on holotype male. 6.4 mm: female not known. Head : exposed, slightly longer than

deep, ventrally truncated with straight ventral margin; lateral cephalic lobe large, broadly rounded; rostrum absent;

eyes apparently absent. Antenna 1 : short, 0.1 times body; peduncular article 1 short (really massive), length

1.2 times breadth, with small midmedial tooth, with large posterodistal tooth, without anterodistal projection;

peduncular article 2 short, 0.1 times article 1, without anterodistal projection; accessory flagellum short, 0.3 times

primary flagellum, 2-articulate. article 1 short, 1 times article 2; flagellum 5-articulate. with strong 1-field

callynophore without spines or setae. Antenna 2 : subequal in length to antenna 1; strongly geniculate between

peduncular articles 3-4, article 3 long, 1.4 times article 4; flagellum well developed, 5-articulate, with thick setal

brush on articles 1 to 3.

Mouthpart bundle : quadrate, projecting anteriorly. Epistome and upper lip : fused, straight. Mandible : incisors

symmetrical, small, with slightly convex margins; left lacinia mobilis present, a short smooth peg; accessory

spine row, left with 3, right with 4 nodular spines on raised setose ridge, left lamina dentata present: molar absent;

150

J. K. LOWRY & H. E. STODDART

mandibular palp attached midway; article 1 short, length 1.6 times breadth; article 2 broad, length 2.4 times

breadth. 0.9 times article 3, without A2-setae, with 4 D2-setae; article 3 spatulate, long, length 3.2 times breadth,

without A3-setae. with 4 distal D3-setae and 2 apical E3-setae. Maxilla 1 : inner plate narrow, with 2 plumose and

3 simple apical setae; outer plate with 10 spine-teeth in modified 7/4 arrangement; outer row with ST1 to ST3

large, stout, weakly cuspidate. ST4-ST5 large, stout. 5-cuspidate, ST6 large, stout, 5- to 6-cuspidate. ST7 absent-

inner row with STA small, displaced from STB-STD, 4- to 5-cuspidate, STB small, slender, 3- to 4-cuspidate,

STC small, slender, 2-cuspidate, medial cusp long and 3-cuspidate, STD small, slender, 2- to 3-cuspidate; palp

vestigial. 1 -articulate. Maxilla 2 : inner and outer plates narrow, inner plate 0.76 times length outer plate.

Maxillipecl : inner plate small, subovate, without nodular spines, oblique setal row absent; outer plate large,

subovate, without subapical notch, without apical setae, medial spines vestigial, submarginal setae short, simple;

palp large. 3-articulatc. articles 1 and 2 fused; article 2 broad, length 2.85 times breadth, 2.5 times article 3;

article 3 short, slender, length 2 times breadth; dactylus reduced, with 1 subterminal seta, unguis present.

Peiaeonites : 1 to 7 dorsally smooth. Gnathopod 1 ; chelate; coxa large, as long as coxa 2, anterior margin

straight, posterior margin straight; basis long, slender, length 2.7 times breadth, anterior margin smooth, with

simple setae; ischium short, length 1.3 times breadth; merus, posterior margin with 1 seta, carpus extremely

compressed, hidden by propodus; propodus massive, subrectangular, length 1.4 times breadth, margins subparallel,

posterior margin smooth, subtly sinusoidal, without spines or setae, palm obtuse, margin straight, serrate,

posterodistal comer with complex spine; dactylus simple, without subterminal teeth or spines. Gnathopod 2 •

minutely chelate; coxa large, subequal in size to coxa 3; ischium long, length 3.3 times breadth; carpus long!

length 3.8 times breadth, posterior margin straight; propodus subrectangular. short, length 1.8 times breadth, palm

slightly obtuse, with straight, serrate margin, posterodistal comer without spines; dactylus reaching comer of

palm, posterior margin serrate.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus with 2 small setae and 1 distal spine

along posterior margin; dactylus long, slender. Peraeopod 4 : coxa as deep as wide, with large posteroventral lobe,

anterior margin rounded, posterior margin sloping anteriorly; merus weakly expanded anteriorly; propodus with

2 small setae and 1 distal spine along posterior margin; dactylus long, slender. Peraeopod 5 : coxa equilobate;

basis expanded, posterodorsal margin arched, forming large posteroproximal tooth, posterior margin with two

prominent teeth; merus expanded with rounded posterior margin; propodus with 3 small spines and 2 distal spines

along anterior margin; dactylus short, stocky. Peraeopod 6 ; coxa small, slightly lobate posteriorly; basis,

expanded, posterodorsal margin rounded, gently sloping into prominent tooth, posterior margin straight, with one

weak tooth; merus expanded with rounded posterior margin; propodus with 3 spines and 2 small distal spines

along anterior margin; dactylus short, stocky. Peraeopod 7 : basis expanded posteriorly, posterior margin rounded

with three posterodistal teeth, posteroventral corner subquadrate, posteroventral margin rounded; merus expanded

convex posterior margin with 3 spines; propodus with 3 spines and 1 distal spine along anterior margin and

1 distal spine along posterior margin; dactylus short, stocky.

Gills : from gnathopod 2 to peraeopod 6. not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 3 : posteroventral comer broadly rounded. Urosomites : dorsally

smooth. Uropod 1 : peduncle with 1 dorsolateral, 1 apicolateral and 1 apicomcdial spines; outer ramus slightly

onger than inner ramus, outer ramus with 1 lateral spine; inner ramus without spines. Uropod 2 : peduncle

without dorsolateral flange, with 1 apicolateral and 1 apicomedial spines; rami subequal in length, without spines;

inner ramus without constriction or proximal flange. Uropod 3 : peduncle short, length 1.2 times breadth, without

dorsolateral flange, without dorsal spines; rami lanceolate, inner ramus reduced, about 0.7 times outer ramus, outer

ramus 2-articulate, article 2 short, outer ramus with 1 medial spine. Telson : shorter than broad, length 0.8 times

breadth, entire, emarginate. with 8 marginal penicillate setae, without simple marginal setae, without marginal

spines.

Etymology. — The specific name refers to the general area where the species has been found.

Remarks. — Coriolisa novacaledonia is easily distinguished from C. sculptidentata by the lack of a mandibu¬

lar molar, cusps on the maxilla 1 inner row spine-teeth, the shape of the propodus and palm of gnathopod 1. the

ack of a dorsal tooth on peraeomte 5, the shape of the basis of peraeopod 5 and a small notch in the telson.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

151

DISTRIBUTION. — Coriolisa novacaledonia is currently known only from the Loyalty Basin, in about 1600 m

depth.

Fig. 14. — Coriolisa novacaledonia gen. nov., sp. nov., holotype male, 6.4 mm (MNHN-Am 4400), west of Lifou,

Loyalty Islands. Scales represent 0.1 mm.

152

J. K. LOWRY & H. E. STODDART

Fig. 15. Coriolisa novacaledonia gen. nov.. sp. nov., hololype male.

Loyalty Islands. Scales represent 0.2 mm.

6.4 mm (MNHN-Am 4400). west of Lifou,

Source : MNHN. Paris

LYSIANASSOID AMPHIPODA FROM THF. TROPICAL WESTERN SOUTH PACIFIC OCEAN

153

Genus CYCLOCARIS Stebbing, 1888

Cyclocaris tahitensis Stebbing, 1888

Cyclocaris tahitensis Stebbing, 1888 : 664. pi. 8; 1906 : 30, fig. 7. . — Della Valle, 1893 : 843, pi. 60, fig. 53. —

J.L. Barnard. 1958 : 91. — Gurjanova, 1962 : 85, fig. 17. — Thurston & Allen, 1969 : 358. — Wilson et al„

1985 : 1248, 1251. — Barnard & Karaman, 1991 : 479. — Vinogradov & Vinogradov, 1991 : 33. —

Vinogradov, 1993 : 43.

not Cyclocaris tahitensis - Chevreux. 1903 : 89; 1935 : 31, pi. 4, fig. 4 (= Cyclocaris sp.).

MATERIAL EXAMINED. — Austral Isles. SMCB, R.V. Marara (J. Poupin & J.K. Lowry coll.) : stn FRP-6,

21°47.9'S, 154°34.8'W, north-east of Maria Island, baited trap. 680 m, 8-9 August 1991 : 4 specimens (AM P42125). —

Stn FRP-7, 21°47.7'S, 154°43.4'W, north-east of Maria Island, baited trap, 500 m, 8-9 August 1991 : 8 specimens

(AM P42126). — Stn FRP-21, 22°29.3'S. 151°21.6'W, off Rurutu, baited trap, 840 m, 10-11 August 1991 :

120 specimens (AM P42127). — Stn FRP-23, 22°29.3'S, 151°21.6'W, off Rurutu, baited trap on sandy bottom, 490 m,

10-11 August 1991 : 13 specimens (AM P42128). — Stn FRP-24, 22°29.3'S. 151°21.9'W, off Rurutu, baited trap on

sandy bottom, 490 m, 10-11 August 1991 : 1 specimen (AM P42129). — Stn FRP-38, 23°18.6'S, 149°29.7,W. off

Tubuai, baited trap, 840 m, 12-13 August 1991 ; about 300 specimens (MNHN-Am 4777), about 1000 specimens

(AM P42130). — Stn FRP-40, 23°19.2'S, 149°28.9'W, off Tubuai, baited trap, 65 m, 12-13 August 1991 : 1 specimen

(AM P42131). — Stn FRP-54, 27°35.3'S, 144°15.5'W, off Rapa, baited trap, 870 m, 17-18 August 1991 : 1 specimen

(AM P42132). — Stn FRP-55, 27°35.3'S, 144°15.5'W. off Rapa, baited trap. 870 m, 17-18 August 1991 ; 505 specimens

(AM P42133). — Stn FRP-64, 27°35.5'S. 144°15.8'W, off Rapa, large baited trap, 750 m, 18-19 August 1991 :

1 specimen (AM P42134). — Stn FRP-65, 27°35.5'S. 144°15.8’W, off Rapa, baited trap, 750 m, 18-19 August 1991 :

1 specimens (AM P42135). — Stn FRP-66. 27°35.5'S, 144°15.8'W, off Rapa, baited trap, 750 m. 18-19 August 1991 :

8 specimens (AM P42136).

Remarks. — The recent use of small-mesh bailed iraps (Vinogradov & Vinogradov, 1991;

Vinogradov, 1993, and this paper) has shown ihal Cyclocaris tahitensis is a widespread, abundant scavenger in

the South Pacific Ocean.

Distribution, — Hamilton and Hess guyots, central North Pacific ocean (1740 to 1790 m); Tahiti (750 m);

Austral Isles (490 to 870 m); Nasca Ridge (560 m); East Pacific vent region west of Sala y Gomez, south-eastern

South Pacific Ocean (2024 to 2038 m).

Genus CYPHOCARIS Boeck. 1871

Cyphocaris bellona sp. nov.

Figs 16-18

Material EXAMINED. — New Caledonia. Biocal : stn CP 61, 24°H.67'S, 167°31.37'E to 24°10.67'S,

167°33.65’E, south of the Isle of Pines, beam trawl, 1070 m, 2 September 1985 ; 1 specimen (AM P42137). —

Stn CP 69, 23°51.38'S, 167°58.68'E to 23°52.21'S, 167°57.82'E, south of die Isle of Pines, beam trawl, 1220-1225 m,

3 September 1985 : 1 6 (probably not fully mature), 18 mm (MNHN-Am 4434).

Loyalty Islands. Biogeocal : stn CP 272, 21°00.04'S, 166°56.94'E, south-west of Pointe Lefevre, Lifou, beam

trawl, 1615-1710 m, 20 April 1987 ; 1 specimen (MNHN-Am 4431).

Chesterfield Islands. MUSORSTOM 5 : stn DC 321, 21°20.40'S, 158°02.20’E, west of Middle Bellona, 1000 m,

14 October 1986 : 1 2. 20 mm (MNHN-Am 4427).

Types. — The female, 20 mm (MNHN-Am 4427). is ihc holotype. The other specimens are paratypes.

Type Locality, — West of Middle Bellona, Chesterfield Islands, 21 °20.40'S, 158°02.20'E, in 1000 m depth.

Diagnosis, — Pcraconite 1 : produced into a long, narrow slightly down-turned process. Gnathopod 2 : palm

acute with convex, serrate margin. Peraeopod 5 : posterovcntral corner of basis produced into elongate.

154

J. K. LOWRY & H. E. STODDART

dorsoproximally serralc spur. Uropod 2 : outer ramus shorter than (about 0.7 times) inner ramus. Telson 1.1 times

as long as uropod 3. cleft 75%.

FIG. 16. — Cyphocaris bellona sp. nov., holotype female, 20 mm (MNHN-Am 4427), west of Bellona Reefs, Chesterfield

Islands, Coral Sea.

Description. — Based on holotype female, 20 mm (MNHN-Am 4427); paratype male, 18 mm (MNHN-Am

4434). Head : positioned under produced peraeonite 1, narrow, much deeper than long, extending well below

insertion of antenna 2; lateral cephalic lobe small, subacute; rostrum absent; eyes oval, not enlarged in adult male.

Antenna 1 : elongate, 0.6 times body; peduncular article 1 short, length 1.1 times breadth, without dorsal crest,

tooth on distomedial margin or posterodistal tooth, with short anterodistal projection; peduncular article 2 short,

0.4 times article 1, without anterodistal projection; peduncular article 3 long, 0.43 times article 1; accessory

flagellum very short, 0.16 times primary flagellum, at least 5-articulate. article 1 long, 4.3 times article 2 (male

long, 6.6 times article 2). not forming cap; flagellum 33-articuIate (male 34-articulate), with strong 2-field

callynophore in female and male, without posterodistal setae or spines, with 1 spine each on flagellar articles 3 and

6. calceoh absent in female and male. Antenna 2 : subequal in length to antenna 1; peduncle with weak brush setae

in female and male, weakly geniculate between peduncular articles 3-4, article 3 short. 0.23 times article 4 (male

weakly geniculate between peduncular articles 3-4, article 3 short. 0.39 times article 4), peduncular article 4

enlarged in male; flagellum well developed, at least 40-articulate (male at least 98), calceoli absent in female and

male.

Mouthpart bundle : subquadratc. Epistome and upper lip : separate; epistome long, straight; upper lip slightly

produced, rounded. Mandible : incisors symmetrical, small, with slightly convex margins; left lacinia mobilis

present, a stemmed distally cusped blade; accessory spine row without distal setal tuft, left and right rows each

with 3 long, slender, "bushy" spines, with 2 "bottle-brush" intermediate setae; molar proximally setose, distally

triturating; mandibular palp attached distally; article 1 short, length 1 times breadth; article 2 slender, length

5.1 times breadth, 1.4 times article 3, with 18 submarginal posterodistal A2-setae (male 20), without B2-setae or

D2-setae; article 3 slender, blade-like, long, length 4.3 times breadth, without A3-setac or B3-setae, with 57 (male

55) D3 -setae along most of posterior margin and 2 apical E3-setac. Maxilla 1 : inner plate tapering distally, at

least half of inner margin setose, with 9 plumose setae; outer plate with 1 1 spine-teeth in 6/5 arrangement; outer

row with ST1 to ST3 large, stout, multicuspidate, ST4 large, stout. 3-cuspidate, ST5 large, stout, 4-cuspidate,

ST6 large, stout, 5-cuspidate, ST7 contiguous with ST6, large, broad, 7-cuspidate medially; inner row with STA

large, slightly displaced from STB-STD. 3-cuspidate, STB large, broad, 3-cuspidatc, STC large, broad, 4-cuspidatc.

Source MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

155

Fig. 17. — Cyphocaris bellona sp. nov.. holotype female, 20 mm (MNHN-Am 4427); paralype male, 18 mm, MNHN-Am

4334; west of Bellona Reefs, Chesterfield Islands, Coral Sea. Scales for A1 and A2 represent 0.5 mm, remainder

represent 0.2 mm.

156

J. K. LOWRY & H. E. STODDART

STD large, broad, 5-cuspidate; palp large, 2-articulaie, with 11 long terminal spines, with 10 subterminal setae,

flag spine present on distolateral comer, distomedial margin smooth. Maxilla 2 : inner plate broad, outer plate

narrow, inner plate 1 times length outer plate. Maxilliped : inner plate large, subrectangular, with 3 apical nodular

spines, with 1 distal spine on lateral face near inner margin, oblique setal row strong with 19 plumose setae; outer

plate small, subovate, without subapical notch, with 7 apical plumose setae, without apical spines or apical teeth,

medial spines present, large, submarginal setae long, simple; palp large, 4-articulate; article 2 broad, length

1.5 times breadth, 1.1 times article 3; article 3 short, broad. length 1.7 times breadth; dactylus very large, blade¬

like, with 4 subterminal plumose setae, unguis vestigial.

Peraeonite 1 produced anteriorly into long, narrow, slightly down-turned process. Gnathopod l : simple; coxa

vestigial; basis very long, slender, length 5.2 times breadth, anterior margin smooth, with simple setae; ischium

short, length 0.9 times breadth, anterior margin smooth; merus, posterior margin with group of long simple setae

and patch of short setae; carpus subrectangular, short, length 1.5 times breadth, shorter than (0.9 times) propodus,

with patch of very fine setae near posterior margin and long simple setae along posterior margin; propodus large,'

subtnangular. length 2. 1 times breadth, tapering distally, posterior margin serrate, straight, with 6 spines, without

denticulate patch near posterior margin, palm absent; dactylus simple, with large subterminal tooth and row of

20 cuticular teeth along posterior margin. Gnathopod 2 : minutely subchelate; coxa vestigial; ischium long,

length 3.2 times breadth; carpus very long, length 4.7 times breadth, posterior margin straight; propodus

subrectangular. long, length 3.2 times breadth, posterior margin without strong distal spines, palm acute with

convex, serrate margin, posterodistal corner with 1 (male 1) medial spine; dactylus reaching corner of palm

posterior margin smooth with 1 large spine.

Peraeopod 3 : coxa vestigial; merus not expanded anteriorly, male and female mcrus-carpus without plumose

setae; propodus with 5 spines and 2 distal spines along posterior margin; dactylus long, slender. Peraeopod 4 :

coxa deeper than wide, with acutely produced posteroventral lobe, anterior margin broadly rounded, posterior

margin sinusoidal; merus not expanded anteriorly, male and female mcrus-carpus without plumose setae; propodus

with 3 spines and 2 distal spines along posterior margin; dactylus long, slender. Peraeopod 5 : coxa equilobate;

basis expanded posterovcntrally to form elongate spur with serrate dorsoproximal margin, merus not expanded

posteriorly; propodus with 9 spines and 2 distal spines along anterior margin; dactylus long, slender. Peraeopod 6 :

coxa small, not lobate posteriorly; basis slightly expanded posteriorly with incised posterior margin, without

anteroventral lobe; merus not expanded posteriorly: propodus with 9 spines and 2 distal spines along anterior

margin; dactylus long, slender. Peraeopod 7 : basis slightly expanded posteriorly, posterior margin sinusoidal,

crenate. posteroventral comer subquadrate, posteroventral margin straight; merus not expanded posteriorly, with

6 spines; propodus with 7 spines and 1 distal spine along anterior margin and 2 spines along posterior margin-

dactylus long, slender. ’

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 7, not pleated.

Pleomtes 1 to 3 dorsally smooth. Epimeron 1 : anteroventral comer rounded. Epimeron 3 : posteroventral

comer subquadrate. Urosomites : urosomite 1 with anterodorsal notch, without lateral flange; urosomite 3 without

small dorsolateral spine. Uropod I : without fine setae; peduncle with 3 dorsolateral and 1 apicolateral spines;

outer ramus, length 0.75 times inner ramus, outer ramus without spines; inner ramus with 5 lateral spines

Uropod 2 : without fine setae; peduncle without dorsolateral flange, with 1 apicolateral spine, without plumose

setae, without spines along distal margin; outer ramus 0.7 limes as long as inner ramus, inner ramus with

5 a,era* ^pmes 111 wcak acclivities; inner ramus without constriction. Uropod 3 : peduncle well developed, long,

ength 2.3 times breadth, without dorsolateral flange, with 1 apicomedial spine, with 8 midlateral setae, without

d is to ventral spines, with plumose setae in female and male; biramous. rami lanceolate, inner ramus reduced, about

.67 times outer ramus, outer ramus 2-articulate, article 2 short, rami without spines, plumose setae present in

lemale and male, lelson : longer than broad, length 3.7 times breadth, deeply cleft (75%), with 1 dorsal spine on

each lobe, without dorsal setae, distal margins truncated, without marginal penicillate setae, simple setae or

E i ymology. — Named for the Bcllona reefs near the type locality.

Source : MNHN. Paris

L.YSIANASSOID AMPHIPODA FROM THF, TROPICAL WESTERN SOUTH PACIFIC OCEAN

157

FlG. 18. — Cyphocaris bellona sp. nov., holotype female, 20 mm (MNHN-Am 4427), west of Bcllona Reefs, Chesterfield

Islands, Coral Sea. Scales represent 0.2 mm.

Source : MNHN. Paris

158

J. K. LOWRY & H. E. STODDART

Remarks. — The only species which appears to be closely related to Cyphocaris bellona is C. johnsoni

described by Shoemaker (1934) from waters above the Puerto Rican trench, and not reported since. Both species

have a posteroventral spur on the basis of peraeopod 5 in which only the dorsal margin is serrate. However, in

C. johnsoni peraeonite 1 is distinctly curved upwards, the palm of gnathopod 2 is extremely acute and concave,

uropod 3 reaches only 0.57 times the length of the telson (0.88 times in C. bellona) and the telson is cleft 90%.

Cyphocaris geyserensis Ledoyer, 1986, has a strongly serrate dorsal margin on the posteroventral spur of the basis

of peraeopod 5 and a weakly serrate ventral margin. It differs further from C. bellona in having a differently shaped

peraeonite 1 projection, a convex posterior margin on the basis of peraeopod 7, subequal rami on uropod 2 and a

more deeply cleft telson.

Distribution. — Chesterfield Islands to the Loyalty Islands, in 1000 to 1700 m depth.

Genus EURYTHENES Smith, 1882

Eurythenes cf. gryllus (Lichtenstein. 1822)

Material EXAMINED. — New Caledonia. R.V. Vauban : stn CA-1, 20°44'S, 166°27'E. baited trap 1000 m A

Intes, 19 February 1977 :2 c?, 1 juvenile (AM P28855).

Loyalty Islands. Biogeocal : stn CP 265, 21°04.09'S, 166°00.40'E, Loyalty Islands Basin, 1760-1870 m

18 April 1987 : 1 immature specimen (MNHN-Am 4465). — Stn CP 317, 20°48.12’S, 166°53.16’E, west of Lifou

1620-1630 m, 1 May 1987 : 1 6 (MNHN-Am 4403).

Wallis and Futuna Islands. Musorstom 7 : stn CP 550. 12°14.8'S, 177°28.0'W, Combe Bank, beam trawl 800-

si? AtVi8 l?92 (N?N”N,'Anm 4778)' ~ Sln CP 627‘ 1 1054-2'S- 179°31.4'W, Bayonnaise Bank, beam trawl,

597-600 m. 29 May 1992 : 2 d,3 9, 1 immature (MNHN-Am 4779). — Stn CP 632, 1 1°54.0'S, 179°31 5'W

Bayonnaise Bank, beam trawl, 595-600 m, 29 May 1992 : 1 6 (MNHN-Am 4780).

Austra! Isles. SMCB, R.V Marara : stn FRP-54. 27°35.3'S, 144°15.5'W, just off Rapa, baited trap in 870 m.

J.K. Lowry & J.M. Poupin, 17-18 August 1991 : 1 (AM P42138).

, Tuamotu Archipelago. SMCB. R.V. Marara : 22°16'S, 138°42'W, Fangataufa atoll, baited trap. 900-1000 m

J.M. Poupin : 2 d, 1 9 (AM P42139).

Remarks. — Although these specimens are very similar to Eurythenes gryllus evidence is accumulating

which suggests that the South Pacific populations are a separate species.

Genus F1GORELLA J.L. Barnard, 1962

Figorella tasmanica Lowry, 1984

Figorella tasmanica Lowry, 1984 : 86. figs 28-30.

m M,A™AL EXAMINED. — New Caledonia. Biogeocal : stn KG 201, 22°40.42'S, 166°32.72'E. south-west of

nZ£: 975 a£! 198, : SSS )4>' - S,n KG 2U- 22"4L8°,S' 166"32'53'E' °(

1 .pS'/imSa 1 ” CP 31?- 20‘4812'S' 16ra'16'E' “f «*•*«» l 1 ** 1987 :

Remarks. This is the first record of Figorella tasmanica since its description and significantly extends the

geographic and depth ranges.

Distribution. — Figorella tasmanica is known from : off the New South Wales coast, eastern Australia (615

to 1200 m); New Caledonia (595 to 1630 m).

Source : MNHN.

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

159

Genus HIPPOMEDON Boeck, 1871

Hippomedon vao sp. nov.

Figs 19-21

MATERIAL EXAMINED. — New Caledonia. Biocal : sin DW 51, 23°05.27'S, 167°44.95'E, south of the Isle of

Pines, 680-700 m, 31 September 1985 : 1 <3, 10 mm (MNHN-Am 4444). — Stn CP 75, 22°18.65'S, 167°23.30'E, north

of the Isle of Pines, 825-860 m, 4 September 1985 : 1 immature specimen (MNHN-Am 4378). — Stn DW 77, 22°15.32'S,

167°15.40'E, north-west of the Isle of Pines. 440 m, 5 September 1985 : 3 <3 (MNHN-Am 4794).

Types. — The male (MNHN-Am 4444) is the holotypc. The other specimens are paratypes.

Type Locality. — South of the Isle of Pines, 23°05.27'S, 167°44.95'E, 680 to 700 m.

Diagnosis. — Eye present. Maxilla 1 : inner plate with 2 apical, plumose setae. Gnathopod 1 : carpus long,

1.5 times propodus; palm acute with slightly convex, serrate margin. Epimcron 3 : posteroventral tooth small,

broad. Telson : length 1.2 times breadth, cleft 75%.

Fig. 19. — Hippomedon vao sp. nov., holotype male. 10 mm (MNHN-Am 4444), south of the Isle of Pines,

New Caledonia.

Description. — Based on male holotype, 10 mm; female not known. Head and body : without setae. Head :

exposed, deeper than long; lateral cephalic lobe large, narrow, subacute; rostrum absent; eyes absent in preserved

material, ventral lens present. Antenna 1 : short, 0.17 times body; peduncular article 1 short, length 1.1 times

breadth, without dorsal crest, with small midmedial swelling, without posterodistal tooth or anterodistal projec¬

tion; peduncular article 2 short, 0.29 times article 1. without anterodistal projection; peduncular article 3 short,

0.23 times article 1; accessory flagellum medium length, 0.42 times primary flagellum, 5-articuIate, article 1

short, 1.3 times article 2; flagellum 12-articulate. with strong 2-field callynophore with 6 large posterodistal

spines, without flagellar spines, calceoli present in adult male, proximal calceoli much larger than rest.

Antenna 2 : 0.7 times body length in male: peduncle with strong brush setae, weakly geniculate between

peduncular articles 3-4, article 3 short, 0.55 times article 4, peduncular articles 4 and 5 not enlarged in male;

flagellum well developed, 31 -articulate in male, calceoli present in adult male.

Mouthpart bundle : subquadrate. Epistome and upper lip : separate; epistome straight; upper lip slightly

produced, rounded. Mandible : incisors symmetrical, small, with slightly convex margins; left lacinia mobilis

present, a stemmed distally serrate blade; accessory spine row without distal setal luft, left and right rows each

160

J. K. LOWRY & H. E. STODDART

with 3 short, slender, "bushy" spines, without intermediate setae; molar columnar with fully triturating surface

large plumose seta absent: mandibular palp attached distally; article 1 short. length 1 times brcadth:°article 2

slender, length 3.5 times breadth. 1.2 times article 3, with 15 submarginal posterodistal A2-setae, without B2-

setac or D2-setae; article 3 lalcate, long, length 3.5 times breadth, without proximal A3-setae without B3-setae

wnh 21 D3-setae along most of posterior margin and 2 apical E3-setae. Maxilla 1 ; inner plate narrow with

- plumose apical setae, outer seta with denticulate row; outer plate with 1 1 spine-teeth in 6/5 arrangement- outer

row with ST1 to ST3 large, stout, weakly cuspidate, ST4 large, stout, 2-cuspidate, ST5 large, stout. 3-cuspidate,

“dfiS represent ^ m“'' 10 ”m (MNHN*A” «44>- “»“■ »f «» of Pines

Source : MNHN, Paris

LYSIANASSOID AMPH1PODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

161

ST6 large, stout, 7-cuspidate, ST7 contiguous with ST6. large, broad, 6-cuspidate; inner row with STA large,

slightly displaced from STB-STD, 3-cuspidate, STB large, broad, 3-cuspidate. STC-STD large, broad. 4-cuspidate;

palp large, 2-articulate. with 9 short terminal spines, with 1 subterminal setae, flag spine present on distolateral

comer, distomedial margin smooth. Maxilla 2 : inner and outer plates broad, inner plate 1 times length outer plate.

Maxilliped : inner plate large, subrectangular, with 3 apical nodular spines, with 2 distal spines on lateral face near

inner margin, oblique setal row strong with 15 plumose setae; outer plate medium size, subovate, without

subapical notch, without apical setae, with 1 apical spine, without apical teeth, medial spines present, large,

submarginal setae short, simple; palp large, 4-articulate; article 2 very broad, length 2.2 times breadth, 1.1 times

article 3; article 3 short, broad, length 1.4 times breadth; dactylus well developed, with 5 subtcrminal setae, unguis

present.

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : subchelate; coxa large, as long as coxa 2, anterior margin

slightly convex, anteroventral corner rounded, posterior margin slightly convex; basis long, slender, length

3.7 times breadth, anterior margin smooth, with simple setae; ischium short, length 1.3 times breadth; merus,

posterior margin with a few simple setae; carpus subrectangular. long, length 2.8 times breadth, longer than

(1.5 times) propodus, with long simple setae along posterior margin; propodus large, subovate, length 1.8 times

breadth, margins slightly converging distally. posterior margin smooth, straight, with 2 spines, without

denticulate patch near posterior margin, palm extremely acute, margin convex, serrate, posterodistal corner with

1 medial and 1 lateral spines; dactylus simple, without subterminal teeth or spines. Gnathopod 2 : minutely

subchelate; coxa large, subequal in size to coxa 3; ischium long, length 2.8 times breadth; carpus long, length

3.1 times breadth, posterior margin straight; propodus subrectangular, short, length 1.7 times breadth, posterior

margin without strong distal spines, palm transverse, with straight, serrate margin, posterodistal corner with

5 medial and 4 lateral spines; dactylus not reaching corner of palm, posterior margin serrate.

Peraeopod 3 : coxa large; merus not expanded anteriorly, male mcrus-carpus without plumose setae, female not

known; propodus with 4 spines and 1 distal spine along posterior margin; dactylus long, slender. Peraeopod 4 :

coxa deeper than wide, with large posteroventral lobe, anterior margin rounded, posterior margin sloping

anteriorly; merus not expanded anteriorly, male merus-carpus without plumose setae; propodus with 4 spines and

1 distal spine along posterior margin; dactylus long, slender. Peraeopod 5 : coxa equilobate; basis expanded with

posterior margin smooth; merus expanded with rounded posterior margin; propodus with 5 spines, 4 setae and

2 distal spines along anterior margin; dactylus long, slender. Peraeopod 6 : coxa small, not lobate posteriorly;

basis slightly expanded posteriorly with weakly crenate posterior margin, without anteroventral lobe; merus not

expanded posteriorly; propodus posterior margin 5 spines, 5 setae and 1 distal spine along anterior margin,

dactylus long, slender, straight and closing along palm of propodus. Peraeopod 7 : basis expanded posteriorly,

posterior margin rounded, minutely crenate, posteroventral comer rounded, posteroventral margin straight; merus

not expanded posteriorly with 2 spines; propodus and dactylus unknown.

Gills : from gnathopod 2 to peraeopod 7, not pleated.

Pleonites I to 3 dorsally smooth. Epimeron 1 : anteroventral corner rounded. Epimeron 3 : posteroventral

corner produced into small, broad tooth. Urosomites : dorsally smooth; urosomite 3 without small dorsolateral

spine. Uropod 1 : without fine setae; peduncle with 9 dorsolateral. 1 apicolateral. 9 dorsomedial and 1 apicomedial

spines; outer ramus slightly longer than inner ramus, outer ramus with 1 lateral spine; inner ramus with 2 medial

and 3 lateral spines. Uropod 2 : without fine setae; peduncle without dorsolateral flange, with 4 dorsolateral.

1 apicolateral, 2 dorsomedial and 1 apicomedial spines, without spines along distal margin; outer ramus slightly

longer than inner ramus, outer ramus with 5 lateral spines in weak acclivities; inner ramus with 3 medial and

5 lateral spines, inner ramus without constriction. Uropod 3 : peduncle well developed, short, length 1.4 limes

breadth, without dorsolateral flange, with 1 apicolateral and 3 apicomedial spines, without midlateral spines or

setae, with 4 distoventral spines, with 8 simple setae; rami lanceolate, subequal in length, outer ramus

2-articulatc, article 2 short, article 1 with 8 lateral spines; inner ramus with 2 medial and 6 lateral spines, plumose

setae present in male. Telson : longer than broad, length 1.2 times breadth, deeply cleft (75%), with 2 dorsal

spines on each lobe, distal margins oblique, without marginal pcnicillate setae or simple marginal setae, with

1 marginal spine on each lobe.

Etymology. — Named for the town of Vao on the Isle of Pines.

162

J. K. LOWRY & H. E. STODDART

F,^c “4)i — °f - - - —

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

163

Remarks. — According to Jarrett and Bousfield (1982) this species belongs in the large genus

Hippomedon. Hippomedon vao occurs in the group of species without the notched tooth on epimeron 3 and with a

short telson. It is most closely related to Hippomedon bandae Pirlot, 1933, from the Banda Sea, Indonesia. Both

species have a weakly developed posteroventral tooth on epimeron 3 and a short telson, but the bases of

peraeopods 5 and 7 are longer and more slender in H. bandae and the inner ramus of uropod 2 is shorter than the

outer.

Distribution. — Hippomedon vao is known from southern New Caledonia, in 440 to 860 m depth.

Genus ICHNOPUS Costa, 1853

Ichnopus malpatun Lowry & Stoddart, 1992

Ichnopus malpatun Lowry & Stoddart, 1992 : 210, figs 15-16.

Material EXAMINED. — New Caledonia. Musorstom 4 : stn DW 150, 19°23.40'S, 163°22.70’E, north of the

Belep Isles, 110 m, 14 September 1985 : 2 6 (MNHN-Am 4402).

CHALCAL 2 : stn DW 80, 23°26.70'S, 168°01.80’E, south of the Isle of Pines, 80-160 m, 31 October 1986 : 1 6

(MNHN-Am 4432).

Distribution. — Ichnopus malpatun is known from outside the barrier reef, Madang Lagoon, northern Papua

New Guinea (95 m); off the Grand Passage, northern New Caledonia (110 to 165 m) and off the Isle of Pines,

southern New Caledonia (80 to 160 m).

Genus KERGUELENIA Stebbing, 1888

Kerguelenia Stebbing, 1888 : 1219; 1906 : 11. . — Sars.1891 : 119. — Della Valle, 1893 : 786. — STEPHENSEN,

1929 : 51. — J.L. Barnard, 1969 : 346. — Ledoyer, 1986 : 770. — Diviacco & Ruffo, 1989 : 488. — Barnard &

Karaman, 1991 : 493.

Diagnosis. — Mandible : incisor, lacinia mobilis, accessory spine row and molar all absent. Maxilla 1 : inner

plate reduced, without apical setae; outer plate with reduced number of spine-teeth. Maxillipcd : inner and outer

plates reduced in size and spination. Gnathopod 1 : simple, linear. Peracopod 4 : coxa with very large

posteroventral lobe. Peraeopod 5 : basis linear to moderately expanded. Uropod 3 : inner ramus reduced or absent.

Type Species. — Kerguelenia compacta Stebbing, 1888, by monotypy.

SPECIES COMPOSITION. — The genus contains 15 taxa : Kerguelenia adeliensis Bellan-Santini, 1972;

K. antarctica K.H. Barnard, 1930; K. antiborealis Bellan-Santini & Ledoyer, 1987; K. borealis Sars, 1891;

K. b. japonica Gurjanova. 1962; K. b. ochotica Gurjanova, 1962; K. compacta Stebbing, 1888; K. eoa

Gurjanova, 1962; K. glacialis Schellenberg. 1926a; K. koutoumo sp. nov.; K. lifou sp. nov.; K. macropoda

Ledoyer, 1986; K. microphthalma Ledoyer, 1986; K. palpalis K.H. Barnard, 1932; K. reducta Ledoyer, 1977.

Distribution. — Kerguelenia is a cosmopolitan genus occurring in 15 to 3700 m depth.

Kerguelenia koutoumo sp. nov.

Figs 22-24

Material EXAMINED. — New Caledonia. Calsub : stn PL 20, 22°52.7'S, 167°23'E, south of the Isle of Pines,

555-616 m, 10 March 1989 : 1 <3, 4.0 mm (MNHN-Am 4782); 1 2, 4.2 mm, 1 juvenile (MNHN-Am 4783).

164

J. K. LOWRY & H. E. STODDART

Types. The male, 4.0 mm (MNHN-Am 4782) is the holotype. The female and juvenile arc paratypes.

Type Locality. — South of the Isle of Pines. New Caledonia. 22°52.7'S, 167°23’E, 555 to 616 m.

Diagnosis. — Eyes apparently absent. Maxilla 1: palp 1 -articulate. Antenna 1 : peduncular article 1 produced

dorsodistally. Peraeopod 5 : basis slightly expanded posteriorly. Peraeopod 7 : merus, anterior and posterior

margins subparallel, strongly produced postcroventrally. extending beyond carpus. Uropod 3 biramous.

FlGNew Oledonm k°U,OUm° Sp' n0V- hololyPe male- 4-0 mm (MNHN-Am 4782). south of the Isle of Ptnes,

Description. — Based on holotype male. 4.0 mm and paratype female 4.2 mm. Head : exposed, slightly

longer than deep, ventrally truncated with straight ventral margin; lateral cephalic lobe small, narrowly rounded;

rostrum absent; eyes apparently absent. Antenna 1 : medium length, 0.2 times body; peduncular article 1 short]

ength 1.3 times breadth, without dorsal crest or tooth on distomedial margin, with anterodistal lobe reaching

al way along article 2; peduncular article 2 short, 0.27 times article 1, without anterodistal projection; peduncular

ar ic e 3 long, 0 23 times article 1; accessory flagellum long, 0.56 times primary flagellum 3- to 4-articulate,

article 1 long, 2 times article 2 (male long. 2.2 times article 2); flagellum 5-articulate (male 6), without

ca ynophore in female (strong 1-field in male). Antenna 2 : subequal in length to antenna 1 (same in male);

peduncle wuhout brush setae in female or male, male weakly geniculate between peduncular articles 3-4, article 3

short, 0.52 times article 4; flagellum well developed, 5-articulate in male.

Mouthpart bundle : subquadrate. Epistome and upper lip : fused, straight. Mandible : incisors absent; laciniae

mobilis absent; molar absent; mandibular palp attached extremely distally; article 1 short, length 0 73 times

breadth; article 2 broad length 3.2 times breadth. 1.2 limes article 3, with 8 submarginal posterodistal A2-sctae,

without D2-setae; article 3 slender, blade-like, long, length 3.6 times breadth, without proximal A3-setae, with

1 1-14 D3-setae along most of posterior margin and 2 apical E3-setae. Maxilla 1 : inner plate absent; outer plate

c^Dkhte ST4 Srr 7/4 arran8emcn,: 0UIer row wi<h ST1 to ST3 small, stout, weakly

cuspidate ST4-ST5 absent, ST6 small, stout. 5-cuspidate, ST7 slightly displaced from ST6, small, stout

2~siihfprm C> i‘nnCr r°w WIth STA-STD absent; palp large, 1-articulate, with smooth apical margin, with

2 sublermiITal setae Hag spine absent, distomedial margin smooth. Maxilla 2 ; inner and outer plates narrow-

inner plate about 0 5 times length outer plate. Maxilliped ; inner plate small, subrectangular, with 1 apical nodula^

spine, oblique seta! row absent; outer plate vestigial, subrectangular, without apical setae or spines, with 1 apical

Source : MNHN. Paris L

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

165

tooth, without medial spines, submarginal setae long, simple; palp large, 4-articulate; article 2 broad, length

1.7 times breadth. 1 times article 3; article 3 long, broad, length 1.8 times breadth; dactylus well developed, with

2 subtcrminal setae, unguis absent.

Peraeoniies : 1 to 7 dorsally smooth. Gnathopod 1 : simple: coxa large, slightly shorter than coxa 2, tapering,

anterior margin slightly convex, anterovcntral comer rounded, posterior margin straight; basis long, slender, length

Fig. 23. — Kerguelenia kouloumo sp. nov., holotype male, 4.0 mm (MNHN-Am 4782); paratype female, 4.2 mm

(MNHN-Am 4783); south of the Isle of Pines, New Caledonia. Scales for MX1, MX2, U3, T represent 0.05 mm,

remainder represent 0.1 mm.

166

J. K. LOWRY & H. E. STODDART

5 limes breadth- antenor margin smoolh. with simple setae; ischium long, length 3.1 times breadth; merus

posterior margin with a few simple setae; carpus subrectangular. very long, length 5.5 times breadth and’l times

without denticulate patch near postcrodistal margin; propodus large, subrectangular, length 7 times breadth'

margins subparallel, posterior margin smooth, slightly concave, with setae, without denticulate patch new

posterior margin, palm absent; dactylus simple, with large plumose seta. Gnathopod 2 : minutely chelate- coxa

Source : MNHN. Paris jl

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

167

large, subequal in size to coxa 3; ischium long, length 3.1 times breadth; carpus long, length 3.7 times breadth,

posterior margin straight; propodus subrectangular, long, length 2.7 times breadth, palm obtuse, with convex,

smooth margin, posterodistal corner without spines; dactylus over-reaching comer of palm, posterior margin

smooth.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus with 2 spines and 2 distal spines along

posterior margin; dactylus short, stocky. Peraeopod 4 : coxa wider than deep, with very large posteroventral lobe,

anterior margin rounded, posterior margin of lobe evenly rounded; merus weakly expanded anteriorly; propodus

with 3 spines and 1 distal spine along posterior margin; dactylus short, stocky. Peraeopod 5 : coxa bilobate,

posterior lobe produced distoventrally; basis slightly expanded posteriorly, without posteroventral lobe; merus

expanded, posterior margin rounded, produced distally along carpus; propodus with 2 spines along anterior margin;

dactylus short, stocky. Peraeopod 6 : coxa large, not lobate posteriorly; basis expanded posteriorly with smooth

posterior margin, without anteroventral lobe: merus expanded, posterior margin rounded, produced distally along

carpus; propodus with 2 spines and 2 distal spines along anterior margin; dactylus short, stocky. Peraeopod 7 :

basis expanded posteriorly, posterior margin almost straight, smooth, posteroventral comer rounded, posteroventral

margin rounded; merus expanded, anterior and posterior margins subparallel, produced posterodistally beyond

carpus; propodus with 1 spine and 2 distal spines along anterior margin and 2 setae along posterior margin;

dactylus short, stocky.

Oosiegites : on peraeopods 4 and 5 (buds). Gills : from gnathopod 2 to peraeopod 6, not pleated.

Pleonites I to 3 dorsally smooth. Epimeron 3 : posteroventral comer produced, narrowly rounded. Urosomites :

urosomite 1 with anterodorsal notch. Uropod I : peduncle with 4 dorsolateral, 1 apicolateral. 3 dorsomedial and

1 apicomedial spines; outer ramus slightly longer than inner ramus, outer ramus with 3 dorsal spines; inner ramus

with 2 dorsal spines. Uropod 2 : peduncle with large dorsolateral flange, with 1 apicolateral and 1 apicomedial

spines, without spines along distal margin; outer ramus slightly longer than inner ramus, outer ramus with

2 dorsal spines; inner ramus with 2 dorsal spines, without constriction. Uropod 3 : peduncle well developed,

short, length 1.1 times breadth, without dorsolateral flange, without dorsal spines, without midlateral spines or

setae, without distoventral spines; biramous, rami lanceolate, inner ramus reduced, about 0.63 times outer ramus,

outer ramus 2-articulate, article 2 short, rami without spines. Telson : shorter than broad, length 0.73 times

breadth, entire, without dorsal setae, distal margin emarginatc, with 5-6 marginal pcnicillate setae, without simple

marginal setae or marginal spines.

Etymology. — Named for Koutoumo. the island nearest the type locality.

Remarks. — Kerguelenia koutoumo is compared to species with biramous third uropods, such as

K. adeliensis, the K. borealis complex, K. microphthalma and K. palpalis. It differs from K. adeliensis by the less

rounded basis and greatly extended merus of peraeopod 7. It differs from the other three species by not having a

posteroventral lobe on the basis of peraeopod 5. Kerguelenia koutoumo is easily distinguished from K. lifou by

the propodus of peraeopods 3 to 7 which arc more spinosc, not setose, uropods 1 to 3 which are more spinose and

uropod 3 which has well developed rami.

Distribution. — Kerguelenia koutoumo is known from southern New Caledonia, in 580 m depth.

Kerguelenia lifou sp. nov.

Figs 25-27

Material EXAMINED. —Loyalty Islands. Biogeocal : stn CP 373, 21°01.53'S, 166°57.41'E, south-west of

Point LefSvre, Lifou, 1920-2040 m. 20 April 1987 : 1 2 4.4 mm (MNHN-Am 4459).

Types. — The unique specimen is the holotype.

Type Locality. — South-east of Point Leffcvre, Lifou. Loyalty Islands, 21°01.53'S, 166°57.41'E, 1920 to

2040 m.

168

J. K. LOWRY & H. E. STODDART

nrn?1AHNHSIS 7 absenI' Maxilla 1 •’ PalP 2-articuIate. Antenna 1 : peduncular article 1 not

p educed dorsodistally. Pcraeopod 5 : basis slightly expanded posteriorly. Peraeopod 7 • mcrus expanded

c~3rlSus.C°nVCX P°S'er0diS,al margin- S'r0ng,y Pr0dUCCd P°steroventrally, extending beyond

Fig 25. Kerguelenia lifou sp. nov„ holotype female, 4.4

Loyalty Islands.

mm (MNHN-Am 4459), south-west of Point Leffevre, Lifou,

medial margTtf finely1 setose **’ ^ *** ^ °"

lcng,h 2, bnres breadth; dac,ylus we.l devehfped. ^ ^

Source : MNHN, Pari ;

LYSIANASSOID AMPH1PODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

169

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : simple; coxa large, as long as coxa 2, anterior margin

slightly convex, anteroventral corner rounded, posterior margin straight; basis long, slender, length 4.4 times

breadth, anterior margin smooth, with simple setae; ischium very long, length 4.7 times breadth; merus, posterior

margin with a few simple setae; carpus subrectangular, very long, length 4.2 times breadth and 1 times propodus,

without denticulate patch near posterodislal margin; propodus large, subtriangular, length 4.9 times breadth,

margins slightly converging distally, posterior margin smooth, subtly sinusoidal, with setae, without denticulate

patch near posterior margin, palm absent; dactylus simple, with large plumose seta. Gnathopod 2 : minutely

chelate; coxa large, subcqual in size to coxa 3; ischium long, length 3.8 times breadth; carpus long, length

3.3 times breadth, posterior margin straight; propodus subrectangular, long, length 2 times breadth, palm obtuse,

with straight, serrate margin, posterodistal corner without spines; dactylus over-reaching comer of palm, posterior

margin serrate.

Fig. 26. — Kerguelenia lifou sp. nov., holotype female, 4.4 mm (MNHN-Am 4459), south-west of Point Lefevre, Lifou,

Loyalty Islands. Scales represent 0.1 mm.

Peraeopod 3 : coxa large; merus slightly expanded anteriorly; propodus with 1 seta and 1 distal spine along

posterior margin; dactylus short, stocky. Peraeopod 4 : coxa wider than deep, with very large posterovcntral lobe,

anterior margin rounded, posterior margin of lobe evenly rounded; merus weakly expanded anteriorly; propodus

with 3 setae and 1 distal spine along posterior margin; dactylus short, stocky. Peraeopod 5 : coxa bilobatc,

posterior lobe strongly produced ventrally; basis slightly expanded posteriorly, without posteroventral lobe; merus

170

J. K. LOWRY & H. E. STODDART

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM TIIE TROPICAL WESTERN SOUTH PACIFIC OCEAN

171

broadly expanded with sloping posterior shoulder and straight posterior margin; propodus with 2 spines and

2 distal spines along anterior margin; dactylus short, stocky. Peraeopod 6 : coxa large, not lobate posteriorly;

basis expanded posteriorly with smooth posterior margin, without anterovcntral lobe; merus broadly expanded with

loping posteroproximal shoulder and straight posterior margin; propodus with 1 spines and 2 distal spines along

nterior margin; dactylus short, stocky. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly

ounded, minutely crenate. posteroventral corner rounded, posteroventral margin rounded; merus distally expanded,

.lightly convex posterior margin with 4 setae; propodus with 1 spine and 2 distal spines along anterior margin and

setae along posterior margin; dactylus short, stocky.

Oostegiies : on peraeopod 5 only (bud). Gills : from gnathopod 2 to peraeopod 6, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 3 : posteroventral comer subquadrate. Urosomites : dorsally

smooth. Uropod 1 : peduncle with 6 dorsolateral and 1 apicomedial spines; outer ramus slightly longer than inner

ramus, outer ramus with 1 dorsal spine; inner ramus with 1 dorsal spine. Uropod 2 : peduncle with large

dorsolateral flange, with 1 dorsolateral spine, without spines along distal margin; outer ramus slightly longer than

nner ramus, outer ramus with 1 dorsal spine; inner ramus without spines, without constriction. Uropod 3 :

leduncle well developed, short, length 1.2 times breadth, without dorsolateral flange, without dorsal spines,

nidlateral spines or setae or distovcntral spines; uniramous, ramus 1 -articulate, without spines. Telson : shorter

han broad, length 0.67 times breadth, entire, without dorsal setae, distal margins truncated, with 6 marginal

penicillate setae, without simple marginal setae or marginal spines.

Etymology. — Named for the island of Lifou in the Loyalty Islands.

Remarks. — Compared to other species with uniramous third uropods K. lifou differs from K. eoa and

K. glacialis in having a slightly expanded linear basis on peraeopod 5 without a posteroventral lobe. It further

differs from K. glacialis in not having a produced posteroventral comer on epimeron 3. It is remarkably similar to

K. comp acta, but K. compacta has well defined eyes and the merus on peraeopods 5 to 7 is less expanded posteri¬

orly. Kerguelenia antarctica has a dorsodistal lobe on the first peduncular article of antenna 1 and K. macropoda

has an enormously expanded and distally produced basis on peraeopod 7.

Distribution. — Kerguelenia lifou is known from the Loyalty Islands Basin, South-West Pacific Ocean, in

1920 to 2040 m depth.

Genus LEPIDEPECREELLA Schellenberg. 1926a

Lepidepecreella Schellenberg, 1926a : 281. — STEPHENSEN, 1931 : 6; 1935 : 101. — J.L. Barnard, 1966 : 68; 1969 :

347. — Ledoyer, 1986 : 774. — Barnard & Karaman. 1991 : 494.

Paracyclocaris K.H. Barnard, 1930 : 321 (type species : Paracyclocaris bidens K.H. Barnard, 1930, original designation).

Diagnosis. — Head deeper than long with well developed anterior keel. Mouthparts subconical. Maxilla 1 :

outer plate with 1 1 spine-teeth in a 7/4 arrangement. Gnathopod 1 : simple, slender, elongate, attenuate with coxa

vestigial. Gnathopod 2 : coxa vestigial. Telson short, entire.

Type Species. — Lepidepecreella ctenophora Schellenberg, 1926, by monotypy.

Species Composition. — Lepidepecreella contains 9 species : L. bidens (K.H. Barnard, 1930); L. charno

J.L. Barnard, 1966; L. ctenophora Schellenberg, 1926a; L. cymba (Goes. 1866); L. emarginata Nicholls, 1938;

L. oval is K.H. Barnard, 1932; L. pamanzi Ledoyer, 1986; L. sarcelle sp. nov. and L. tridactyla Bellan-Santini,

1972.

Distribution. — Lepidepecreella is mainly a bipolar genus (tropically submergent in bathyal basins)

occurring down to 2500 m depth.

172

J. K. LOWRY & H. E. STODDART

Lepidepecreella sarcelle sp. nov.

Figs 28-29

MATERIAL EXAMINED. — New Caledonia. Biocal : sin CP 75 7?°iRfiS's iA7°m7n'c ,, . , , ,

Pines. 825-860 m. 4 September 1985 : 1 specimen, sex not known. 3+ mm (incomplete) (MNHN-AmTs^^ " * 6 °f

Types. — The unique specimen is the holotype.

Type Locality. — North of the Isle of Pines, New Caledonia, 22°8.65'S. 167°23.30'E, 825 to 860 m.

Diacnosls. - Gnathopod 1 : dactylus without convex posterior margin. Gnathopod 2 : palm slightly obtuse

with serrate margin. Peraeopod 5 : basts with weakly developed posterodistal lobe.

Descr^ion Based on holotype, sex not known. Head : exposed, much deeper than long extending well

e^e°WovaT"l°"e™rr"sah;rlh 7“ f 'eVd 1 inser,i0n: laleral cephaIic l0be smaI1' subac“"!: ™«mm small;

3 Anle'"m 1 ■ short' P^uncular article 1 medium, length 1.5 times breadth, without dorsal crest

ldmedial swe ling or anterodtstal projection; peduncular article 2 short, 0.36 times article 1 without anterodistal

projec,,on: pcduncu'ar article 3 long. 0.24 times article 1; accessory flagellum long, 0 61 “pTm^

ageUum S-articufate. article 1 long, 2 times article 2; flagellum 7-articulate, with strong 2-field callynophore

ithoul setae or spines. Antenna 2 : subequal in length to antenna 1; peduncle without brush setae weaklv’

=sr,s"ce^clcs ~ 3 037

UPPe' U? : faSCd- f0m,ing 3 Str°“8'y projecling s“bac“le too'h

octwecn antennae 1 and 2. Mandible : incisors symmetrical, small, with slightly convex margins- left lacinn

1 S' timShTdiJ CUSp'fa'e f g: molar ves,i§ial: mandibular palp attached proximally; article 1 short length

^ .cl T alcme j , , SlCndCr- leng,h 3 ,imCS brcad,h- °‘78 limcs ^‘cle 3, without A2-se«ae or ^2^1

article 3 falcate, strongly tapering distally, long, length 3.9 times breadth, without proximal A3-setae with

w, Thorne ^ MaXiUa 7 , 1 TCI PlalC narT°W Wi,h0U' 3piCal SC,ae: OU,er broad

n ii spine teeth in 7/4 arrangement; outer row with ST1 to ST3 small, stout, multicuspidate ST4-ST5 small

SP de uST6 SmaU’ Sl0U'- 4'CUSp,dale- ST7 Conliguous with ST6, small, shorter ln^T6 sS

j Pld medially, inner row with STA large, broad. 1-cuspidate. STB-STD small stout 2-cusnidate- min

slipssissisii

slighUy oto” tta78'n T** PrOPOd,,S Subrectal'8“lar' '™8- 'a"8'b 3-1 times breach, palm

comerof palm, '^jslerior margin selrateSetTalC mar8'"- P°aer°diSIal ”i,h ‘ mb*al ■>«*■ «***

sSSSSSs sSSSHSSSSS

Source : MNHN, Pari ;

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

173

174

J. K. LOWRY & H. E. STODDART

prop^u^ WIIh 5 spines and 2 distal spines along posterior margin; dactylus long, stocky.

rPOd5: bl °,bt ’ poslenor lobe strong'y produced ventrally; basis weakly expanded, posterior margin

concave, pos.eroventral lobe extending along ischium; mems expanded, posterior margin rounded, produced d.stally

a ong carpus, propodus with 5 spines along anterior margin and 2 distal spines; dactylus long, stocky.

« “ k”°™' (MNHN-Am 4383), „f *. We ol

Source : MNHN. Pari i

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

175

Peraeopod 6 : coxa small, strongly lobate posteriorly; basis expanded posteriorly with smooth posterior margin,

without anteroventral lobe; merus expanded, posterior margin rounded, produced distally along carpus; propodus

with 5 spines along anterior margin and 2 distal spines; dactylus long, stocky. Peraeopod 7 : basis expanded

posteriorly, posterior margin almost straight, minutely crenate, posteroventral comer rounded, posteroventral

margin rounded; merus broadly expanded with rounded posteroproximal shoulder and straight posterior margin,

produced postered istally beyond carpus; propodus with 5 spines and 2 distal spines along anterior margin, and

2 setae along posterior margin; dactylus long, stocky.

Gills : from gnathopod 2 to peraeopod 7, not pleated. Pleonites , Urosomites and Telson unknown.

Etymology. — Named for the Sarcelle Passage between New Caledonia and the Isle of Pines.

Remarks. — As J.L. Barnard (1966) stated, Lepidepecreella bidens is the species most distinct from the

type species, L. ctenophora because of the posteroventral lobe on the basis of peraeopod 5. Lepidepecreella sarcelle

also has a posteroventral lobe and comes from the same geographic area as L. bidens. It is unfortunate that the

urosome is missing because it is extremely distinctive in this genus. Nonetheless L. sarcelle differs from L. bidens

as follows : the dactylus of gnathopod 1 does not have an excavate, convex posterior margin; the carpus and

propodus of gnathopod 2 are not as long and linear and the palm is slightly obtuse and convex, not concave; the

posteroventral lobe on the basis of peraeopod 5 is poorly developed and does not extend past the ischium.

Distribution. — Lepidepecreella sarcelle is known from southern New Caledonia, in 825 to 860 m depth.

Genus ONF.S1MOIDES Stebbing, 1888

Onesimoides abyssalis sp. nov.

Figs 30-32

Material EXAMINED. — Loyalty Islands. BlOCAL : stn CP 17, 20°34.54’S, 167°24.68'E to 20°34.62’S,

167°25.46'E, north-east of Lifou, 3680 m, 14 August 1985 : 1 9. 13.2 mm (MNHN-Am 4426); 1 6. 11.5 mm (MNHN-

Am 4784); 64 specimens (MNHN-Am 4461); 10 specimens (AM P42140). — Stn CP 72, 22°09.02'S, 167°33.18'E to

22°10.65'S, 167°33.78'E, north-east of Cape Coronation. Loyalty Islands Basin, 2100-2110 m, 4 August 1985 : 1 9

(MNHN-Am 4460).

Types. — The female (MNHN-Am 4426) is the holotype. The other specimens are paratypes.

Type Locality. — North-east of Lifou, Loyalty Islands. 20°34.54’S, 167°24.68'E to 20°34.54'S,

167°24.68'E, 3680 m.

Diagnosis. — Antennae : calccoli present in adult male. Gnathopod 1 : in male with large setal patch on

merus and propodus, palm changing with age from transverse to a midpalmar tooth and posterior cavity.

Pleonite 3 without dorsal carina. Urosomite 1 without lateral flange. Epimeron 3 : posteroventral comer narrowly

rounded. Uropod 3 : inner ramus about 0.6 times outer ramus.

Description. — Based on holotype female, 13.2 mm, MNHN-Am 4426; paratype male, 1 1.5 mm, MNHN-

Am 4784. Head : exposed, deeper than long; lateral cephalic lobe large, broadly rounded; rostrum absent; eyes

apparently absent. Antenna I : medium length; peduncular article 1 short, length about 1.2 times breadth;

peduncular article 2 short, 0.38 times article 1; peduncular article 3 long, 0.34 times article 1; accessory flagellum

medium length, 0.47 times primary flagellum. 4-articulate, article 1 long, 8.5 times article 2 (male long,

7.6 times article 2). forming cap covering callynophore; flagellum 16-articulate (male 14), with strong 2-field

callynophore in female and male, without flagellar spines, calceoli absent (present in adult male, 2). Antenna 2 .

slightly longer than antenna 1 (same in male); peduncle without brush setae (same in male), weakly geniculate

between peduncular articles 3-4, article 3 short, 0.40 times article 4 (male weakly geniculate between peduncular

176

J. K. LOWRY & H. E. STODDART

articleS 3-4. article 3 short 0.3 times article 4). peduncular articles 4 and 5 not enlarged in male or female-

adult maTeX P ' ,6'arUculate (malc I2)- wlthout >hi<* setal bmsh, calceoli absent in female (4 present in

Mouthpan bundle : subquadratc. Epistome and upper Up : separate; epistome slightly convex- unoer lin

slightly produced, rounded. Mandible : incisors symmetrical, small, with slightly convex margins; leftTcinia

Source : MNHN, Pari

LYSIANASSOID AMPHIPODA FROM TIIE TROPICAL WESTERN SOUTH PACIFIC OCEAN

177

mobilis present, a cuspidate peg; accessory spine row without distal setal tuft, left and right row each with 3 short,

thin, simple spines, without intermediate setae; molar with reduced column and convex triturating surface;

mandibular palp attached midway; article 1 short, length 1.1 times breadth; article 2 slender, length 4.1 times

breadth, 1.3 times article 3, with 20 submarginal posterodistal A2-setae (male 14), without B2-setae or D2-setae;

article 3 falcate, long, length 3.4 times breadth, with 2 (male 2) proximal A3-setae, without B3-setae, with

19 (male 21) D3-setac along most of posterior margin and 3 apical E3-setae. Maxilla 1 : inner plate narrow with

2 plumose apical setae, outer seta without denticulate row; outer plate with 1 1 spine-teeth in 6/5 arrangement;

outer row with ST1 to ST3 large, stout, multicuspidate, ST4 large, stout, 2-cuspidate, ST5 large, stout,

3-cuspidate, ST6 large, stout, 6-cuspidate, ST7 slightly displaced from ST6, large, broad, 6-cuspidate; inner row

with STA large, slightly displaced from STB-STD, 4-cuspidatc, STB-STC large, broad. 4-cuspidate, STD long,

slender, 4-cuspidate; palp large, 2-articulate. with 1 1 long terminal spines, with 1 subterminal seta, flag spine

present on distolateral comer, distomedial margin smooth. Maxilla 2 : inner plate narrow, outer plate broader,

inner plate 1 times length outer plate. Maxilliped : inner plate very large, subrectangular, with 3 apical nodular

spines, with 1 distal spine on lateral face near inner margin, oblique setal row strong with 1 1 plumose setae; outer

plate small, subovate, without subapical notch, with many fine apical setae, with 1 apical spine, medial spines

present, small, submarginal setae long, simple; palp large, 4-articulate: article 2 broad, length 1.6 times breadth.

1.3 times article 3; article 3 short, broad, length 1.5 times breadth; dactylus well developed, with 4 subterminal

setae, unguis present.

Peraeoniies : 1 to 7 dorsally smooth. Gnathopod 1 : sexually dimorphic; female, chelate; coxa reduced, anterior

margin slightly concave, anteroventral corner rounded, posterior margin slightly concave; basis long, slender,

length 3 times breadth, anterior margin smooth, with simple setae; ischium long, length 1.8 times breadth;

merus, posterior margin lined with long simple setae; carpus subtriangular, short, length 1.5 times breadth,

shorter than (0.7 times) propodus, without denticulate patch near posterodistal margin; propodus large,

subrectangular, length 1.9 times breadth, margins subparallcl, posterior margin smooth, strongly sinusoidal, with

7 groups of setae, without denticulate patch near posterior margin, palm obtuse, margin convex, smooth,

posterodistal comer with 1 medial and 1 lateral spines; dactylus simple, with subtcrminal tooth. Gnathopod 1 in

male, subchelate; basis long, slender, length 3.1 times breadth; merus with large brush of setae on medial face;

carpus subtriangular, short, length 1.2 times breadth, shorter than (0.61 times) propodus; propodus massive,

subrectangular, length 1.6 times breadth, slightly tapering distally, posterior margin smooth, convex, with dense

brush of setae on medial face, palm transverse anteriorly and posteriorly with large vertical step medially, smooth,

posterodistal comer with 1 medial and 1 lateral spines; dactylus simple, without subterminal teeth or spines.

Gnathopod 2 : minutely chelate; coxa intermediate in size between coxa 1 and coxa 3; ischium long, length

2.4 times breadth; carpus long, length 2.6 times breadth, posterior margin broadly lobate; propodus subquadrate,

short, length 1.3 times breadth, posterior margin without strong distal spines, palm obtuse, with straight, serrate

margin, posterodistal comer with 1 (male 1) medial spine; dactylus reaching comer of palm, posterior margin

rugose.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly, male and female merus-carpus without plumose

setae; propodus with 8 spines and 2 distal spines along posterior margin; dactylus short, stocky. Peraeopod 4 :

coxa deeper than wide, with large posteroventral lobe, anterior margin slightly rounded, posterior margin slightly

sloping anteriorly; merus weakly expanded anteriorly, male and female merus-carpus without plumose setae,

propodus with 9 spines and 2 distal spines along posterior margin; dactylus short, stocky. Peraeopod 5 . coxa

equilobatc; basis expanded with posterior margin minutely crenate; merus expanded with rounded posterior margin;

propodus with 6 setae and 2 distal spines along anterior margin: dactylus short, stocky. P eraeopod 6 : coxa small,

slightly lobate posteriorly; basis expanded posteriorly with minutely crenate posterior margin, basis and ischium

with anteroventral lobe; merus expanded with rounded posterior margin; propodus with 6 setae and 2 distal spines

along anterior margin; dactylus short, stocky. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly

rounded, minutely crenate, posteroventral corner rounded, posteroventral margin rounded; merus expanded

posterodistally with straight posterior margin; propodus with 5 spines and 2 distal spines along anterior margin

and 2 setae and 4 distal setae along posterior margin; dactylus short, stocky.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 6. not pleated.

178

J. K. LOWRY & H. E. STODDART

y y 6C3leS ,0r U1*3- T rePresent 0.2 mm, remainder represent 0.5 mm.

Source : MNHN, Pari\

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

179

Pleonite 3 dorsally smooth, without dorsal carina. Epimeron I : anteroventral comer rounded. Epimeron 3 :

posteroventral comer narrowly rounded. Urosomites : urosomite 1 with anterodorsal notch and low rounded boss,

without dorsal carina, without lateral flange; urosomite 3 without small dorsolateral spine. Uropod I : peduncle

with 3 dorsolateral, 1 apicolateral, 6 dorsomedial and 1 apicomedial spines; outer ramus slightly longer than inner

ramus, outer ramus with 3 dorsal spines; inner ramus with 3 dorsal spines. Uropod 2 : peduncle without

dorsolateral flange, with 7 dorsolateral, 1 apicolateral. 2 dorsomedial and 1 apicomedial spines; rami subequal in

length, outer ramus with 4 dorsal spines; inner ramus with 3 dorsal spines, without constriction. Uropod 3 :

peduncle well developed, short, length 1.2 times breadth, with dorsolateral flange, with 2 dorsolateral and

1 apicolateral spines, with 7 midlateral setae and 2 distoventral spines; biramous, rami lanceolate, inner ramus

reduced, about 0.6 times outer ramus, outer ramus 2-articulate, article 2 short, article 1 with 2 lateral and 1 medial

spines; inner ramus with 4 lateral spines, plumose setae absent in female (absent in male). Telsor : shorter than

broad, length 0.9 times breadth, entire, without dorsal spines or dorsal setae, distal margin rounded, with

2 marginal penicillatc setae and 2 simple marginal setae, without marginal spines.

Fig. 32. — Onesimoides abyssalis sp. nov., A. paratype male, 7.0 min; B. paratype male, 8.2 mm; C. paratype male,

8.8 mm; 1). paratype male, 9.5 mm; E. paratype male, 11.5 mm (MNHN-Am 4461); north-east of Lifou,

Loyalty Islands.

180

J. K. LOWRY & H. E. STODDART

Etymology. — The specific name refers 10 the depth at which this species lives.

r 0nesimoides ah'ssalis differs from 0. carinaius Stebbing. 1888. in having calceoli in the male-

a hough the shape of the palm of gnathopod 1 is similar in the male, the propodus is longer a"d not s wide’

there ts no canna on pleom.e 3; and no lateral flange on urosomite 3. Onesimoides abyssal* is easily

distinguished from O. castellatus Lowry & Stoddart. 1993. because of the strong patch of setae on the merus and

propodus of the male gnathopod 1 and the very different shape of the palm. Onesimoides abyssalis is most similar

' b"‘ ,hC ShaPe °f ‘he Pa‘m a',d ,he lenglh IO ”d,h ra,i0S °f lhc I"*—

Distribution. — Onesimoides abyssalis is known from the Loyalty Basin, in 3680 m depth.

Onesimoides carinatus Stebbing. 1888

Onesimoides carinaius Stebbine, 1888 • 648 nl 14- iqoa- a-> r,-„ o ,, _

- Thurston & Allen. 1969 : 363. -Lowry & Stoddart 1993 77 figs 13-^’ ' 7%' pL 6°' figS 39‘4L

Onesimoides cavimanus Pirlot, 1933 : 129, figs 4041. ’ 6

not Onesimoides cavimanus - Dahl, 1959 : 214, fig. 3 (= O mindoro)

no. Onesimoides cavimanus - Ledoyer. 1978 : 375, figs 9- 10b; 1986 : 794. fig. 309 (= Onesimoides sp )

1 juvenile (AM P42141). y -10.12,2 juveniles (MNHN-Am 4785); 1 <J .

w,,hRrerd s^I;T,ltST0DDART (‘,9? have tMntly carlmm and compared i,

related species. In these specimens the dorsal carina on pleonite 3 and urosomite 1 is present but verv weak

This record extends the genus Onesimoides onto the Pacific plate. ^

IslaSsZTo'sdO m,ln<l0n'Sia ^ ^ (9°° ‘° 2560 ■* R°““d' — Wadis and Fuluna

Genus ORCHOMENEI.LA Sars 1890

SSSpsi

Orchomenella abyssorum (Stebbing, 1888)

0nSST ■bym SKbb,n|' 1888 : 676' * 21-Lowav * Bullock, 1976 : 94. -BAR»RD & kakkm* 199, -

0rCmm^‘^ZZ'im:u'r >*» : 23, ,903 : 92, ,905b : 7, 1935 : 59.

Anonyx orm 8 £ ~ ST™NS™- 1925 : >“— D*»u 1954 : 282.

Walker,

Source : MNHN.

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

181

Orchomenella abyssorum - RUFFO, 1949 : 10 (list). — SCHELLENBERG, 1955 : 192. — J.L. Barnard, 1958 : 96; 1964b :

86, 89 (key). — Dahl, 1959 : 225. — Birstein & Vinogradov, 1960 : 188, fig. 8; 1962 : 41; 1964 : 164. —

Gurianova, 1962 : 433 (key). — Hurley, 1963 : 125, 126. — Thurston & Allen, 1969 : 364. — Sanderson,

1973 : 37. — Arnaud, 1974 : 572.

Orchomenopsis (Orchomene) abyssorum - COSTELLO el al., 1989 : 32.

Orchomene (Abyssorchomene) abyssorum - Barnard & Ingram, 1990 : 26, figs 15-17.

Abyssorchomene abyssorum - Thurston, 1990 : 262, 263. — PALERUD & VaDER, 1991 : 32.

? Orchomenella abyssorum - K.H. Barnard, 1932 : 69, figs 27b, 28. — NlCHOLLS, 1938 : 35, fig. 15. — Lowry, 1982 :

320.

? Orchomene abyssorum - AUSTIN, 1985 : 601.

not Orchomenopsis chilensis abyssorum Schellenberg, 1926a : 291, fig. 27 (= Orchomenella sp.).

MATERIAL EXAMINED. — Marquesas Islands. SMCB. R.V. Marara : stn 293, 9°47.30'S, 139°11.80'W, off

Hiva-Oa Island, baited trap at 900 m. J.M. POUPIN, 30 August 1990 : 1 <3, 7.0 mm (AM P42142).

Remarks. — The specimen from the Marquesas differs from Stebbing's illustrations in the following

points : the palp of maxilla 1 has 5 apical spines; the inner of the two apical spines on the maxilliped outer plate

is broader; articles 2 and 3 of the maxillipedal palp are slightly broader; there are a few setae on the anterior margin

of gnathopod 1 basis; the dorsal boss on urosomite 1 is slightly longer; uropod 1 inner ramus has a small spine

on the medial margin; uropod 2 outer ramus has fewer spines; the proximal spine of the three dorsal spines on

each lobe of the telson is very weak.

DISTRIBUTION. — Orchomenella abyssorum is known from off Buenos Aires, South Atlantic Ocean (3475 m);

north-eastern North Atlantic Ocean (1414 to 4849 m); Kcrmadec Trench (8210 to 8300 m), Marquesas Islands

(900 m). South Pacific Ocean; Galapagos vents, East Pacific Ocean (2491 m); Southern Ocean (210 to 3700 m).

Orchomenella distinctus Birstein & Vinogradov, 1960

Orchomenella distinctus Birstein & Vinogradov, 1960 : 191, fig. 10.

Orchomene ( Abyssorchomene ) distinctus - Barnard & INGRAM, 1990 : 22, figs 12-14. — VINOGRADOV, 1993 : 43.

Abyssorchomene ?distincta - THURSTON, 1990 : 263.

Orchomene distinctus - Barnard & Karaman, 1991 : 508.

Material EXAMINED. — Loyalty Islands. BlOGEOCAL : stn CP 225, 22°44.94'S, 166°19.84’E, Loyalty Islands

Basin, 2200-2280 m. 11 April 1987 : 1 9, 18 mm (MNHN-Am 4466).

Remarks. — There are no dorsal spines on the telson of this specimen; Birstein & Vinogradov (1960)

show one pair of spines, and Barnard & Ingram (1990) show two pairs.

Distribution. — Orchomenella distinctus is known from near Palau, south-western North Pacitic Ocean (0 to

2000 m); East Pacific vent region, south-eastern North Pacific Ocean (2635 m); East Pacific vent region west of

Sala y Gomez, south-eastern South Pacific Ocean (2024 to 2038 m); Loyalty Islands Basin, south-western Pacific

Ocean (2200 to 2280); tropical Atlantic Ocean (2300 to 4942 m).

Orchomenella gerulicorbis (Shulenberger & Barnard, 1976)

Orchomene affinis - BIRSTEIN & VINOGRADOV, 1955 ; 223, fig. 9.

Orchomene gerulicorbis Shulenberger & Barnard, 1976 : 243, figs 1-3. — THURSTON, 1979 : 56. Barnard &

Karaman, 1991 : 508.

Orchomenella gerulicorbis - THURSTON, 1990 : 265.

Orchomenella ( Orchomenopsis ) gerulicorbis - PALERUD & VaDER, 1991 : 41.

Material EXAMINED. — Austral Isles. SMCB. R.V. Marara , (J.K. Lowry & J.M. POUPIN coll.) ; sin FRP-55.

27°35.3'S, 144°15.5'W, just off Rapa, bailed trap in 870 m, 17-18 August 1991 : 22 specimens (AM P42143). —

Stn FRP-64, 27°35.5'S, 144°15.8’W, just off Rapa, large baited trap, 750 m, 18-19 August 1991 : 1C specimens

182

J. K. LOWRY & H. E. STODDART

(MNHN-Am 4786) : 48 specimens (AM P42144). — Stn FRP-66 27°35 5'S 144° 15 R'W i„ci rr d ,, , .

750 m, 18-19 Angus. 1991 : 4 specimens (AM P42145) J P3’ Sma11 bmIed

870°^ dN' 7 °rCh0mf'lC!la 8erulic°rbis is known from the Austral Isles, South Pacific Ocean (750 to

Atlantic S SS ”, m,: °" K“ril-KamChata- ^ Pactfic Ocean: norih-eas,

Genus PARAMBASIA Walker & Scott, 1903

Parambasia acuticaudata Ledoyer, 1984

Parambasia acuticaudata Ledoyer. 1984 : 84, fig. 41.

151^M8EWIAlLonAMfNE,D'r^AUS,nal Isles- SMCB- R V- Marara • (J-K- Lowry coll.) : stn FRP-16 22°28 5'S

lSl^^sIw^lag^on1 in^r^Tof s^ecirmnw Tam'^M^I^).11— ^lnWFRP-18,at22™8'n^,^

the reef platform, 1 m.^o' August, 1991 : ^pecime'ns SNHN5Zfl4787).‘llke br°Wn a‘8a' eXtremely common Jusl inside’

thp ' ~ ™eS? sPccimens 316 difficult to identify. They are not P. nui Myers, 1985 because article 2 of

gnathop^Tan^tJ £ t

the aitaaUon in Ledoyer, female material of P. acuticaudata la undetetmini We can see » ZS|

“ r Asi* from - s"shuy “

Distribution. — New Caledonia and Austral Isles, in less than 10 m depth.

Genus PROCYPHOCARIS J.L. Barnard, 1969

Procyphocaris indurata (K.H. Barnard, 1925)

Uristes induratus K.H. Barnard, 1925 : 333 pi. 34, fig 3

Procyphocaris primula J.L. Barnard, 1961 : 49 fig 18

Pprr°nr,Pn,Tr'5 ‘^Ura,US nGR,FFI™S' 1975 : 149. - LEDOYER, 1986 : 800, fig 312

Procyphocaris indurata - Barnard & Karaman, 1991 : 520. S

P- 23"44'51'S' ,66"54-!,4'E >» 23°56.52'S,

23°56.52'S, 166°40.55'E to 23°55 86'S ' ' specmen (MNHN-Am 4433). - Stn CP 58,

1 specimen (MNHN-Am 4435). ’ ' h °f Ihc Isle of Pines’ 2660-2750 m, 2 September 1985 :

1280™^.'’™^"°“''“ ,ndUm,a ^ kn°W" ^ S°“,hem Africa- A““»* a”“ Caledonia, in

Genus SOC ARNES Boeck. 1871

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

183

Diagnosis. — Upper lip produced beyond epistome in large rounded lobe. Mandible : lacinia mobilis present;

molar setose with or without distal vestigial triturating surface. Maxilla 1 : ST7 symmetrical; palp terminally

serrate. Gnathopod 1 : simple.

Type Species. — Lysianassa vahlii Krpyer, 1838. by monotypy.

Species Composition. — Socarnes contains 10 taxa : S. bidenticulatus (Bate, 1858); S. bidenticulatus

japonicus Gurjanova, 1962; ? S. eugenovi (Gurjanova, 1934); S. hartmani Hurley, 1963; S. rurulu sp. nov.;

S. tiendi sp. nov.; S. tuscarora sp. nov.; S. illudens (Hurley, 1963); ? S. unideniatus Schellenberg, 1931;

S. vahlii (Krpyer, 1838).

HURLEY (1963), originally placed S. illudens in Socarnoides because it had an incised inner ramus on uropod 2

(we do not regard this as a generic character). He pointed out that in other respects it resembled S. vahlii. We are

transferring S. illudens to Socarnes because of the shape of the epistome/upper lip complex, the midway insertion

of the mandibular palp and the non-swollen base of the maxilla 1 palp.

Socarnoides eugenovi possibly belongs in Socarnes. It is currently not well enough described to place with

certainty in any genus, but the upper lip/epistome complex is uncharacteristic of Socarnoides and similar to that of

Socarnes. The base of the maxilla 1 palp is not swollen and the mandibular palp appears to be inserted more

distally than in Socarnoides. Socarnes unidentatus has been placed in the genus Socarnoides (Hurley, 1963;

Barnard & Karaman, 1991), again probably because of the incised inner ramus of uropod 2. The species is

poorly described and illustrated, but based on the illustration of the epistome/upper lip it should not be considered

as a Socarnoides and is more likely to be in the genus Socarnes.

Barnard & Karaman (1991) assigned, with some doubt, Orchomene morbihanensis Bellan-Santini &

Ledoyer, 1974, to the genus Socarnes. However, the species belongs in (he genus Lysianella Sars, 1883. Among

other Lysianella characters it has the characteristic swollen peduncular article 4 of antenna 2 and the short

mandibular palp article 2.

Remarks. — Lincoln (1979) synonymized Socarnes and Socarnopsis Chevreux, 1911, because the type

species of both genera have the gills pleated on both sides. We agree that these genera are closely related, but we

think that consistent differences, such as the presence or absence of a lacinia mobilis, the triturating surface of the

molar and the terminal ornamentation of the maxilla 1 palp, indicate that the genera should remain separate. All

species attributed to these genera need to be carefully examined.

Distribution. — Socarnes is known from north-eastern Atlantic ocean; north-western and south-western

North Pacific Ocean; tropical south-western South Pacific Ocean; from immediate sublittoral to 550 m depth.

Socarnes rurulu sp. nov.

Figs 33-35

MATERIAL EXAMINED. — Austral Isles. SMCB, R.V. Marara, (J.K. Lowry, & J.M. POUPIN coll.) : sin FRP-7,

21°47.7'S, 154°43.4'W, north-east side of Maria Island, baited trap, 500 m, 8 August 1991 : 6 specimens (MNHN-Am

4788). — Stn FRP-23, 22°29.3'S, 151°21.9'W, off Rurutu, baited trap on sandy bottom, 490 m, 10 August 1991 : 1 ?$,

6.4 mm (AM P42147); 2 specimens (AM P42148). — Stn FRP-56, 27°36.2'S, 144°16.3’W, off Rapa, baited trap.

290 m, 7-18 August 1991 : 5 specimens (AM P42149).

Types. — The ?female, 6.4 mm (AM P42147) is the holotype. The other specimens are paratypes.

TYPE Locality. — Off Rurutu, Austral Isles, French Polynesia. 22°29.3'S, 151°21.9'W, 490 m.

Diagnosis. — Maxilla 1 : palp with vestigial terminal spines. Maxilliped : inner plate without nodular

spines. Gnathopod 2 : minutely subchelate, palm extremely obtuse with straight margin. Peraeopod 7 :

posteroventral comer of basis rounded. Telson moderately cleft, about 50%.

184

J. K. LOWRY & H. E. STODDART

Fig. 33. - Socarnes rurulu sp. nov., paratype ?female. 5.8 mm (AM P42148), off Rurutu, Austral Isles. French Polynesia.

Description. — Based on holotype ?female. 6.4 mm; male not known. Head : exposed, deeper than long;

lateral cephalic lobe large, broad, subacute; rostrum absent; eyes reniform. Antenna 1 : medium length, 0.24 times

body; peduncular article 1 short, length 1 times breadth, without tooth on distomcdial margin, posterodistal tooth

or anterodistal projection; peduncular article 2 short, 0.36 times article 1, without anterodistal projection;

peduncular article 3 long, 0.23 times article 1; accessory flagellum long, 0.5 times primary flagellum, 5-articulate.

article long, 2 times article 2, not forming cap; flagellum 9-articulate, with strong 2-field callynophore without

setae or spines, without flagellar spines, calceoli absent. Antenna 2 ; subcqual in length to antenna 1; peduncle

without brush setae, weakly geniculate between peduncular articles 3-4, article 3 short, 0.46 times article 4

peduncular articles 4 and 5 not enlarged; flagellum well developed, 10-articulate, calceoli absent.

Mouthpan bundle : subquadrate. Epistome and upper lip : separate; epistome concave; upper lip produced

apically rounded. Mandible ; incisors symmetrical, large, with strongly convex margins; left lacinia mobilis

present, a long slender peg; accessory spine row without distal setal tuft, left and right rows each with 4 slender to

stout serrate spines without intermediate setae; molar a strongly setose tongue; mandibular palp attached

prox.mally article 1 short, length 1.3 times breadth; article 2 slender, length 4.1 times breadth, 1.8 times

article 3 with 11 submarginal posterodistal A2-setae; article 3 falcate, long, length 3.4 times breadth, without

proximal A3-setae. with 7 proximal and 4 distal D3-se.ae and 2 apical E3-setae. Maxilla 1 ; inner plate narrow

with 2 plumose apical setae, outer seta without denticulate row; outer plate with 11 spine-teeth in

6A arrangement; outer row with ST1 to ST3 large, stout, multicuspidate, ST4 large, slender, 7-cuspidate, ST5-

ST6 large, slender. 9-cuspidate. ST7 slightly displaced from ST6, large, broad, 9-cuspidate; inner row with STA

large, broad, 4-cuspidate. STB large, broad, 6-cuspidate, STC large, broad, 5-cuspidate, STD small, slender

n ’ palp '^ge. 2-articulate, with 4 vestigial spines on seirate apical margin, without subtcrminal setae,

g spine present on distolateral comer (vestigial), distomcdial margin serrate. Maxilla 2 : inner plate narrow

Wi hnn, nL , r' ‘"T P °‘73 t,meS leng,h °U,Cr pla,e- MaxilliP^ : inner plate large, subrectangular.

suhnvaL ^ TT apparf?t y fused)- obl,que seIal row stron8 wiIh ^ plumose setae: outer plate small,

opines’ wi ho6, Ky 'rUnCa,Cd SHghtly COnCave’ with subapical nolch- without apical setae

or spmes, wthout medial spines, submarginal setae vestigial; palp large, 4-ar.iculate; article 2 slender, length

with breadth>.1-6 ,imes miclc 3= 3 'ong, slender, length 2.3 times breadth; dactylus well developed

with 3 subterminal setae, unguis present. p ’

conrlTlt! : 1,t0i7 d0rsally T°0lh' Gna,h°P°d 1 : simP,c: coxa iargc, as long as coxa 2, anterior margin

concave, anteroventral comer produced, rounded, posterior margin slightly concave; basis long, slender, length

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

185

3 6 times breadth, anterior margin smooth, with simple setae; ischium short, length 1.1 times breadth, anterior

margin smooth; merus, posterior margin with group of long simple setae; carpus subtriangular, short, length

1 6 times breadth, shorter than (0.9 times) propodus, without denticulate patch near posterodistal margin;

propodus large, subrectangular. length 2.4 times breadth, tapering distally, posterior margin smooth, straight,

withll spines, without disto-medial setae, without denticulate patch near posterior margin, palm absent; dactylus

simple, without subterminal teeth or spines. Gnathopod 2 : minutely chelate; coxa large, subcqual in size to

FIG. 34. — Socarnes rurutu sp. nov„ holo.ype ?female, 6.4 mm (AM P42147). off Ruru.u, Austral Isles. French

Polynesia. Scales represent 0.2 mm.

186

J. K. LOWRY & II. E. STODDART

^PofynSf' ’Srre^.a2Z': h°l0t,Pe ?fem“e' 6'“ m" <AM P42417)' rft R"™“- A“"»> I'1“-

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

187

coxa 3; ischium long, length 2.9 times breadth; carpus long, length 3.2 times breadth, posterior margin broadly

lobate; propodus subrectangular, long, length 2.2 times breadth, palm extremely obtuse, with straight, smooth

margin, posterodistal comer with 3 medial spines; dactylus over-reaching comer of palm, posterior margin

smooth.

Peraeopod 3 : coxa large; merus expanded anterodistally along carpus, merus-carpus without plumose setae;

propodus with 8 setae along posterior margin; dactylus short, slender. Peraeopod 4 : coxa deeper than wide, with

large posteroventral lobe, anterior margin slightly rounded, posterior margin slightly sloping anteriorly; merus

expanded anterodistally along carpus, merus-carpus without plumose setae; propodus with 9 setae along posterior

margin; dactylus short, slender. Peraeopod 5 : coxa equilobate (also very large); basis expanded with posterior

margin minutely crenate; merus expanded with rounded posterior margin; propodus with 4 setae along anterior

margin; dactylus short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis, anterior margin rounded

proximally, straight distally, basis expanded posteriorly with smooth posterior margin, without anteroventral lobe;

merus expanded with rounded posterior margin; propodus with 4 setae and 1 distal spine along anterior margin;

dactylus short, slender. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly rounded, minutely

crenate, posteroventral corner rounded, posteroventral margin rounded; merus not expanded posteriorly with

4 spines; propodus with 8 spines and 2 distal setae along anterior margin and 10 setae along posterior margin;

dactylus short, slender.

Oostegites : [unknown]. Gills : from gnathopod 2 to peraeopod 6, with weak horizontal pleating.

Pleonites 1 to 3 dorsally smooth. Epimeron 1 : anteroventral comer produced, narrowly rounded. Epimeron 3 :

posteroventral comer narrowly rounded. Urosomites : dorsally smooth; urosomite 3 with 1 small dorsolateral

spine. Uropod 1 : with long fine setae; peduncle with 10 dorsolateral, 1 apicolateral, 4 dorsomedial and

1 apicomedial spines; rami subequal in length, outer ramus with 7 lateral spines; inner ramus with 2 lateral and

2 medial spines. Uropod 2 : with long fine setae; peduncle with 3 dorsolateral. 1 apicolateral and 1 apicomedial

spines, without spines along distal margin; rami subequal in length, outer ramus with 6 lateral spines in weak

acclivities; inner ramus with 3 lateral spines and weak constriction. Uropod 3 : peduncle well developed, short,

length 1.7 times breadth, without dorsolateral flange, with 1 apicolateral spine, without midlateral spines or setae,

with 1 distoventral spine, without plumose setae; rami lanceolate, inner ramus reduced, about 0.84 times outer

ramus, outer ramus 2-articuIate. article 2 short, article 1 with 1 medial spine; inner ramus with 2 lateral and

1 medial spines, plumose setae present. Telson : longer than broad, length 1.5 times breadth, moderately cleft

(52%), without dorsal spines or setae, distal margins incised, without marginal penicillate or simple setae, with

1 marginal spine on each lobe.

Etymology. — Named for the island of Rurutu, near the type locality.

Remarks. — Socarnes rurutu and S. tuscarora are the first records of the genus on the Pacific Plate. They arc

closely related species and the differences between them are complicated because no specimens of S. rurutu appear

to be fully mature. Nonetheless S. rurutu differs from S. tuscarora and 5. tiendi. from New Caledonia, in not

having nodular spines on the inner plate of the maxilliped. having a rounded posteroventral corner on the basis ol

peraeopod 7. having stronger, though less numerous, spines on the rami of uropod 2 and having the telson cleft

only about 50%. It differs further from 5. tiendi in having an apically notched outer plate on the maxilliped and a

minutely chelate gnathopod 2.

DISTRIBUTION. — Socarnes rurutu is known only from Rurutu, Austral Isles, in 290 to 500 m depth.

Socarnes tiendi sp. nov.

Figs 36-38

MATERIAL EXAMINED. - New Caledonia. Lagon : sin 394, 22°44'S. 167°06'E, east of T.endi reef. Grand Recif

Sud 309 m, 23 January 1985 : 1 d, 12.5 mm (MNHN-Am 4386). f „• Ann m

B IOCAL : sin DW 43, 22°46.21'S, 167°14.50'E to 22°46.22'S, 167°14.50'E, south of the Isle of Pines, 400 m.

30 August 1985 : 1 juvenile (MNHN-Am 4376).

188

J. K. LOWRY & H. E. STODDART

Musorstom 4 : stn DW 234, 22°15.50'S, 167°08.30'E, off the Havannah

1 2, 16.0 mm (MNHN-Am 4429).

Channel, 350-365 m, 2 October 1985 :

Types. The female (MNHN-Am 4429) is the holotype. The other specimens are paratypes.

Type Locality. East of Tiendi reef. Grand Rdcif Sud, New Caledonia, 22°15.50'S, 167°08.30'E 350-

365 m depth.

Diagnosis. — Maxilla 1 : palp with vestigial terminal spines. Maxillipcd : inner plate with nodular spines.

Gnathopod 2 : subchelate, palm acute with large concave margin. Peraeopod 7 : postero ventral corner of basis

subquadrate. Telson moderately cleft, about 60%.

Fig. 36. — Socarnes tiendi sp. nov., holotype female 16 mm (MNHN-Am 4429), off Havannah Channel, New Caledonia.

Description — Based on holotype female. 16.0 mm; male paratype, 12.5 mm. Head : exposed, deeper than

ong lateral cephalic lobe large, broad, distally truncated (male slightly more acute); rostrum absent; eyes

rent form, not enlarged in adult male. Antenna 1 : short, 0.14 times body; peduncular article 1 short, length

. hmes breadth, without dorsal crest, without tooth on distomedial margin, posterodistal tooth or anterodistal

projec.ton; peduncular article 2 short, 0.27 times article 1, without anterodistal projection; peduncular article 3

, ° ~ U™es ^le accessory flagellum long, 0.7 times primary flagellum, 1 1-articulate, article 1 short,

.8 times article 2 (male short) not forming cap; flagellum 9-articulate (male 10+), with strong 2-field

ca ynophore in female and male, without flagellar spines, calceoli absent in female and male. Antenna 2 : length

1.7 times antenna 1 (0.7 times body length in male); peduncle without brush setae (same in male), female weakly

geniculate between peduncular articles 3-4, article 3 short. 0.51 times article 4 (male strongly geniculate between

peduncular articles 4-5. article 3 short, 0.52 times article 4), peduncular article 4 enlarged in male; flagellum well

developed. 26-art.culate (male 71), without thick setal brush, calceoli absent in female and male.

•mi ^aThPar,Abr<!!/ : -^quadrate. Epistome and upper lip : separate; epistome concave; upper lip produced,

”de ' Mandible : incisors symmetrical, large, with strongly convex margins; left lacinia mobil.s

present, a long slender peg; accessory spine row without distal setal tuft, left and right rows each with 4 long

slender bushy spines, intermediate setae absent; molar a strongly setose tongue; mandibular palp attached

T CnSh 2 limCS bread'h: artiCle 2 Slcndcr' leng,h 4 ,imes breadth, with 32 submarginal

With 15+ frnllMT? m f ’ W,th0UI D2-se,ae; micle 3 falca‘e. long, with 0 (male 1) proximal A3-setac,

i h 15+ (male 17) D3-sctae along most of posterior margin and 2 apical E3-setae. Maxilla J : inner plate narrow

with 2 plumose apical setae, outer seta without denticulate row; outer plate with 11 spine-teeth in

/5 drrangement- outer row with ST1 to ST3 large, stout, multicuspidate. ST4 large, slender, 10-cuspidate, ST5

Source : MNHN, Paris

LYSIANASSOID AMPIIIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

189

'"Ifflnhiurtu

Fig 37. — Socarnes liendi sp. nov., holotype female, 16 mm (MNHN-Am 4429), off Havannah Channel. New Caledonia,

paratypc male. 12.5 mm (MNHN-Am 4386); Grand Recif Sud. easi of Tiendi reef. New Caledonia. Scales represent

0.5 mm.

190

J. K. LOWRY & H. E. STODDART

large, slender, 9-cuspidate, ST6 large, stout, multicuspidate, ST7 slightly displaced from ST6, large, broad,

8- to 10-cuspidate; inner row with STA large, broad. 6-cuspidate, STB large, broad, 5-cuspidate, STC large, broad.

7 -cuspidate, STD large, broad. 6-cuspidate; palp large, 2-articulate, with 4-5 vestigial spines on serrate apical

margin, without subterminal setae, flag spine present on distolateral comer (vestigial), distomedial margin serrate.

Maxilla 2 : inner plate narrow, outer plate broader, inner plate 0.73 limes length outer plate. Maxilliped : inner

plate large, subrectangular, with 3 apical nodular spines, oblique sctal row strong with 14 plumose setae; outer

plate small, subovate, without subapical notch, without apical setae, apical spines, apical teeth or medial spines

submarginal setae vestigial; palp large, 4-articulate; article 2 slender, length 3.7 times breadth, 1.6 times article 3;’

article 3 long, slender, length 2.9 times breadth: dactylus well developed, with 5 subterminal setae, unguis present.

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : simple; coxa large, as long as coxa 2, anterior margin

concave, anteroventral comer produced, rounded, posterior margin straight; basis long, slender, length 3.3 times

breadth, anterior margin smooth, with simple setae; ischium short, length 1.2 times breadth, anterior margin

smooth; merus. posterior margin with group of long simple setae and patch of short setae; carpus subtriangular,

short, length 1.6 times breadth, longer than (1.1 times) propodus. without denticulate patch near posterodistal

margin; propodus large, subrectangular. length 2.1 times breadth, tapering distally, posterior margin smooth,

straight, with 13 spines, without disto-medial setae, without denticulate patch near posterior margin, palm absent;

dactylus simple, without subterminal teeth or spines. Gnathopod 2 : subchelate; coxa large, subcqual in size to

coxa 3; ischium long, length 3.3 times breadth; carpus long, length 2.6 times breadth, posterior margin broadly

lobate; propodus subrectangular. short, length 1.7 times breadth, palm acute, with large, concave, smooth margin,

posterodistal corner without spines; dactylus not reaching corner of palm, posterior margin serrate.

Peraeopod 3 : coxa large; merus expanded anterodistally along carpus, male and female mcrus-carpus without

plumose setae; propodus with 10 setae and 2 distal spines along posterior margin; dactylus short, slender.

Peraeopod 4 : coxa deeper than wide, with large posteroventral lobe, anterior margin slightly rounded, posterior

margin slightly sloping anteriorly; merus expanded anterodistally along carpus, male and female merus-carpus

without plumose setae; propodus with 9 setae and 2 distal setae along posterior margin; dactylus short, slender.

Peiaeopod 5 : coxa cquilobate (very large); basis expanded with posterior margin minutely crenate; merus expanded

with rounded posterior margin; propodus with 12 spines and 2 distal spines along anterior margin; dactylus short,

slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis, anterior margin rounded proximally, straight

distally, basis expanded posteriorly with minutely crenate posterior margin, without anteroventral lobe; merus

expanded with rounded posterior margin; propodus with 10 spines and 2 distal spines along anterior margin;

dactylus short, slender. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly rounded, minutely

crenate, posteroventral corner subquadrate, posteroventral margin straight; merus not expanded posteriorly with

9 spines; propodus with 7 spines and 2 distal spines along anterior margin and 10 setae along posterior margin-

dactylus long, slender. 6 ’

pleaUng6^5 ! fr°m gna'h0p0d 2 '° PeraeoPod 5- Gills '■ from gnathopod 2 to peraeopod 7, with strong horizontal

Pleomtes 1 to 3 dorsally smooth. Epimeron 1 : anteroventral comer produced, narrowly rounded. Epimeron 3 :

posteroventral corner narrowly rounded. Urosomites : urosomite 1 with anterodorsal notch, urosomite 3 without

small dorsolateral spine. Uropod 1 : with long fine setae; peduncle with 28 dorsolateral, 1 apicolateral.

6 dorsomedial and 1 apicomcdial spines; rami subequal in length, outer ramus with 7 lateral and 7 medial spines;

inner ramus with 16 lateral spines. Uropod 2 : with long fine setae; peduncle with 8 dorsolateral and 2 dorsomedial

spines, without spines along distal margin; rami subequal in length, outer ramus with 21 lateral spines in weak

acclivities; inner ramus with 4 medial and 5,1 lateral spines and weak constriction. Uropod 3 : peduncle well

developed long, length 2 times breadth, without dorsolateral flange, with 3 dorsolateral, 2 apicolateral and

apicomcdial spines, with 12 midlateral spines and 1 distoventral spines, with plumose setae in female and male-

ram, lanceolate. inner ramus reduced, about 0.84 times outer ramus, outer ramus 2-articulate, article 2 short'

article 1 with 1 medial spine; inner ramus without spines (male with 2 lateral spines), plumose setae present in

emale and male. Telson : longer than broad, length 1.6 times breadth, moderately cleft (60%), without dorsal

spines with row ol 5 dorsal setae on each lobe, distal margins truncated, with 1 marginal penicillate seta and

1 simple marginal seta on each lobe, without submarginal spines.

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

191

192

J. K. LOWRY & H. E. STODDART

Etymology. — Named for Tiendi reef near ihe lype locality.

Remarks. — Socarnes tiendi differs from S. rurutu and S. tuscarora in having a smoothly rounded apical

margin on the outer plate of the maxilliped and a large gaped palm on gnathopod 2 in the male and female. In

other respects S', tiendi and S. tuscarora are similar. Socarnes tiendi differs further from S. rurutu in having nodular

spines on the inner plate of the maxilliped. a subquadrate comer on the basis of peraeopod 7. more numerous

spines on the rami of uropod 2 and a more deeply cleft telson.

distribution. — Socarnes tiendi is known only from southern New Caledonia, in 350 to 365 m depth.

Socarnes tuscarora sp. nov.

Figs 39-41

Material EXAMINED. — Wallis and Futuna Islands. Musorstom 7 : stn DW 555 11°47’S 178°19'W

Tuscarora Bank. South Pacific Ocean, 540-542 m. 19 May 1992 : adult <J, 19.5 mm (MNHN-Am 4789)

Indonesia. Karubar : DW 32. 5°17'S, 132°51'E, Kei Islands. 170-206 m. 26 October 1991 : 1 6 (PPPO).

TYPES. — The male, 19.5 mm (MNHN-Am 4789) is the holotype.

Type Locality. — Tuscarora Bank, South Pacific Ocean, 1 1°47'S, 178°19'E, 540-542 m depth.

Diagnosis. — Maxilla 1 : palp with vestigial terminal spines. Maxilliped : inner plate with nodular spines.

Gnathopod 2 : minutely subchelate, palm slightly obtuse with straight margin. Peraeopod 7 : posterovcntral

comer of basis subquadrate. Telson moderately cleft, about 60%.

FlG'Ocean~ SoCarmS ,uscarora SP‘ nov" holo'yP*= male, 19.5 mm (MNHN-Am 4789). Tuscarora Bank. South Pacific

Description. Based on holotype male, 19.5 mm; female not known. Hpnd ■ pynneed ihon

Source : MNHN. Paris

LYSIANASSOID AMPH1PODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

193

peduncular article 3 short, 0.19 times article 1; accessory flagellum medium length, 0.35 times primary flagellum,

8-articulate, article 1 short, 2.4 times article 2, not forming cap; flagellum 20-articulate, with strong 2-field

callynophore in male, without flagellar spines, 9 calceoli present in adult male. Antenna 2 : much longer than

body in male; peduncle with strong brush setae in male; male strongly geniculate between peduncular articles 4-5.

article 3 short, 0.51 times article 4; peduncular article 4 short, broad, length 3.4 times breadth; flagellum well

developed, at least 138-articulate in male, calceoli present in adult male.

FIG. 40. -Socarnes tuscarora sp. nov., holotype male. 19.5 mm (MNHN-Am 4789), Tuscarora Bank. South Pacific

Ocean. Scales represent 0.5 mm.

194

J. K. LOWRY & H. E. STODDART

,USCar"a sp- nov- h0l0type male- 19.5 mm (MNHN-Am 4789), Tuscarora Bank, South Pacific

ucean. Scales represent 0.5 mm.

Source : MNHN . Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

195

Mouthpart bundle : subquadrate. Epislome and upper lip : separate; epistome straight; upper lip produced,

apically rounded. Mandible : incisors symmetrical, large, with strongly convex margins; left lacinia mobilis

present, a long slender peg; accessory spine row wilhoul distal setal tuft, left and right rows each with 4 short,

slender, "bushy" spines, without intermediate setae; molar setose with rudimentary distal triturating surface;

mandibular palp attached proximally; article 1 short, length 1.6 times breadth; article 2 broad, length 3.8 times

breadth, 1.5 times article 3, with 31 submarginal posterodistal A2-setae; article 3 falcate, long, length 4.9 times

breadth, with 1 proximal A3-seta, with 12 proximal and 7 distal D3-setae and 3 apical E3-setae. Maxilla 1 : inner

plate narrow, with 2 plumose apical setae, outer seta without denticulate row; outer plate with 1 1 spine-teeth in

6/5 arrangement; outer row with ST1 to ST3 large, stout, multicuspidate, ST4, large, slender, 9-cuspidate, ST5

large, slender, 1 1-cuspidate, ST6 large, broad, 12-cuspidate, ST7 slightly displaced from ST6, large, broad,

1 1-cuspidate; inner row with STA large, broad. 4-cuspidate, STB large, broad, 7-cuspidate, STC large, broad,

6-cuspidate, STD long, slender, 7-cuspidate; palp large, 2-articulate, with vestigial spines on serrate apical margin,

without subterminal setae, flag spine present (vestigial) on distolateral corner, distomedial margin serrate.

Maxilla 2 : inner plate narrow, outer plate broader, inner plate 0.69 times length outer plate. Maxilliped : inner

plate large, subrectangular, with 3 apical nodular spines, oblique setal row strong with 4 distal plumose setae and

double row of 10 medial plumose setae; outer plate small, subovate, distomedial margin obliquely truncated and

slightly concave, without subapical notch, without apical setae, apical spines or medial spines, submarginal setae

absent; palp large, 4-articulate; article 2 extremely slender, length 3.7 times breadth, 1.7 times article 3; article 3

long, slender, length 2.8 times breadth: dactylus well developed, with 4 subterminal setae, unguis present.

Peraeoniies : 1 to 7 dorsally smooth. Gnathopod 1 : simple; coxa large, as long as coxa 2, anterior margin

concave, anteroventral corner produced, rounded, posterior margin straight; basis long, slender, length 3.9 times

breadth, anterior margin smooth, with simple setae; ischium short, length 1.2 times breadth, anterior margin

smooth; merus, posterior margin with group of long simple setae and patch of short setae; carpus subtriangular,

short, length 1.5 times breadth, subequal in length to propodus, without denticulate patch near posterodistal

margin; propodus large, subtriangular, length 2.1 times breadth, tapering distally, posterior margin smooth, subtly

sinusoidal, with 14 spines, without disto-medial setae, without denticulate patch near posterior margin, palm

absent; dactylus simple, without subterminal teeth or spines. Gnathopod 2 : minutely subchelate; coxa large,

subequal in size to coxa 3; ischium long, length 3.5 times breadth; carpus long, length 3.1 times breadth,

posterior margin broadly lobate; propodus subrectangular, long, length 2.3 times breadth, palm slightly acute,

with straight, smooth margin, posterodistal corner without spines; dactylus reaching corner of palm, posterior

margin serrate.

Peraeopod 3 : coxa large; merus expanded anteriorly, male mcrus-carpus with plumose setae, female not

known; propodus with 28 setae along posterior margin; dactylus short, slender. Peraeopod 4 : coxa deeper than

wide, with large posterovenlral lobe, anterior margin rounded, posterior margin sloping anteriorly; merus expanded

anteriorly, male merus-carpus with plumose setae; propodus with 26 setae along posterior margin; dactylus short,

slender. Peraeopod 5 : coxa equilobate (also very large); basis expanded with posterior margin minutely crenate;

merus expanded with rounded posterior margin; propodus with 10 spines and 2 distal spines along anterior margin;

dactylus short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis, anterior margin rounded

proximally, straight distally, basis expanded posteriorly with minutely crenate posterior margin, without

anteroventral lobe; merus expanded with rounded posterior margin; propodus with 14 spines and 1 distal spine

along anterior margin; dactylus short, slender. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly

rounded, minutely crenate, posterovenlral corner subquadrate, posteroventral margin straight; merus not expanded

posteriorly with 9 setae; propodus with 13 spines and 2 distal spines along anterior margin and 18 setae along

posterior margin; dactylus short, slender.

Gills : from gnathopod 2 to peraeopod 7, with weak horizontal pleating.

Pleonites I to 3 dorsally smooth. Epimeron 1 : anteroventral corner produced, narrowly rounded. Epimeron 3 :

posteroventral comer narrowly rounded. Urosornites : urosomite 1 with antcrodorsal notch; urosomite 3 with

1 small dorsolateral spine. Uropod 1 : with long fine setae; peduncle with 35 dorsolateral, 8 long dorsomedial and

1 apicomedial spines; outer ramus slightly longer than inner ramus, outer ramus with 26 lateral spines; inner

ramus with 8 medial and 6 lateral spines. Uropod 2 : without long fine setae; peduncle with 13 dorsolateral and

1 apicolateral spines, with 3 large midmedial spines, without plumose setae, without spines along distal margin;

196

J. K. LOWRY & H. E. STODDART

rami subequal in length, outer ramus with 26 lateral spines; inner ramus with 3 medial and 4,1 lateral spines, and

with weak constriction. Uropod 3 : with long fine setae; peduncle well developed, short, length 1.7 times breadth,

without dorsolateral flange, with 3 dorsolateral and 6 dorsomedial spines, without midlateral spines or setae, with

1 distoventral spine, with 9 simple setae, plumose setae present; rami lanceolate, subequal in length, outer ramus

2-articulate, article 2 short, article 1 with 1 medial spine; inner ramus without spines; plumose setae present in

male. Telson : longer than broad, length 1.6 times breadth, moderately cleft (61%), without dorsal spines, with

sparse dorsal setae, distal margins rounded in male, each with 1 simple marginal seta and 1 marginal spine.

Eitmology. — Named forTuscarora Bank, the type locality.

Remarks. — Socarnes tuscarora sp. nov. differs from S. rurutu in having a slightly obtuse palm on

gnathopod 2, and a subquadrate posteroventral comer on the basis of peraeopod 7. Socarnes tuscarora differs from

S. tiendi in having an incised apicomedial margin on the outer plate of the maxilliped and minutely subchelate

gnathopod 2.

Distribution. — Socarnes tuscarora is known from Tuscarora Bank. South Pacific Ocean and the Kei Islands,

Banda Sea, in 170 to 540 m depth.

Genus SOCARNOPSIS Chevreux, 1911

Socarnopsis Chevreux. 1911 : 164. — Chevreux & FaGE, 1925 : 48. — J.L. Barnard, 1969 : 363. — Barnard &

Karaman, 1991 : 532.

Diagnosis. — Epistome and upper lip equally produced. Mandible : lacinia mobilis absent; molar setose with

well developed triturating surface. Maxilla 1 : ST7 asymmetrical; palp with terminal spines. Gnathopod 1 :

slightly subchelate.

Type Species. — Socarnopsis crenulata Chevreux, 191 1 (= Anonyx ftlicornis Heller, 1866), by monotypy.

Species Composition. — Socarnopsis contains 7 species ; S. allecta (Andres, 1981); S. dissimulantia

(Imbach, 1967); S. erythrophthalma (Robertson, 1892); S. ftlicornis (Heller, 1866); S. honiara sp. nov.; S. obesa

Chevreux. 1919; S. tandai sp. nov.

Remarks. — Socarnopsis filicornis (Heller, 1866) now includes S. schmardae (Heller, 1866) and S. crenulata

Chevreux, 1911, according to Krapp-Schickel, 1974, and Diviacco, 1984.

WALKER (1904) reported one specimen of S. filicornis (as Socarnes schmardae) from Sri Lanka, northern

Indian Ocean. It seems unlikely that this is S. filicornis. Walker did not illustrate his material. The very brief

description is not sufficient to place it in a genus but does not preclude it from Socarnopsis.

K.H. Barnard (1916) and Griffiths (1974, 1975) have recorded S. crenulata from southern Africa. These

records may not be the species now known as S. filicornis (they are not included in Diviacco & Ruffo's 1989

summary of distribution) but do seem to be in the genus Socarnopsis.

Ledoyer (1986) recorded several lots of material (previously reported in Ledoyf.r, 1967, 1972, 1973 and

1979) as Socarnes obesa. However, the material of Ledoyer, 1967 (originally recorded as Orchomene sp.) has an

entire telson and is clearly not a species of Socarnopsis. The material of LEDOYER, 1972 (also originally recorded

as Orchomene sp.) is possibly a species of Socarnopsis but differs from S. obesa in having a broadly rounded

lateral cephalic lobe. The material Ledoyer, 1973 (the illustration of male in Ledoyer, 1986) is clearly a

Socarnopsis but differs from S. obesa in having a straight epitome, a peak on the anterior margin of gnathopod 1

ischium, a much broader gnathopod 1 carpus and a differently shaped gnathopod 2 propodus. The material of

Ledoyer, 1979 (the illustrations of female in Ledoyer, 1986) differs from 5. obesa in having a broadly rounded

lateral cephalic lobe and an almost chelate gnathopod 2. Although it has some characters of a Socarnopsis.

Source :

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

197

especially the epistome/upper lip complex and gnathopod 1 propodus, the text suggests that the maxilla 1 palp and

mandibular molar are more like those of Socarnes.

Distribution. — Mediterranean Sea; eastern North and South Atlantic Oceans; Red Sea; south-western North

Pacific Ocean, north-western South Pacific Ocean; south-western Indian Ocean, ? northern Indian Ocean;

immediate sublittoral to 1544 m depth.

Socarnopsis Honiara sp. nov.

Figs 42-43

MATERIAL EXAMINED. — Solomon Islands. Stn SI-7, Rove, Honiara, Guadalcanal. 9°25.2’S, 159°56'E, baited

trap on limestone reef with some live coral heads and sand patches, 3 m, R.T. SPRINGTHORPE, 28-29 September 1991 :

10 specimens (MNHN-Am 4790); 44 specimens (AM P42150). — Stn SI-8, Rove, Honiara, Guadalcanal, 9°25.2'S,

159°56'E, baited trap on limestone reef with some live coral heads and sand patches, 4 m, R.T. SPRINGTHORPE,

28-29 September 1991 : 137 specimens (AM P42151). — Stn SI-9, Rove, Honiara, Guadalcanal, 9°25.2'S, 159°56'E,

baited trap on limestone reef with some live coral heads and sand patches, 5 m, R.T. SPRINGTHORPE, 28-29 September

1991 : 1 2, 4.1 mm (AM P42152); 1 6. 2.5 mm (AM P42153); 180 specimens (AM P42154).

Types. — The female, 4.1 mm (AM P42152) is the holotype. The other specimens are paratypes.

Type Locality. — Rove, Honiara, Guadalcanal, Solomon Islands, 9°25.2'S, 159°56'E, 5 m.

Diagnosis. — Antenna 1 with midmedial spines on callynophore. Antenna 2 of male subequal in length to

antenna 1. Gnathopod 1: ischium with sharp peak on anterior margin. Epimeron 3: posteroventral comer narrowly

rounded. Telson deeply cleft, about 70%.

Description. — Based on holotype female, 4.1 mm (AM P42152); male paratype, 2.5 mm (AM P42153).

Head : exposed, deeper than long; lateral cephalic lobe large, broad, distally truncated; rostrum small; eyes

reniform, dark brown/black in alcohol, not enlarged in adult male. Antenna 1 : peduncular article 1 short, length

1 times breadth, without tooth on distomedial margin, posterodistal tooth or anterodistal projection; peduncular

article 2 short, 0.33 times article 1, without anterodistal projection; peduncular article 3 short. 0.2 times article 1;

accessory flagellum long, 0.5 times primary flagellum, 5-articulate, article 1 long, 1.8 times article 2, not

forming cap; flagellum 9-articulate (male 7), with strong 2-field callynophore with midmedial spines in female and

male, flagellum without flagellar spines, calceoli absent in female and male. Antenna 2 : subequal in length to

antenna 1 (same in male); peduncle without brush setae in female and male, weakly geniculate between peduncular

articles 3-4, article 3 short, 0.52 times article 4 (male weakly geniculate between peduncular articles 3-4, article 3

short, 0.47 times article 4), peduncular articles 4 and 5 not enlarged in male or female; flagellum well developed.

9-articulate (male 6), without thick setal brush, calceoli absent in female (1 present in adult male).

Mouthpart bundle : subquadrate. Epistome and upper lip : separate; epistome concave; upper lip slightly

produced, rounded. Mandible : incisors symmetrical, small, with strongly convex margins; laciniae mobilis

absent; accessory spine row without distal setal tuft, left row with 3, right wilh 4 short, robust, 'bushy spines,

without intermediate setae; molar setose with well developed triturating surface; mandibular palp attached

proximally; article 1 short, length 1.2 times breadth; article 2 slender, length 4.2 times breadth, 1.5 times

article 3, with 5 (male 3) submarginal posterodistal A2-setae. without D2-setae; article 3 falcate, long, length

4.3 times breadth, without proximal A3-setae, with 3 (male 2) proximal and 2 (male 2) distal D3-setae and

2 apical E3-setae. Maxilla 1 : inner plate narrow with 2 plumose apical setae, outer seta without denticulate row;

outer plate with 11 spine-teeth in 6/5 arrangement; outer row with ST1 to ST3 large, stout, weakly to

multicuspidate, ST4 large, stout, 4-cuspidate, ST5 large, stout, 4- to 5-cuspidate, ST6 large, very broad,

multicuspidate distomedially, left and right ST7 asymmetrical, slightly displaced from ST6, left large, broad,

5-cuspidate distally, right large, very broad, 7-cuspidate distomedially; inner row with STA large, very broad,

slightly displaced from STB-STD, multicuspidate along entire medial margin, STB-STD large, broad.

198

J. K. LOWRY & H. E. STODDART

mullicuspidate along eniire medial margin; palp large, 2-articulate, with 5 short terminal spines, without

subterminal setae, flag spine present on distolateral comer, distomedial margin serrate. Maxilla 2 : inner plate

narrow, outer plate broader, inner plate 1 times length outer plate. Maxilliped : inner plate large,

subrectangular.with 3 apical nodular spines, oblique setal row reduced with 6 plumose setae; outer plate small.

Socarnopsis Honiara sp. nov„ holotype female, 4.1 mm (AM P42152); paratype male, 2.5 mm (AM P42153);

Rove, Honiara, Guadalcanal, Solomon Islands. Scales represent 0.1 mm.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM TOE TROPICAL WESTERN SOUTH PACIFIC OCEAN

199

subovate, without subapical notch, without apical setae or spines, medial spines vestigial, submarginal setae

short, simple; palp large, 4-articulate; article 2 slender, length 2.1 times breadth, 1.4 times article-3; article 3

long, slender, length 2. 1 times breadth; dactylus well developed, with 2 subterminal setae, unguis present.

Peraeoniles : 1 to 7 dorsally smooth. Gnathopod 1 ; subchelate; coxa large, slightly shorter than coxa 2,

anterior margin concave, anteroventral comer produced, rounded, posterior margin slightly convex; basis long,

slender, length 3 times breadth, anterior margin smooth, with simple setae; ischium short, length 1 times breadth,

anterior margin with sharp peak; merus, posterior margin with patch of short setae and with a few simple setae;

carpus subrectangular, short, length 1.9 times breadth, longer than (1.1 times) propodus, with patch of very fine

setae near posterior margin; propodus large, subrectangular, length 2.1 times breadth, tapering distally, posterior

margin smooth, straight, with 2 spines, without denticulate patch near posterior margin, palm extremely acute,

margin straight, smooth, posterodistal corner with 1 medial and 1 lateral spines; dactylus complex, with large

subterminal tooth and row of at least 25 medial spines near anterior margin. Gnathopod 2 : minutely subchelate;

coxa large, subequal in size to coxa 3; ischium long, length 2.6 times breadth; carpus long, length 2.6 times

breadth, posterior margin broadly lobate; propodus subquadratc, short, length 1.4 times breadth, palm slightly

obtuse, with straight, serrate margin, posterodistal comer without spines; dactylus reaching corner of palm,

posterior margin serrate.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly, merus-carpus without plumose setae; propodus

with 4 setae and 2 distal spines along posterior margin; dactylus short, slender. P eraeopod 4 : coxa deeper than

wide, with large posteroventral lobe, anterior margin slightly rounded, posterior margin slightly sloping

anteriorly; merus weakly expanded anteriorly, merus-carpus without plumose setae; propodus with 4 setae and

2 distal spines along posterior margin; dactylus short, slender. Peraeopod 5 : coxa equilobate, very large; basis

expanded with posterior margin scalloped, anterior margin with small glands at base of distal setae; merus

expanded with rounded posterior margin; propodus with 4 spines and 2 distal spines along anterior margin;

dactylus short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis expanded posteriorly with smooth

posterior margin, without anteroventral lobe; merus slightly expanded posteriorly; propodus with 1 seta and

2 distal spines along anterior margin; dactylus short, slender. Peraeopod 7 : basis expanded posteriorly, anterior

margin with small glands at base of proximal setae, posterior margin slightly rounded, minutely crenate.

posteroventral corner rounded, posteroventral margin rounded: merus slightly expanded, convex posterior margin

with 3 short spines; propodus with 2 distal spines on anterior margin and 3 setae on posterior margin; dactylus

short, slender.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 6, strongly pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 3 : posteroventral comer narrowly rounded. Urosomites : dorsally

smooth. Uropod I : without fine setae; peduncle with 5 dorsolateral. 1 apicolateral, 2 dorsomedial and

1 apicomedial spines; rami subequal in length, outer ramus with 3 dorsal spines; inner ramus with 2 dorsal

spines. Uropod 2 : without fine setae; peduncle without dorsolateral flange, with 1 dorsolateral. 1 apicolateral and

1 apicomedial spines, without plumose setae, without spines along distal margin; outer ramus slightly shorter

than inner ramus, outer ramus with 2 dorsal spines; inner ramus with 2 dorsal spines, without constriction.

Uropod 3 : peduncle well developed, long, length 2 times breadth, without dorsolateral flange, with 1 dorsolateral

and 1 apicolateral spines, without midlateral spines or setae, without distovcntral spines or plumose setae; rami

lanceolate, subequal in length, outer ramus 2-articulate, article 2 short, rami without spines, plumose setae absent

in female and male. Telson : longer than broad, length 1.9 times breadth, deeply cleft (72%), without dorsal

spines, with sparse dorsal setae, distal margins incised slightly, without marginal penicillate setae, with

1-2 simple marginal setae on each lobe, without marginal spines.

Etymology. — Named for the town of Honiara, near the type locality.

Remarks. — This is the second report of midmedial spines on the callynophore of a lysianassoid. The other

occurrence, in Aristias uokonia, is in a genus not considered to be closely related to Socarnopsts. which indicates

that the spines are independently derived. The spines in 5. honiara are not as robust as those ol A. uokonia.

Socarnopsts honiara and S. tandai apparently differ from all other species in the genus in having midmedial spines

on the callynophore.

200

J. K. LOWRY & H. E. STODDART

Source : MNHN. Pahs

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

201

Socarnopsis honiara is one of two species in the genus in which the male second antenna does not become

elongate. The other is S. erythrophthalma. from which S. honiara differs in having dark brown eyes, a well-

developed callynophore in both female and male and a more deeply cleft telson.

Distribution. — Socarnopsis honiara is known from Guadalcanal, Solomon Islands, in 3 to 5 m depth.

Socarnopsis tandai sp. nov.

Figs 44-45

MATERIAL EXAMINED. — Solomon Islands. Stn SI-11, Rove, Honiara, Guadalcanal, 9°25.2'S, 159°56'E, baited

trap near small coral head on sand bottom, 25 m, R.T. Springthorpe, 29-30 September 1991 : 1 9 , 4.8 mm (AM

P42252); 1 6, 4.2 mm (AM P42253); 80 specimens (AM P42155); 20 specimens (MNHN-Am 4795). — Stn SI-12,

Rove, Honiara, Guadalcanal, 9°25.2'S, 159°56'E, baited trap near small coral head, algal beds on sandy slope, 20 m,

R.T. SPRINGTHORPE, 29-30 September 1991 : 22 specimens (AM P42156). — Stn SI-13, Rove, Honiara, Guadalcanal,

9°25.2'S, 159°56’E, baited trap near small coral head on sandy slope, 16 m, R.T. SPRINGTHORPE, 29-30 September 1991 :

9 specimens (AM P42157). — Stn SI-19, 300 m north-west of Tandai Point, Guadalcanal, 9°23’S, 159°52.5'E, baited trap

on sand/rubble bottom under overhang near bed of coralline algae, 24 m, R.T. SPRINGTHORPE, 6-7 October 1991 :

3 specimens (AM P42158). — Stn SI-20, 300 m north-west of Tandai Point, Guadalcanal, 9°23'S, 159°52.5'E, baited

trap near small coral head, 18 m, R.T. Springthorpe, 6-7 October 1991 : 6 specimens (AM P42159).

Types. — The female. 4.8 mm (AM P42252) is the holotype. The other specimens are paratypes.

Type Locality. — Rove, Honiara, Guadalcanal, Solomon Islands, 9°25.2'S, 159°56’E, 25 m.

Diagnosis. — Antenna 1 with midmedial spines on callynophore. Antenna 2 of male elongate. Gnathopod 1:

ischium with sharp peak on anterior margin. Epimeron 3: posteroventral comer broadly rounded. Telson deeply

cleft, about 70%.

Description. — Based on holotype female, 4.8 mm (AM P42252); paratype male, 4.2 mm (AM P42253).

Head : exposed, deeper than long, lateral cephalic lobe large, broad, distally truncated; rostrum small; eyes

reniform, colour faded red in alcohol, not enlarged in adult male. Antenna I : peduncular article 1 short, length

1 times breadth; peduncular article 2 short, 0.36 times article 1, without anterodistal projection; peduncular article

3 short, 0.2 times article 1; accessory flagellum medium length, 0.36 times primary flagellum, 5-articulate,

article 1 long, 1.9 times article 2 (male long, 2 times article 2), not forming cap; flagellum 14-articulate

(male 14), with strong 2-field callynophore with midmedial spines in female and male, without flagellar spines,

calceoli absent in female and male. Antenna 2 ; subequal in length to antenna 1 (longer than body in male),

peduncle without brush setae in female and male, weakly geniculate between peduncular articles 3-4, article 3

short, 0.42 times article 4 (male weakly geniculate between peduncular articles 3-4, article 3 short, 0.6 times

article 4), peduncular article 4 enlarged in male; flagellum well developed. 12-articulate (male 51), calceoli absent

in female (6 present in adult male on articles 38, 40, 42, 44, 46, 48).

Mouthpart bundle : subquadrate. Epistome and upper lip : separate, epistome concave, upper lip slightly

produced, rounded. Mandible : incisors symmetrical, small, with strongly convex margins; lacinia mobilis absent;

accessory spine row without distal setal tuft, left row with 3. right row with 4 short, robust, multiscrrate setae;

intermediate setae absent; molar setose with well developed triturating surface; mandibular palp attached

proximally; article 1 short, length 1.5 times breadth; article 2 slender, length 5.9 times breadth. 1.5 times

article 3, with 5 submarginal posterodistal A2-setae (male 5), without D2-setae; article 3 lalcate, long, length

5.3 times breadth, without A3-sctae. with 4 proximal D3-sctae (male 5), with 2 distal D3-setae (male 2) and

2 apical E3-setae. Maxilla 1 : inner plate narrow, with 2 plumose apical setae, outer seta without denticulate row:

outer plate with 1 1 spine-teeth in 6/5 arrangement, outer row with ST1 to ST3 large, stout, weakly to

multicuspidate. ST4 large, stout, 4-cuspidate. ST5 large, stout, 5-cuspidate. ST6 large, very broad. 8-cuspidate

distomedially, left and right ST7 asymmetrical, slightly displaced from ST6, left large, broad. 5-cuspidate distally,

right very broad, 9-cuspidate distomedially; inner row with STA large, very broad, slightly displaced from STB.

202

J. K. LOWRY & H. E. STODDART

tZTriS*Tda\ S|,',n0V” hT°!0lyPC female- 4-8 mm- AM P42252; Para‘yPe male, 4.2 mm (AM P42253);

Hove, Honiara, Guadalcanal, Solomon Islands. Scales represent 0.1 mm.

Source : MNHN . Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

203

multicuspidate along entire medial margin. STB-STD large, broad, mullicuspidate along entire medial margin;

palp large, 2-articulate, with 7 short terminal spines, without subterminal setae, flag spine present on distolateral

corner, distomedial margin serrate. Maxilla 2 : inner plate narrow, outer plate broader, inner plate 0.8 limes length

outer plate. Maxilliped : inner plate large, suhrectangular, with 3 apical nodular spines, oblique setal row reduced

with 7 plumose setae; outer plate small, subovate, without subapical notch, without apical setae or spines, medial

setae vestigial, submarginal setae short, simple; palp large. 4-articulate, article 2 broad, length 2.4 times breadth.

1.6 times article 3, article 3 long, slender, length 2.4 times breadth, dactylus well developed, with 2 subterminal

setae, unguis present.

Peraeonites 1 to 7 dorsally smooth. Gnathopod l : weakly subchelate; coxa large, slightly shorter than coxa 2.

anterior margin concave, anteroventral comer produced, rounded, posterior margin slightly convex; basis long,

slender, length 4.3 times breadth, anterior margin smooth, with simple setae; ischium short, length 1.2 times

breadth, anterior margin with sharp peak; merus, posterior margin with patch of short setae and a few simple setae;

carpus subrectangular, long, length 2 times breadth, longer than (1.1 limes) propodus, with patch of very fine

setae near posterior margin; propodus large, subrectangular. length 2.3 times breadth, tapering distally, posterior

margin smooth, straight, with 2 spines, palm extremely acute, margin straight, smooth, posterodistal corner with

1 medial and 1 lateral spine; dactylus complex, with large subterminal tooth and row of 28 medial spines near

anterior margin. Gnathopod 2 : minutely subchelate; coxa large, subcqual in size to coxa 3; ischium long, length

2.8 times breadth; carpus long, length 3 times breadth, posterior margin broadly lobate; propodus subquadrate,

short, length 1.5 times breadth, palm slightly obtuse, with straight, serrate margin, posterodistal corner without

spines; dactylus not reaching corner of palm, posterior margin serrate.

Peraeopod 3 : coxa large; merus expanded anterodistally along carpus; merus-carpus without plumose setae in

male and female; propodus with 5 slender setae and 2 distal spines along posterior margin; dactylus short, slender.

Peraeopod 4 : coxa deeper than wide, with large posteroventral lobe, anterior margin slightly rounded, posterior

margin slightly sloping anteriorly; merus expanded anterodistally along carpus; merus-carpus without plumose

setae in male and female; without posterodistal spur; propodus with 4 slender setae and 2 distal spines along

posterior margin; dactylus short, slender. Peraeopod 5 : coxa equilobate (very large): basis expanded with posterior

margin minutely crenate; merus expanded with rounded posterior margin; propodus with 4 slender setae along

anterior margin and 2 distal spines; dactylus short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis,

anterior margin rounded, basis expanded posteriorly with minutely crenate posterior margin; merus slightly

expanded and rounded posteroproximally, straight posterodistally with 2 setae; propodus with 7 spines long

anterior margin; dactylus short, slender. Peraeopod 7 : basis expanded posteriorly, anterior margin with small

glands at the base of proximal setae, posterior margin slightly rounded, minutely crenate, posteroventral corner

rounded, posteroventral margin rounded: merus slightly expanded, convex posterior margin with 3 spines;

propodus with 7 spines along anterior margin and 9 slender setae along posterior margin; dactylus short, slender.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 6. with strong horizontal

pleating.

Pleonites I to 3 dorsally smooth. Epimeron l : anteroventral corner rounded. Epimeron 3 : posteroventral cor¬

ner broadly rounded. Urosomites ; dorsally smooth. Uropod I : with long fine setae; peduncle with 5 dorsolateral.

1 apicolateral, 2 dorsomedial and 1 apicomcdial spine; rami subequal in length; outer ramus with 4 dorsal spines,

inner ramus with 2 dorsal spines. Uropod 2 : without fine setae; peduncle without dorsolateral flange, with

1 dorsolateral, 1 apicolateral and 1 apicomcdial spine, without plumose setae, without spines along distal margin;

outer ramus slightly shorter than inner ramus; outer ramus with 3 dorsal spines, inner ramus with 3 dorsal spines;

inner ramus without constriction. Uropod 3 : with long fine setae: peduncle long, length 2.2 times breadth, with¬

out dorsolateral flange, with 1 dorsolateral and 1 apicolateral spine, without midlateral setae or spines, without dis-

toventral spines, without plumose setae; rami lanceolate, subequal in length, outer ramus

2-articulate, article 2 short, rami without spines, slender plumose setae absent in female and male. Telson : longer

than broad, length 2.2 times breadth, deeply cleft (68%), without dorsal spines, with sparse dorsal setae, distal

margins incised slightly, with 1 marginal penicillate seta on each lobe, with 1-2 marginal simple setae on each

lobe, without marginal spines.

Etymology. — Named for Tandai Point, one of the localities at which the species is found.

204

J. K. LOWRY & H. E. STODDART

F'CuL7, S£?n°PSiS 'T/n'oSP- n°V" h0l°‘yPe femalC' 4 8 mm (AM P42252)- Rove> Honiara- Guadalcanal. Solomor

islands. Scales represent 0.2 mm.

Source : MNHN, Paris

LYSIANASSOID AMPIIIPODA FROM TOE TROPICAL WESTERN SOUTH PACIFIC OCEAN

205

Remarks. — Socarnopsis tandai is easily distinguished from S. allecta, S. dissimulantia and S. obesa by the

sharp peak on the anterior margin of the gnathopod 1 ischium. It is very similar to S. filicornis , but differs in

having a slightly more concave epistome, a smaller triturating area on the molar, a broader article 2 of the

maxillipedal palp, an anterodistal bulge on the propodus of gnathopod 2 and a slightly rounder posteroventral

comer on epimeron 3.

Socarnopsis tandai is extremely similar to S. honiara. There are small differences in the relative length of the

flagellum and accessory flagellum of antenna 1. the breadth of articles 2 and 3 of the maxillipedal palp and the

taper of the head lobe. The two species can be separated by differences in adult size, eye colour and the adult male

antenna 2 which is elongate with an enlarged peduncular article 4 in S. tandai but subequal to that of the female in

S. honiara. Lincoln (1979) recognized the same close relationship, including the male antenna 2, in the North

Atlantic species in S. filicornis (as S. crenulatus) and S. erythrophlhalma.

Although we have only a small number of samples, from a limited area, it seems that the two species of

Socarnopsis from the Solomon Islands occur in different habitats. Socarnopsis honiara was taken only in traps set

on limestone reefs, in 3 to 5 metres depth: S. tandai was taken in traps set on sandy bottoms in 18 to 25 metres

depth.

DISTRIBUTION. — Socarnopsis tandai is known from Guadalcanal, Solomon Islands, in 18 to 25 m depth.

Genus STEPHONYX Lowry & Stoddart. 1989

Stephonyx sp.

MATERIAL EXAMINED. — New Caledonia. ClIALCAL 2 : stn DW 76, 23°40.50'S, 167°45.20’E, south of the Isle of

Pines, 470 m, 30 October 1986 : 1 specimen (MNHN-Am 4428).

REMARKS. — Lowry & STODDART arc preparing a monograph on the genus Stephonyx. This species will be

described in that report.

Genus TRISCH1ZOSTOMA Boeck. 1861

Trischizostoma richeri sp. nov.

Figs 46-48

Material EXAMINED. — Loyalty Islands. B too FOCAL : stn CP 321. 21°12.0'S. 166°59.85'E. Loyalty Islands

Basin. 2190-2205 m, 3 May 1987 : 1 9, 41 mm, ovigerous, 8 eggs (MNHN-Am 4458).

Types. — The unique specimen is the holotype.

Type Locality. — Loyalty Islands Basin, 21°12.0'S. 166°59.85'E, 2190 to 2205 m.

Diagnosis. — Maxillipedal palp 4-articulate. styliform. longer than outer plate. Gnathopod 1 : propodus oval.

0.45 times as long as broad. Telson entire, distally subacute.

Description. — Based on holotype female, 41 mm; male not known. Head : exposed, deeper than long; lateral

cephalic lobe absent; rostrum large; eyes covering most of head, expanded dorsally and nearly confluent.

Antenna 1 : short. 0.16 times body; peduncular article 1 short, length 0.9 times breadth, without tooth on

distomedial margin; peduncular article 2 short. 0.2 times article 1. without anterodistal projection; peduncular

article 3 short, 0.2 times article 1: accessory flagellum medium length. 0.4 times primary flagellum, at leas!

4-articulate. article 1 long. 8.8 times article 2. forming cap partially covering callynophore; flagellum

1 1-articulate, with strong 2-field callynophore. with 1 long spine on article 3, calceoli absent in female.

206

J. K. LOWRY & H. E. STODDART

Antenna 2 : length 1.6 times antenna 1; peduncle with weak brush setae, peduncular article 1 greatly enlarged, not

covering article 2. weakly geniculate between peduncular articles 3-4. article 3 short, 0.32 times article 4;

flagellum well developed. 29-articulate, without thick setal brush, calceoli absent in female.

Fig. 46. Trischizostoma richeri sp. nov., holotype female, 41 mm (MNHN-Am 4458), Loyalty Islands Basin.

Mouthpart bundle : conical. Epistome and upper lip : fused, sinusoidal. Mandible : incisors symmetrical, very

small at tip of styliform projection; laciniae mobilis absent; accessory spine row absent; molar absent; mandibular

palp attached proximally; article 1 short, length 0.6 times breadth; article 2 broad, length 2.8 times breadth.

1.1 times article 3, with many A2-setae. with about 25 D2-setae on distal third of posterior margin; article 3

falcate, long, length 2.9 times breadth, without proximal A3-setae, with 19 D3-setac along most of posterior

margin and 1 apical E3-seta. Maxilla 7 ; inner plate narrow with 1 simple apical seta; outer plate narrow with

8 spine-teeth in modified 8/3 crown arrangement; outer row with ST1 to ST3 large, stout, without cusps. ST4

large, stout, smooth. ST5-ST7 absent; inner row with STA large, stout, displaced from STB-STD, without cusps.

STB-STD short, slender, without cusps; palp small, 1 -articulate, with 2 apical setae, without subterminal setae,

flag spine absent, distomedial margin smooth. Maxilla 2 : inner and outer plates narrow, inner plate 1 times

length outer plate. Maxilliped : inner plate very large, substyliform, with 2 apical vestigial nodular spines; outer

plate small, subovate, without subapical notch, apical setae, apical spines or medial spines, submarginal setae

vestigial; palp large, 4-articulate. styliform, geniculate between articles 2 and 3; article 2 broad, length 2 times

breadth. 1 times article 3; article 3 long, broad, length 1.9 times breadth; dactylus longest of all, slender,

lanceolate with smooth anterior margin, with 2 terminal setae, unguis absent.

Peraeonites : 1 to 7 dorsally smooth. Gnathopod l : subchelate; coxa vestigial; basis long, slender, length

4.9 times breadth, anterior margin smooth, without setae; ischium short, length 1.5 times breadth; merus and

carpus rotated, propodus and dactylus inverted in adult, posterior margin without setae; carpus subtriangular,

compressed, shorter than propodus, without denticulate patch near posterodistal margin; propodus massive,

subovate, length 0.45 times breadth, margins diverging distally. posterior margin smooth, convex, without spines

or setae, without denticulate patch near posterior margin, palm slightly acute, margin convex, lined with row of

short, thick spines, posterodistal corner with 3 medial and 2 lateral spines; dactylus simple, without subterminal

teeth or spines. Gnathopod 2 : minutely subchelate; coxa large, larger than coxa 3, anteroventral comer rounded,

ventral margin straight; ischium long, length 3.75 times breadth; carpus long, length 3.6 times breadth, posterior

margin straight; propodus produced anterodistally beyond dactylus, short, length 1.7 times breadth, palm slightly

acute, with straight, smooth margin, without spines; dactylus, posterior margin smooth.

Peraeopod 3 : coxa large; merus weakly expanded anteriorly; propodus with 3 setae along posterior margin;

dactylus short, slender. Peraeopod 4 : coxa deeper than wide, with weak posteroventral lobe, anterior margin

I.YSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

207

EP3

. _ /

Fig. 47. — Trischizostoma richcri sp. nov., holotype female, 41 mm (MNHN-Am 4458), Loyalty Islands Basin.

Scales represent 0.5 mm.

Source MNHN . Paris

208

J. K. LOWRY & H. E. STODDART

■ 48. — Trischizostoma richeri

Scales represent 1.0 mm.

holotype female, 41 mm (MNHN-Am 4458), Loyalty Islands Basin

nov

Source : MNHN. Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

209

broadly rounded, posterior margin slightly sloping anteriorly; merus expanded anteriorly and posteriorly; propodus

with 3 setae along posterior margin; dactylus short, slender. Peraeopod 5 : coxa equilobate; basis expanded with

posterior margin smooth; merus expanded with rounded posterior margin; propodus with 3 setae along anterior

margin; dactylus short, slender. Peraeopod 6 : coxa small, not lobate posteriorly; basis expanded with broad

posterovcntral lobe, without antcroventral lobe; merus expanded proximally, posterior margin straight, converging

distally, with 4 spines; propodus with 3 setae along anterior margin; dactylus short, slender. Peraeopod 7 : basis

expanded posteriorly, posterior margin slightly rounded, minutely crenate, posteroventral margin rounded; merus

expanded proximally, posterior margin straight, converging distally, with 6 spines; propodus with 6 setae along

anterior margin and 2 setae along posterior margin; dactylus short, slender.

Oosiegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 7, with strong horizontal

pleating.

Pleonites 1 to 3 dorsally smooth. Epimeron 1 : anteroventral corner broadly rounded. Epimeron 3 :

posteroventral corner subquadrate. Urosomites : urosomite 1 with anterodorsal notch. Uropod 1 : peduncle with

9 dorsomedial and 1 apicomcdial spines; outer ramus slightly shorter than inner ramus, outer ramus with 6 lateral

spines; inner ramus with 2 lateral spines. Uropod 2 : peduncle without dorsolateral flange, with 1 apicolatcral

spine, without spines along distal margin; outer ramus slightly shorter than inner ramus, rami without spines;

inner ramus without constriction. Uropod 3 : peduncle well developed, short, length 0.92 times breadth, without

dorsolateral flange, without dorsal spines, midlateral spines or setae or distoventral spines; biramous, rami

lanceolate, subequal in length, outer ramus 2-articulate, article 2 short, rami without spines. Telson : longer than

broad, length 1.1 times breadth, entire, without dorsal setae, margins converging distally. apically subacute,

without marginal penicillate or simple setae, without marginal spines.

ETYMOLOGY. — Named for Bertrand Richer de Forges, carcinologist, and architect of the ORSTOM

collecting program in the tropical western Pacific Ocean.

Remarks. — Trischizosloma richeri is a distinctive species which appears most similar to 7'. longirostre

Chevreux, 1919, and T. macrochela Vinogradov. 1990. In these three species the telson is entire, the maxillipedal

palp is much longer than the outer plate, the propodus of gnathopod 1 is oval and much deeper than long and the

palm is smooth. Trischizosloma richeri has a shorter rostrum than docs T. longirostre (described more fully by

Chevrf.ux, 1927). the dorsodistal protuberance on the propodus of gnathopod 2 is stronger and the telson is

subacute distally, not truncated. Although T. richeri and T. macrochela share the above characters they are very

different. In T. macrochela the mandibular palp is weakly setose, the palp of maxilla 1 is completely absent, the

maxillipedal palp, although much longer than the outer plate, is weakly developed and not styliform. the palm of

gnathopod 1 is weakly spinose. the peracopods are slender and the telson is rounded. Trischizosloma richeri differs

from the other species in the Indo-west Pacific, T. crosnieri Lowry & Stoddart, 1993, as follows : the

antcroventral comer of coxa 2 is rounded; the merus is not as well developed in peracopods 3 and 4; the basis ot

peraeopod 5 has a rounded posteroventral corner and the telson is entire and subtriangular.

Species of Trischizosloma are known as ectoparasites on fish. The host species for T. richeri is not known.

DISTRIBUTION. — 'Trischizosloma richeri is currently known from the Loyalty Islands Basin in 2200 m depth.

Genus TRYPHOSELLA Bonnier. 1893

Tryphosella ama sp. nov.

Figs 49-51

MATERIAL EXAMINED. — New Caledonia. CAI.SUB : sin PL 20, 22°52.7'S, 167°23'E, south of the Isle of Pines,

555-616 m, 10 March 1989 : 1 female, 9.5 mm. ovigerous (MNHN-Am 4791); 5 specimens (MNHN-Am 4791); 2

specimens (AM P42160).

Types. — The female, 9.5 mm (MNHN-AM 4792), is the holotype. The other specimens are paratypes.

210

J. K. LOWRY & H. E. STODDART

Type Locality. — Soulh of Ihe Isle of Pines, New Caledonia, 22°52.7'S, 167°23'E, 555 to 616 m.

Diagnosis. — Lateral cephalic lobe large, broad, down-turned. Eyes apparently absent. Epistome : slightly

produced, rounded. Maxillipcd : outer plate with 3 vestigial apical spines. Gnathopod 1 : carpus slightly longer

than propodus; palm acute. Epimeron 3 : posteroventral corner narrowly rounded. Urosomite 1 rounded, with

rounded boss. Uropod 2 : inner ramus with moderate constriction.

Fig. 49. — Tryphosella ama sp. nov., holotype female. 9.5 mm (MNHN-Am 4791), south of the Isle of Pines,

New Caledonia.

Description. — Based on holotype female, 9.5 mm; male not known. Head and body : without setae. Head :

exposed, deeper than long; lateral cephalic lobe large, broad, down-turned, distally rounded; rostrum absent; eyes

apparently absent. Antenna I : medium length. 0.2 times body; peduncular article 1 short, length 1.3 times

breadth, without tooth on distomedial margin, posterodistal tooth or anterodistal projection; peduncular article 2

short, 0.22 times article 1. without anterodistal projection; peduncular article 3 short, 0.15 times article 1:

accessory flagellum long, 0.51 times primary flagellum. 9-articulate, article 1 long, 3.1 times article 2, not

forming cap; flagellum 16-articulate, with strong 2-field callynophore, with 2 small posterodistal setae, without

flagellar spines, calceoli absent. Antenna 2 : subcqual in length to antenna 1; peduncle without brush setae,

weakly geniculate between peduncular articles 3-4. article 3 short, 0.42 times article 4, peduncular articles 4 and 5

not enlarged; flagellum well developed. 16-articulate, calceoli absent.

Mouthpart bundle : subquadrate. Epistome and upper lip : separate; epistome slightly produced, rounded; upper

lip slightly produced, rounded. Mandible : incisors symmetrical, large, with strongly convex margins; left lacinia

mobilis present, a stemmed distally serrate blade; accessory spine row without distal setal tuft, left row with 4.

right with 3 short, slender, "bushy" spines, with 3 "whip-like" intermediate setae; molar proximally setose,

distally triturating; mandibular palp attached midway; article 1 short, length 1.2 times breadth; article 2 slender,

length 6.1 times breadth, 2.3 times article 3. with 16 submarginal posterodistal A2-setae, without B2-setae or

D2-setae; article 3 talcate, long, length 3.4 times breadth, with 1 proximal A3-seta, without B3-setac, with

16 D3-setae along most of posterior margin and 2 apical E3-setae. Maxilla 1 : inner plate narrow with 2 simple

apical setae, outer seta with denticulate row; outer plate with 1 1 spine-teeth in 6/5 arrangement; outer row with

ST1 large, stout, multicuspidate. ST2 and ST3 weakly cuspidate. ST4 large, stout. 4-cuspidatc, ST5 large, stout,

6-cuspidate, ST6 large, stout, multicuspidate, ST7 slightly displaced from ST6, large, broad, with convex

multicuspidate medial margin; inner row with STA large, very broad, slightly displaced from STB-STD,

multicuspidate along entire medial margin, STB-STD large, broad, multicuspidate along entire medial margin;

palp large. 2-articulate, with 7 short terminal spines, with 1-2 subterminal setae, flag spine present on distolateral

corner, distomedial margin smooth. Maxilla 2 : inner and outer plates narrow, inner plate 0.8 times length outer

plate. Maxilliped : inner plate large, subrectangular, with 3 apical nodular spines, with 2 distal spines on lateral

Source :

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

211

FlG. 50. — Trypliosella ama sp. nov.. holotype female, 9.5 mm (MNHN-Am 4791). south of ihe Isle of Pines.

New Caledonia. Scales represent 0.2 mm.

Source : MNHN, Paris

212

J. K. I.OWRY & H. fi. STODDART

face near inner margin, oblique seial row strong with 17 plumose setae; outer plate medium size, subovate,

without subapical notch or apical setae, with 3 vestigial apical spines, medial spines present, small, submarginal

setae short, simple; palp large. 4-articulate; article 2 slender, length 3.2 times breadth, 1.6 times article 3; article 3

long, slender, length 2.3 times breadth; dactylus well developed, with 3 subterminal setae, unguis present.

Peraeonites 1 to 7 dorsally smooth. Gnathopod I : subchelate; coxa large, slightly shorter than coxa 2.

tapering distally. anterior margin concave, anteroventral corner produced, rounded, posterior margin distally angled

F,V*. Tryphosella ama sp. nov., holotype female, 9.5 mm (MNHN-Am 4791),

New Caledonia. Scales represent 0.5 mm.

south of the Isle of Pines.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

213

towards anterior margin; basis long, slender, length 4.9 times breadth, anterior margin smooth, with simple setae;

ischium long, length 3 times breadth, anterior margin smooth; merus. posterior margin with patch of short setae;

carpus subrectangular. long, length 3.3 times breadth, longer than (1.2 times) propodus, with patch of very fine

setae near posterior margin; propodus large, subrectangular, length 2.7 times breadth, margins subparallel,

posterior margin smooth, straight, with setae, with very fine setae near posterior margin, palm acute, margin

straight, serrate, posterodistal corner with 1 medial and 1 lateral spines; dactylus complex, with large subterminal

tooth and row of many medial spines near anterior margin. Gnathopod 2 : minutely subchelate; coxa large,

subcqual in size to coxa 3; ischium long, length 2.9 times breadth; carpus long, length 3.3 times breadth,

posterior margin broadly lobate; propodus subquadrate, short, length 1.6 times breadth, posterior margin without

strong distal spines, palm transverse, with straight, serrate margin, posterodistal corner with 1 medial spine;

dactylus reaching comer of palm, posterior margin serrate.

Peraeopod 3 : coxa large; merus, propodus and dactylus unknown. Peraeopod 4 : coxa deeper than wide, with

large posterovcntral lobe, anterior margin slightly rounded, posterior margin slightly sloping anteriorly; merus not

expanded anteriorly, merus-carpus without plumose setae; propodus with 7 spines and 2 distal spines along

posterior margin; dactylus short, slender. Peraeopod 5 : coxa bilobatc, posterior lobe slightly produced ventrally;

basis expanded with posterior margin minutely crenatc; merus slightly expanded posteriorly; propodus with

10 spines and 2 distal spines along anterior margin; dactylus short, slender. Peraeopod 6 : coxa small, not lobate

posteriorly; basis expanded posteriorly with minutely crenatc posterior margin, without antcroventral lobe; merus

slightly expanded and rounded posteroproximally, straight posterodistally with 3 setae; propodus with 1 1 spines

and 2 distal spines along anterior margin; dactylus short, slender. Peraeopod 7 : basis expanded posteriorly,

posterior margin slightly rounded, minutely crenatc, posterovcntral comer rounded, posterovcntral margin rounded;

merus anterior and posterior margins subparallel; propodus with 8 spines and 2 distal spines along anterior margin

and 2 distal spines along posterior margin; dactylus short, slender.

Oostegites : from gnathopod 2 to peraeopod 5. Gills : from gnathopod 2 to peraeopod 7. not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 1 : anteroventral corner narrowly rou-ded. Epimeron 3 :

posterovcntral corner narrowly rounded. Urosomites : urosomite 1 with rounded boss, without lateral flange,

urosomite 3 without small dorsolateral spine. Uropod 1 : without fine setae; peduncle with 6 dorsolateral.

1 apicolatcral, 6 dorsomedial and 1 apicomedial spines; rami subequal in length, outer ramus with 6 dorsal spines;

inner ramus with 6 dorsal spines. Uropod 2 : without fine setae; peduncle without dorsolateral flange, with

8 dorsolateral, 1 apicolatcral, 3 dorsomedial and 1 apicomedial spines, without plumose setae, without spines

along distal margin; rami subequal in length, outer ramus with 7 dorsal spines; inner ramus with 4 dorsal spines.

with° moderate constriction. Uropod 3 : peduncle well developed, short, length 1.6 times breadth, without

dorsolateral flange, with 2 apicolateral and 1 dorsal spines, without midlateral spines or setae, with 3 distoventral

spines with plumose setae; biramous. rami lanceolate, inner ramus reduced, about 0.8 times outer ramus, outer

ramus 2-articulate. article 2 short, article 1 with 5 lateral and 1 medial spines; inner ramus with 4 medial and

2 lateral spines, plumose setae present in female. Tel son : longer than broad, length 1.5 times breadth, deeply

cleft (81%), with 2 dorsal spines on each lobe, with sparse dorsal setae, distal margins incised, with 1 marginal

penicillate setae on each lobe, without simple marginal setae, with 1 marginal spine on each lobe.

Etymology. — Named for Ama Island, the southern-most island in the Isle of Pines group.

REMARKS. — Because of the size of the genus the following comparison is restricted geographically.

Tryphosella ama is atypical of the genus in having a long ischium on gnathopod 1 and a constricted inner ramus

on uropod 2. We have other Indo-west Pacific species under study which also have these characters and we cannot

yet assess their importance at the generic level. Tryphosella ama and T. oupi both have narrowly rounded

posterovcntral corners on epimeron 3. Tryphosella ama differs from T. oupi in having a more slender antenna

with more articles in the accessory and primary flagella, a shorter article 3 on the mandibular palp, 3 or 4 vestigial

apical spines on the outer plate of the maxilliped and a more slender maxillipedal palp, a more acute palm on

gnathopod 1 and large dorsal spines on the telson.

Distribution. — Tryphosella ama is known only from southern New Caledonia, in 555 to 616 m depth.

214

J. K. LOWRY & II. E. STODDART

Tryphosella oupi sp. nov.

Figs 52-53

Material EXAMINED. —New Caledonia. Biocal : stn CP 75. 22°18.65'S, 167°23.30'E. north of the Isle of

Pines. 825-860 m, 4 September 1985 :1 c?, 5.5 mm (MNHN-Am 4381).

Types. — The unique specimen is the hololype.

Type Locality. — North of the Isle of Pines. New Caledonia. 22°18.65'S. 167°23.30'E, 825 to 860 m.

Diagnosis. — Lateral cephalic lobe subacute. Eyes apparently absent. Epistome : slightly produced, rounded.

Maxilliped : outer plate with 2 apical spines. Gnathopod 1 : carpus length 1 times propodus; palm slightly acute.

Epimeron 3 : posteroventral comer narrowly rounded. Urosomite 1 rounded, with anterodorsal notch.

Description. — Based on holotype male. 5.5 mm: female unknown. Head : exposed, deeper than long; lateral

cephalic lobe large, broad, subacute; rostrum absent; eyes apparently absent. Antenna 1 : peduncular article 1

short, length 1 times breadth, without tooth on distomedial margin, postcrodistal tooth or antcrodistal projection;

peduncular article 2 short. 0.19 times article 1. without anterodistal projection; peduncular article 3 long,

0.23 times article 1; accessory flagellum long, 0.52 times primary flagellum. 4-articulate, article 1 long.

3.1 times article 2. not forming cap; flagellum male 8-articulate, with strong 2-field callynophore without

posterodistal setae or spines, without flagellar spines, calceoli absent. Antenna 2 : subcqual in length to antenna 1;

peduncle without brush setae in male, weakly geniculate between peduncular articles 3-4. article 3 short,

0.49 times article 4. peduncular articles 4 and 5 not enlarged; flagellum well developed, 1 1 -articulate, calceoli

present.

Mouthpart bundle : subquadratc. Epistome and upper Up : separate: epistome slightly produced, rounded; upper

lip not produced, straight. Mandible : incisors symmetrical, large, with strongly convex margins; left lacinia

mobilis present, a stemmed distally serrate blade: accessory spine row without distal setal tuft, left and right rows

each with 3 long, slender, "bushy" spines, with 4 "whip-like" intermediate setae; molar proximally setose, distally

triturating; mandibular palp attached midway; article 1 short, length 1.1 times breadth: article 2 slender, length

4.2 times breadth. 1.3 times article 3. with 8 A2-setae, without B2-setae or D2 -setae; article 3 falcate, long,

length 3.9 times breadth, with 1 proximal A3-seta, without B3-setae. with 12 D3-setac along most of posterior

margin and 2 apical E3-setac. Maxilla 1 : inner plate narrow with 2 plumose apical setae, outer seta with

denticulate row; outer plate with 11 spine-teeth in 6/5 arrangement; outer row with ST1 to ST3 large, stout,

weakly cuspidate, ST4 large, stout. 3-cuspidatc. ST5 large, stout. 5-cuspidate, ST6 large, stout, multicuspidate!

ST 7 slightly displaced Irom ST6. large, broad, with convex multicuspidate medial margin; inner row with STA

large, very broad, slightly displaced from STB-STD, 6-cuspidate. STB large, broad, 6-cuspidatc. STC large, broad,

7 -cuspidate. STD large, broad, multicuspidate along medial margin; palp large, 2-articulate. with 4 long terminal

spines, with 1 subtcrminal seta, flag spine present on distolatcral comer, distomedial margin smooth. Maxilla 2 :

inner and outer plates narrow, inner plate 0.9 times length outer plate. Maxilliped : inner plate large,

subrectangular. with 3 apical nodular spines, with 2 distal spines on lateral face near inner margin, oblique setal

row strong with 9 plumose setae; outer plate small, subovate. without subapical notch or apical setae, with

2 apical spines, medial spines present, small, submarginal setae short, simple; palp large, 4-articulate; article 2

very broad, length 2 times breadth. 1.4 times article 3; article 3 short, broad, length 1.75 times breadth’; dactylus

well developed, with 3 subterminal setae, unguis present.

Peraeonites 1 to 7 dorsal ly smooth. Gnathopod 1 : subchelate; coxa large, slightly shorter than coxa 2,

tapering distally, anterior margin straight, posterior margin distally angled towards anterior margin; basis long,

slender, length 3.2 times breadth, anterior margin smooth, with simple setae; ischium short, length 1 times

breadth, anterior margin smooth; merus, posterior margin with patch of short setae; carpus subrectangular, short,

length 1.8 times breadth and 1 times propodus, without patch of very fine setae near posterior margin; propodus

without very fine setae near posterior margin: propodus large, subrectangular. length 2 times breadth, margins

subparallcl. posterior margin smooth, straight, with setae, without denticulate patch near posterior margin, palm

LYSIANASSOID AMPMIPODA FROM THE TROPICAL WESTERN SOUTH PACIFIC OCEAN

215

slightly acute, margin sinusoidal, serrate, posterodistal comer with 1 medial and 1 lateral spines; dactylus simple,

with subterminal tooth. Gnaihopod 2 : minutely subchelate; coxa large, subequal in size to coxa 3; ischium long,

length 2.7 times breadth; carpus long, length 3.7 times breadth, posterior margin straight; propodus

Fig. 52. — Tryphosella oupi sp. nov., holotypc male, 5.5 mm (MNHN-Am 4381), north of the Isle of Pines,

New Caledonia. Scales for Al, 2, H+E represent 0.2 mm, remainder represent 0.1 mm.

Source MNHN. Pans

216

J. K. LOWRY & H. E. STODDART

subrcctangular, short, length 1.7 times breadth, palm slightly obtuse, with straight, serrate margin, posterodistal

comer with 1 medial spine; dactylus reaching comer of palm, posterior margin smooth proximally with serrate tip.

Fig. 53. — Tryphosella oupi sp.

New Caledonia. Scales for U3, T

nov., holotype male, 5.5 mm (MNHN-Am 4381), north of the Isle of Pines,

represent 0.1 mm, remainder represent 0.2 mm.

Source : MNHN, Paris

LYSIANASSOID AMPHIPODA FROM T1IE TROPICAL WESTERN SOUTH PACIFIC OCEAN

217

Peraeopod 3 : coxa large; merus weakly expanded anteriorly, merus-carpus without plumose setae; propodus

with 3 spines and 2 distal spines along posterior margin; dactylus short, slender. Peraeopod 4 : coxa deeper than

wide, with large posteroventral lobe, anterior margin slightly rounded, posterior margin slightly sloping

anteriorly; merus weakly expanded anteriorly, merus-carpus without plumose setae; propodus with 3 spines and

2 distal spines along posterior margin; dactylus short, slender. Peraeopod 5 : coxa equilobate; basis expanded with

posterior margin minutely crenate; merus slightly expanded posteriorly; propodus and dactylus unknown.

Peraeopod 6 : coxa small, not lobate posteriorly; basis expanded posteriorly with minutely crenate posterior

margin, without antcroventral lobe; merus slightly expanded posteriorly with 3 spines; propodus and dactylus

unknown. Peraeopod 7 : basis expanded posteriorly, posterior margin slightly rounded, minutely crenate.

posteroventral corner rounded, posteroventral margin rounded; merus slightly expanded posterodistally with

2 spines; propodus and dactylus unknown.

Gills : from gnalhopod 2 to peraeopod 7, not pleated.

Pleonites 1 to 3 dorsally smooth. Epimeron 3 : posteroventral corner narrowly rounded. Urosomites :

urosomite 1 with rounded boss, without lateral flange; urosomite 3 without small dorsolateral spine. Uropod 1 :

unknown. Uropod 2 : unknown. Uropod 3 : peduncle well developed, short, length 1.9 times breadth, without

dorsolateral flange, with 1 apicolateral and 2 apicomedial spines. without midlatcral spines or setae, with

2 distoventral spines, without plumose setae; rami lanceolate, inner ramus reduced, about 0.8 times outer ramus,

outer ramus 2-articulate, article 2 short, article 1 with 2 lateral and 1 medial spines; inner ramus with 1 lateral and

2 medial spines, plumose setae possibly present (broken). Telson : longer than broad, length 1.5 times breadth,

deeply cleft (71%). without dorsal spines, with 1 dorsal seta on each lobe, distal margins incised, without marginal

penicillate or simple setae, with 1 marginal spine on each lobe.

Etymology. — Named for the Passe de Oupi through the reefs north of the Isle of Pines.

Remarks. — For differences from T. ama see remarks under that species.

DISTRIBUTION. — Tryphosella oupi is known only from southern New Caledonia, in 825 to 860 m depth.

Genus WAI.DECKIA Chevreux. 1906

Waldeckia sp. 1

MATERIAL EXAMINED. — Austral Isles. SMCB, R.V. Marara, J.K. Lowry and J.M. POUPIN coll.: stn FRP-1.

21°47.7'S, 154°42'W, north-east side of Maria Island, between Isle du Nordet and Isle Centrale, rubble and coarse sand

samples from lagoon, 1 m, 7 August 1991 : 6 specimens (AM P42161). — Stn FRP-39, 23°19.0'S, 149°29.3 W, otf

Tubuai, baited trap, 485 m, 12-13 August 1991 : 1 9 (AM P42163). — Stn FRP-63, 27°36.55,S, 144°18.65'W. Ha'urei

Bay, Rapa, baited trap on reef with large patches of brown alga, 3 m. 17-18 August 1991 : I juvenile (AM P42164).

Remarks. — Lowry & Stoddart are preparing a monograph on the genus Waldeckia. This species will be

described in that report.

Waldeckia sp. 2

MATERIAL EXAMINED. — Chesterfield Islands. Corail 2 : stn DW 4, 20°52.30'S, 161°36.56E, Fairway Reef,

between New Caledonia and Chesterfield Islands, Halimeda flakes, 64 m, 18 July-6 August, 1988 : I 2, ovigerous

(MNHN-Am 4436). — Stn DW 35. 19°21.65'S. 158°21.5'E, Chesterfield Lagoon, sand. 52 m, 18 July-6 August. 1988 :

1 2 (MNHN-Am 4447).

Remarks. — Lowry & S toddart arc preparing a monograph on the genus Waldeckia. This species will be

described in that report.

218

J. K. LOWRY & H. E. STODDART

ACKNOWLEDGEMENTS

Alain CROSNIER originally encouraged us to study the amphipods from the MUSORSTOM Expeditions and

arranged for Lowry to sort the collections in Paris. Joseph Poupin arranged for Lowry to participate in the

Austral Isles cruise aboard the RV Marara. We are particularly grateful to both of them for their generous

hospitality, expertise and time. We thank Hans Georg ANDRES, Claude DE BROYER and Mike THURSTON who

carefully and critically read and improved our manuscript; Stephen Keable and Kate Dempsey who illustrated the

species and Roger Springthorpe who composed and inked the plates; and the Australian Museum Trust, the

Australian Research Council and ORSTOM, who provided funds which supported the study.

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JLTATS DES CAMPAGNES MUSORSTOM, VOLUME 12 —

RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 - RESULT ATS DE

Crustacea Isopoda : Bopyridae in the MUSORSTOM

collections from the tropical Indo-Pacific

I. Subfamilies Pseudioninae (in part), Argeiinae

Orbioninae, Athelginae and Entophilinae

John C. MARKHAM

Arch Cape Marine Laboratory

Arch Cape, Oregon 97102-0105

USA

ABSTRACT

MUSORSTOM collections from New Caledonia, the Strait of Makassar (Indonesia), the Philippines and near Madagascar

contained many new records and some new species of pseudionines from non-anomuran hosts a new genus and species ot

argeiinc, orbionine parasites of penaeoid shrimps, a single athelgine from 2 hosts and the only entoph.l.ne known

Species of Pseudioninae reported are Pseudione indica Chopra, 1930 (new to the Pacific), infesting Pies, on, ka narval

(Fabricius) (new host record) in the Strait of Makassar; Ionella maculata sp. nov. infesting Calianassa sp. at

New Caledonia; and Giganlione elconaxii sp. nov., infesting Elconaxius sp. at New Caledonia. In the Argennae

Eragia profunda, gen. nov., sp. nov., a parasite of Prionocrangon sp. nov. at New Caledonia. Species of 0rt’,on'lja^

reported (all new geographical and host records, except as noted) are Orb, one hahpori N.erstrasz & Brender a Brandts,

1923, from the Philippines, infesting Haliporoides sibogae (de Man) (not a new host record); Orb, one cf. kempt Chopra

1923, from New Caledonia, infesting Sicyonia Iruncaia (Kubo); Epipenaeon ftssurae Kensley. I974-fr0,n the Slra'‘ °

Makassar, infesting Parapenaeus longipes Alcock (no. a new host record); Parapenaeon japomca <Th*temann .1910)

from near the Strait of Makassar (Indonesia), infesting Metapenaeopsis smica Liu & Zhong, near Panay Island

Philippines infesting Metapenaeopsis velutinus (Dana); NW of Madagascar (not a new locality), infes ing

Metapenaeopsis hilarula (de Man). Parapenaeon brevicoxalis Bourdon, 1981, from Chesterfield Islands, "'jesting

Hymenopenaeus halli Bruce, 1966; Parapenaeon expanse Bourdon. 1979. from the Strait of Makassar, infes i g

Metapenaeopsis sinica Liu & Zhong, from the Seychelles, infesting Metapenaeopsis faoum R“mada"’^"Vf

(not a new record), infesting Metapenaeopsis mogiensis consobrma (Nob.li). and from New _Cal«d°J'®’ ‘nt g

Metapenaeopsis gaillardi Crosnier; and Parapenaeon secunda Nierstrasz & Brender a Brandis, 19 3, near Manila, I p

plT infesSng Metapenaeopsis palmensis (Haswell). and the Strait of Makassar, infesting Metapenaeopsts s.ntca

Markham, J. C„ 1994. - Crustacea Isopoda ; Bopyridae in the MUSORSTOM collections from thetropicallndo^

Pacific I. Subfamilies Pseudioninae (in part), Argeiinae Orbioninae, Athelginae and Entophilinae. /« ‘ ^CROSNIER

(ed.), Resultats des Campagnes MUSORSTOM, Volume 10. Mem. Mas. natn. Hist, nat., 161 . —5-253. Par

85653-212-8.

226

J. C. MARKHAM

Liu & Zhong. In the Athelginae is Pseudostegtas setoensis Shiino, 1933. from Chesterfield Islands, a new locality, where

ImScLc 7Wu 7' Tr‘rZPagUrUS ?V(,° O dcscr,bed as 8en- nov - *P- nov.). In the Entophilinae, Entophilus

ZL Richardson, 1903. ^reported from Chesterfield Islands, a new locality, as a parasite of Muni da "incerta"

Henderson, a previously reported host. Nearly all species are illustrated and described or redescribed.

RESUME

Crustacea Isopoda : Bopyridae des

tropical. I. Sous-familles Pseudioninae

Entophilinae.

collections Musorstom recoltes dans

(en partie), Argeiinae, Orbioninae,

I'lndo-Pacifique

Athelginae et

Des collections MUSORSTOM fanes en Nouvelle-Calddonie. dans le detroit de Makassar (Indonesie), aux Philippines et

Psend3 re8‘°n, ma f Ch£' Pfr"^U°n' de nomhreuscs observations nouvelles et renferment quelques espies nouvelles de

. ludioninac trouvees sur des Crustaces non anomourcs. un nouveau genre et une nouvelle espece d’Argeiinac plusieurs

espfeces d Orbioninae parasites de crevetles peneides, une espece d' Athelginae trouvee sur deux esp&ces^e pagures et Ie

n'avah"nash"nae X",, Lcs, esPeccs de Pseudioninae, dont il est trade, sonl Pseudione indica Chop™ §1 930 (qui

n avail pas encore et 6 signalee dans le Pacifique), recoltee sur Plesionika narval (nouvel hole) dans le detroiule Makassar

“ maCUla,a "°v- ,ro“v^ « Callianassa sp. en Nouvelle-Caledonie. e, Gigamione elconaTsp no7 tX

. r Elconaxius sp. en Nouvelle-Caledonie. Dans les Argeunac. nous decrivons Eragia profunda, gen nov sp nov

Cite cela eTind oT"|SP' “r *" Nouvelle-Calddonie- Les especes d'Orbioninae examinees provfennent toutes, sauf

BraX 1923 ^ des PhT ’ u ,?e°8raPhiquf et d'h6les nouveaux : Or None halipori N.erstrasz & Brender a

dcNonvell‘rilnh,PPmcS' sur Hal,P°ro,de*slb08ae (de Ma-’) (h6te deja connu); Or None cf. kempi Chopra, 19^3

elle-Calcdomc, sur Sicyoma truncata (Kubo); Epipenaeon fissurae Kensley, 1974, du detroit de Makassar sur

Parapenaeus longipes Alcock (hole deja connu); Parapenaeon japonica (Thielemann, 1910) du detroit de Makassar’ sur

STmT1 Sm‘T LlU,f:.Zh0ng' pr6s de Panay Island- Philippines, sur Metapenaeopsis velutinus (Dana) du nord

ouest de Madagascar (zone ddja connue), sur Metapenaeopsis hilarula (de Man); Parapenaeon brevicoxalis Bourdon 1981

es sigmde des des Chesterfield sur Hymenopenaeus halli Bruce. 1966; Parapenaeon expansa BourdoT 979 du dSrod de

Makassar, sur Melapenaeopsts s nica Liu & Zhong. des Seychelles, sur Metapenaeopsis faouzii Ramadan *>1 MadagascS

(local, te non nouvel e) sur Metapenaeopsis mogiensis consobrina ( Nobili), et de No“caldlme su

Metapenaeopsis gat Hard, Crosmer; et Parapenaeon secunda Nierstrasz & Brender a Brandis 190 3 pres de Manille

D <HaSWd,)- C1 dU ddlr°“ dc Mak—' sur ^etapenaeop^LFJa L^i^l

S 0U1 sur un nonvelX Sh""°' 1933' dcs iles Chesterfield, une localite nouvelle ou il®

Pn.lT. r r f Trizopagurus sp. (espece nouvelle qui va etre I'espece type dun genre nouveau) Dans les

MNHNi^onmT ,COlleCti°ns of °RST0N1 housed in ihe Museum national d'Histoire nalurelle (designated

^st^anCs°Thiserenoiie(1pnknCa|1hl0gUeti|an(1 Uniden,ified ^ isoPod*- Pushes of many different decapod

crustaceans. This report deals with a small part ol that collection, with other portions to follow later.

FAMILY BOPYRIDAE Rafinesque. 1815

SUBFAMILY PSEUDIONINAE Codreanu. 1967

Pseudione indica Chopra, 1930

Fig. I

31 : 1 f! ^ °°’,5'S* U7°5TE' 150 m’

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

227

Fig 1 . — Pseudione indica Chopra, 1930. A-J, female. K-R; male. A, Dorsal view. B. Ventral view. C. Right maxilhped

I). Palp of same. E. Plectron of same. F. Barbula. right side. G. Right oostegite 1. external view. H, Same interna

view. I, Right pereopod 1. J. Right pereopod 7. K. Dorsal view. L. Ventral view. M. Right antenna 1. N, Right

antenna 2. O. Right pereopod 1. P. Right pereopod 2. Q. Right pereopod 7. R, Pleon in ventral view.

Scale : 1.0 mm for A-C. F-H. K, L; 0.31 mm for O-R; 0.15 mm for D. E. I. J. M, N.

Source :

228

J. C. MARKHAM

DESCRIPTION of male. — Lenglh 2.31 mm. maximal widlh 0.71 mm. head length 0.40 mm. head width

0.54 mm. pleonal length 0.71 mm. All body regions and segments distinct (Fig. 1K-L). No pigment spots except

for small eyes.

Head suboval, extending well beyond front of body. Eyes as irregular streaks near posterolateral margins of

head. Antennae (Fig. 1M-N) extended far beyond anterior margin of head; first of 3 articles, second of 7 articles,

both setose distally, with only rare setae otherwise.

Pcreon with nearly straight sides, separated laterally by only shallow indentations. Small but conspicuous

mid ventral tubercle on each percomcre. Pereopods (Fig. 10-Q) relatively small and tightly bunched ventrally, not

extending beyond sides of pereon. All articles of all pereopods distinct, dactyli and propodi of first two markedly

larger than others, meri and carpi of pereopods 1 broader and shorter than others; all carpi sparsely setose distally.

Pleon (Fig. 1R) tapering slightly posteriorly, all 6 plcomeres bluntly pointed and shallowly separated laterally.

Midventral tubercles on pleomercs 1 and 2 only, similar to those on pereomeres. Pleopods as 5 pairs of uniramous

subcircular tubercles rising slightly from surface of plcomeres. Final pleomerc broader than long, posteriorly

concave, produced into swollen setose lumps posterolaterally.

Discussion. — This is only the second record for Pseudione indica. Because its type-locality was in the

eastern Indian Ocean, this find considerably extends its known range. An attempt to borrow the type-specimen,

which is in the collection of the Zoological Survey of India. Calcutta, proved unsuccessful because the curator

considered it too badly damaged to be lent; thus remarks on it are based on the published description. The present

female, whose reduced oostegites and lack of eggs mark it as immature, shows a few differences from the holotypc.

the only specimen previously known. Its body is slightly less distorted, and it is proportionately a little broader

(Fig. 1 A-B). The anterior margin of the head is notched medially rather than anterolaterally. Its maxilliped

(Fig. 1C-E) has a longer palp and, on the corner behind the palp, more slender, elaborate compound setation: the

plectron is more extended. Its pereomeres arc more distinctly demarcated dorsally. The present male of P. indica

(Fig. 1K-R) is the first reported. It closely resembles the allotype of P. cognala Markham (1985a), except that the

latter has a quadrilateral-shaped head with second antennae of only 5 articles, not 7, and its pleopods are longer.

Because the host of the types of Pseudione indica was a crangonid, and the present host is in the pandalid genus

Plesionika, one might expect its parasite to be Pseudione affinis (G. O. Sars), which has been reported infesting

6 species of Plesionika around the world. This material is clearly distinct from P. affinis , however, thus : The

temale ot P affinis has a prominent frontal lamina, its barbula has two long projections on each side, the internal

ridge of its first oostegite is ornamented, its pleopodal rami are sharply pointed rather than rounded, and its uropods

are much shorter. The male ol P. affinis has a shorter head, more rounded body and larger pleopods. Despite the

differences cited, the present material belongs to P. indica, though it also has close similarities to P. cognala

Markham (1985a). another parasite of a crangonid. along the coasts of Florida, U. S. A. Although most bopyrids

branchially infesting carideans belong to the subfamilies Bopyrinae and Argeiinae, those found on deep-water

crangomds and pandalids are often pseudionines. as in the case of these three similar species.

Ionella maculata sp. nov.

Fig. 2

iofioM^ERIAL,E"AMINED--NeW Caledonia Lagon : stn DW 1165, 19°15.4'S, 163°19.2'E, 65 m, 30 October

U89. Infesting Callianassa sp., host det. M. DE Saint.Laurent : 1 9. holotypc. 1 6, allotype (MNHN Ep-825).

Description. — Holotype female (Fig. 2A-J). Length 3.6 mm. maximal width 5.2 mm. head length 1.1 mm.

pleon length 1.4 mm. distortion 8° dextrally. Body outline suboval, all body regions distinct (Fig. 2A-B). Circle

of dark pigment spots near edges of dorsal surfaces of several pereomeres.

Head large, indistinctly bilobate, subtrapezoidal in outline, deeply set into pereon; no true frontal lamina, but

fold resembling frontal lamina surrounding anterolateral margins and sides. Antennae minute, first pair in anterior

concavity of head, second pair extending slightly forward (Fig. 2C). Eyes small, near anterolateral corners, visible

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

229

anteriorly not dorsally (Fig. 2C). Maxillipcd (Fig. 2D) of relatively large suboval anterior segment densely lined

with uniformly short setae, lacking palp but slightly produced anteromedially; posterior segment markedly reduced

but bearing extended plectron. Barbula (Fig. 2E) with pair of short broad projections on each side, neither

subdivided, though medial one posteriorly notched.

1,1 ^Scde^To mm for A. B. D. F. G. K. L; 1.0 mm for C. E, J; 0.3 mm for H. 1, O-Q; 0.15 mm for M, N.

230

J. C. MARKHAM

Pereon broadly rounded, all percomeres short and broad, both ends of percon concave and enclosing bolh head

and pleon. Coxal plates on bolh sides of most anterior pcreomeres; pereomcres 6 and 7 produced laterally into

slender falcate, posterolaterally extending projections. Large overlapping oostegites completely enclosing brood

pouch; first oostegite (Fig. 2F-G) suboval, its posterior segment much broader but quite short and lacking

posterolateral projection, its internal ridge digitate laterally; fifth oostegite (Fig. 2J) digitate along posterior

margin. Pereopods (Fig. 2H-I) with all articles distinct, all of nearly same size and structure, though dactyli

somewhat smaller and bases and ischia broader posteriorly; meri slightly setose anteriorly.

Pleon (Fig. 2J) short and ventrally mostly hidden beneath oostegites. Six pleomeres visible dorsally, not

produced into lateral plates. Biramous foliate pleopods tightly bunched together, anterior ones much larger than

others, especially on longer side of body, those of first 3 pairs on that side extending far laterally. Other pleopods

and uniramous uropods reduced and tightly clustered beneath last oostegite.

Allotype male (Fig. 2K-Q). Length 3.44 mm, maximal width 1.04 mm, head length 0.44 mm, pleon length

0.80 mm. Slender and straight, with all segments clearly demarcated (Fig. 2K-L). Dorsal pigment spots on side of

pereomere 1 and near sides of anterior pleomeres.

Head semicircular, slightly narrower than pereon. Prominent eyespots near posterolateral angles. Antennae

(Fig. 2M-N) tiny, first of 3 articles, second of 5 articles, both with sparse but long setae (ends of second antennae

obscured, so presence of terminal setae uncertain).

Pereon with nearly parallel sides, lirst and last pcreomeres slightly narrower than others, all markedly separated

by deep anterolateral indentations. Pereopods (Fig. 20-P) small, all of nearly same size, with carpi distally

sparsely setose; first 5 pairs with large dactyli and reduced carpi peculiarly connected to sides (not distal margins)

ot men so as to touch ischia.

Pleon triangular in outline, of 6 pleomeres. All pleomeres of same length, each about half as long as any

pereomere; final pleomere reduced to small knob. Pleopods 1 and 2 (Fig. 2Q) as biramous flaps; pleopods 3-5 as

uniramous knobs to flaps. Uropods distally setose extended uniramous flaps.

E i i moi.ogy. — The name maculata (= spotted) refers to the dorsal pigment spots forming a circle on the

female and those scattered on the male.

Discussion. — The genus Ionella Bonnier contains 2 previously described species. The type-species

/ agassizi Bonnier, 1900. which has been thoroughly described and redescribed, seems to be a fairly common

parasite of Callianassa uncinata H. Milne Edwards along the coast of Chile (Bonnier, 1900: SmiNO 1964'

STUARDOer a/., 1986). /. murchisoni Danforth. 1970, which has been recorded only once infesting a

Callianassa sp. in Hawaii (Danforth, 1970a). was, and remains, inadequately described, so its proper generic

placement is uncertain. I. maculata illustrates the generic characters shown by /. agassizi. and. where known most

shown by /. murchisoni. Females : broadly oval, barely distorted body; proportionately broad quadrilateral head

deeply inset into pereon; suboval palpless maxilliped, its anterior segment densely fringed clear around its margin

y s ort setae and much the larger; first oostegite with large subcircular anterior segment, its internal ridge

ornamented only laterally, and its posterior segment much the shorter and broader, without posterolateral point;

absence ot lateral plates from most pereomcres and pleomeres; posterior pereopods with wide bases and ischia'

brood pouch completely enclosed by oostegites (fifth ones posteriorly fringed); first pleomere forming broadest

part ot body, others rapidly narrowing, with final (sixth) one small, triangular and nearly or completely set into

tifth, biramous foliate pleopods tightly clustered on short pleon. anterior ones on long side greatly extended

aterally and distorting general body outline; and reduced uniramous uropods. Males : extended head, complete

separation of all body regions, lack of midventral tubercles, reduced pereopods. biramous anterior pleopods, and

posteriorly extending uniramous uropods. All three species infest species of Callianassa. 1. agassizi differs from

' maculata in that its female is more nearly circular, its barbula is more elaborately toothed, its maxilliped is not

produced antcromcdially. all of its pcreomeres lack lateral plates, its pleomeres arc not distinctly separated, and its

ina pleomere is completely embedded in the preceding one; the male of /. agassizi is proportionally broader, all of

its pleopods are biramous, and its uropods are much smaller. The female of/, murchisoni has indistinctly separated

pleomeres. the last ot which is not extended, and has deeply digitate plcopodal rami; the male of that species is

BOPYRIDAE FROM THE TROPICAL INDO-PAC1FIC

231

more oval in outline, its antennae are longer, and its plcopodal rami much more separated and extended. Inclusion

of I. metadata in Ionella produces two slight exceptions to the original generic diagnosis because the female has

lateral plates on two percomeres. and only the first two pleopods of the male are biramous; the necessary resultant

modifications of the diagnosis are not serious, especially in light of the other extensive consistency of characters

of the species listed above.

Damage prevented specific identification of the host, so it is unknown whether this species of Callianassa is

previously known as a bopyrid host.

Gigantione elconaxii, sp. nov.

Figs 3-4

MATERIAL EXAMINED. — New Caledonia. BlOGEOCAL : sin. CP 290, 20°36.91'S, 167°03.34E. 920-760 m,

27 April 1987. Infesting Elconaxius sp.. host (damaged) del. M. DE Saint Laurent : 1 9, holotypc, 1 6 . allotype

(MNHN Ep-820).

FIG. 3. - Gigantione elconaxii , sp. nov., holotype female. A, Dorsal view. B Right antennae. C. Left maxillfpecL

D. Right side of barbula. E, Right oostegite 1, external view. F. Same, internal view. G, Right pereopod 1. H, g

pereopod 7. I, Pleon in ventral view. „TI-10 r a

Scale : 1.00 mm for A, D. E, F, I; 1.80 mm for C; 0.35 mm for G. H; 0.18 mm for B.

Source : MNHN . Paris

232

J. C. MARKHAM

Description. — Holotype female (Fig. 3). Length 2.8 mm. maximal width 3.2 mm. head length 0.9 mm.

pleon length 0.6 mm. distortion 19° dextrally. Body outline suboval, all body regions distinct (Fig. 3A-B). No

pigment spots.

Head large and deeply set into percon, trapezoidal in outline, bisecting first pereomere dorsally. Antennae

(Fig. 3B) arising from deep anterior concavity; first one of 3 articles, of which 2 proximal ones very broad; second

antenna of 8 articles, extending far beyond margin of head, both sparsely setose on distal ends of some articles.

Maxilliped (Fig. 3C) suboval, lacking palp, anterior article much larger than posterior one and lined with fringe of

long setae. Barbula (Fig. 3D) with 2 short simple projections on each side, lateral one much narrower than medial

one.

Pereon broadest across pereomere 4. that also longest pereomere. Pereon encircling both head and pleon.

Dorsolateral bosses on both sides of pereomeres 1-4; pereomeres 4-7 produced into slender curved points.

Oostegitc 1 (Fig. 3E-F) as broad as long, with entire internal ridge, no posterolateral projection, posterior border

fringed by tiny setae. Pereopods (Fig. 3G-H) relatively small, all of about same size, though bases broader

posteriorly; articles variously setose.

Pleon short, with anterior pleomeres overreaching posterior ones. All pleomeres except last (sixth) produced

into sharp extended lateral plates. Pleomcre 6 long and slender, deeply embedded into pleomere 5. Pleopods

(Fig. 31) reduced to irregular biramous Baps tightly bunched behind brood pouch. Uropods, visible only in ventral

view, biramous. each ramus extending posteriorly as small lanceolate flap.

Allotype male (Fig. 4). Length 1.89 mm, maximal width 0.86 mm. head length 0.39 mm. pleon length

0.45 mm. Body tapering toward both ends, with all segments clearly demarcated (Fig. 4A). No pigment spots.

Head quadrilateral, prominently extended from pereon. No eyes. Antennae (Fig. 4B) both very similar to those

ol female in structure, size, setation and great lateral extension from sides of head, except antenna 2 of more (11)

articles.

Pereon broadest across pereomere 5, sides of pereomeres only moderately separated. Pereopods (Fig. 4C-D)

reduced, not reaching beyond sides of body, all of about same size; all dactyli long and propodi broad relative to

other articles.

Pleon ol 6 pleomeres. all much shorter than pereomeres; last pleomere barely extending beyond fifth; lateral

margins of most pleomeres setose. Five uniramous pleopods as small Haps to knobs. Uniramous uropods’similar

to pleopods but larger: no appendages extending to margins of pleon.

Fig. 4. — Giganiione elconaxii, sp. nov., allotype male. A, Dorsal view. B, Left

pereopod 7. E. Left margin of pleon in ventral view.

Scale : 1.00 mm for A; 0.18 mm for B E.

antennae. C, Left pereopod 1. I), Right

BOPYRIDAE FROM THE TROPICAL INDO-PAC1FIC

233

Etymology. — The name elconaxii is the genitive of the generic name of its host, in the axiid genus

Elconaxius.

Discussion. — So far, 1 1 species of Gigantione have been described, but some remain poorly known. The

female of G. elconaxii differs from all other species by having a relatively narrow trapezoidal (rather than

rectangular or suboval) head, which bisects the first pereomere and bears dorsally placed antennae; its male differs

from other species in having very long second antennae, in being more oval, with its pereomercs less separated and

its plcopods shorter and its uropods less extended. The females of the other species differ thus : G. bouvieri

Bonnier, 1900. as redcscribed by Monod (1932). has a more extended head, broader lateral extensions on the

longer side of the pereon and much more conspicuous uropods; G. giardi Nobili (1906), as described by Nobili

(1907), has a more extended head and uropods, a more elaborate internal ridge on the first oostegitc and digilate-

margined pleonal lateral plates; G. hemaiiensis Danforth. 1970 (as described without name by DANFORTH, 1963),

differs the same ways as G. bouvieri ; G. ishigakiensis Shiino (1941) and G. moebii Kossmann (1881), which may

be synonyms, have more extended heads, longer but less sharply tapered coxal plates and lateral plates and more

prominent uropods; G. mortenseni Adkison (1984). the only species known from a large series, has a conspicuous

frontal lamina, longer coxal plates and more prominent uropods; G. pikei Page (1985) is subrcctangular with all

segmentation obscure and uropods conspicuous but not at all extended; G. pratti Danforth (1967) has an extended

head with small antennae and a posteriorly distorted body; G. rathbunae Stebbing (1910) is similar to G. bouvierr,

G sagamiensis Shiino (1958) and G. i iberlackerae Adkison (1984). which are similar, have longer coxal plates and

more extended final pleomeres and uropods. This is the first record of bopyrid infestation of any species of

Elconaxius , though species in 3 other axiid genera arc known as hosts.

The two genera considered here, Gigantione Kossmann, 1881, and lonella Bonnier, 1900, are dilticult to

distinguish. The problem of generic definition was aggravated by the very inadequate description of the type-

species of Gigantione. G. moebii by Kossmann (1881), a parasite of a xanthid crab on Mauritius. G. moebn was

not reported again until BOURDON (1969) published an account of additional material from another xanthid in

Mauritius and presented the first proper description. BONNIER (1900) stated that "Deux caracteres suffisent a

caractdriscr ce genre : la femelle adultc possedc des lames pleurales sur tous les somites, tant ceux du thorax que

ccux de l'abdomen, ct ses uropodcs sont birames." Without referring to Gigantione. Bonnier (1900) diagnosed

lonella : "Ce genre...est nettement caractcrise par l'absence des lames pleurales sur les somites du pleon. par les

uropodcs simples et enfin par les pleopodes qui sont biram6s. non seulement dans le sexe femelle, mais aussi dans

le sexe male..." The two genera are evidently closely related. Females of both are only slightly distorted, then

bodies are at least as wide as long, their large heads are deeply inset, their palplcss maxillipeds are densely setose;

some of their pereopods arc produced into sharp points; and their pleonal appendages arc reduced and tightly

bunched ventrally. The males are more different, but both have distinct segmentation, extended heads, six

pleomeres and conspicuous naplike plcopods. Females of lonella have reduced antennae, pleons distorted to one

side with plcopods on that side much enlarged and uniramous uropods; those of Gigantione have very long

antennae symmetrical pleons and biramous uropods. Males of lonella have most or all plcopods b.ramous, while

males of Gigantione have variable pleopods. Known hosts of lonella spp. are all species ol Calltanassa : reported

hosts of Gigantione spp. are brachyurans except for three in the thalassinidean family Axndae (two Ax, ops, s spp.

and an Elconaxius).

SUBFAMILY ARGEIINAE Markham. 1977

Genus ERAGIA nov.

Diagnosis. — Female : Body only slightly distorted. Quadrilateral head embedded in pereon, bearing very

reduced frontal lamina; maxillipcd with anterior article about twice size of posterior one, articulating pa p sma .

Pereon suboval, broader than long; some anterior dorsolateral bosses weakly developed, first oosteg.te with nejly

unomamented internal ridge and very slender posterolateral projection; pereopods 1 and 2 much smaller than others.

234

J. C. MARKHAM

c ose to head percopod 3 or percopods 3 and 4 on sides of body: other pereopods aligned behind brood pouch

Pleon. of 6 pleomeres. greatly extended, with ends of long lateral plates forming nearly parallel sides: 5 pairs of

3S lmy k"°bS: S,nJClure of uropods ambiguous. Male : Unknown. Host : In crangonid genus

~ E!ar iS an anagram 0f Ihe gcneric names Argeia and Gareia- aPP'ied 10 previously described

genera of the subfamily Argcunae. Gender feminine.

Type-species. — By present designation. Eragia profunda sp. nov.

Eragia profunda sp. nov.

Fig- 5

MATERIAL EXAMINED. — New Caledonia. Biocal : stn CP 77 ',',<’I0'S lfi7°VTF oinooiin a c ,

1985. Infesting Prionocrangon sp. nov., host det. L. B. Holthuis. 1 $ .‘hoTotype (MNHN-Ep‘829). m* SepIembcr

Description - Holotype female (Fig. 5). Length 4.60 mm. maximal width 4.67 mm. head length 0 95 mm

pleonal length 2.12mm. All body regions separated, head embedded in oval pereon. but pleon greatly extended

posteriorly (Fig. 5A), making body key-shape overall.

fFipHl5R rwSidalHmfUCh W'der ‘rar l0ng' bCaring very reduccd fro,,IaI lamina antcrodorsally. No eyes. Antennae

uL H~d F c sCn p 7', °ne,0f 3 ar,ldCS- ,SeC°nd °f 4 ar,idcs- eadl anIen"a wiIh ^arse terminal setae

sTto^e min n suboval anterior article about twice as long as posterior one: short broad articulating

trih 2 small fa. en0r mar8,n maXllliped‘ no1 reachi"8 mos> anterior par. of maxillipcd. Barbula (Fig 5G)

with 2 small falcate projections on each side. v 8 1

Pereon oval much broader than long, pereomeres 1-3 only obscurely separated, and pereomere 7 concealed

orsa ly beneath pleon. Coxal plates slightly developed on pereomere 1. Indistinct dorsolateral bosses on both

sid s of pereomeres 1-3 Oostegites completely surrounding brood pouch bu, no, fully covering TZstcgTl

irnr ^ ,r<Td<7 cach Pla,c *<1™^ ^ng. with internal ridge unornamented except for~ single club ike

CSSr rTTl1 P,r°JCCli0n: fifIh 00S,CgiteS Wi,h S0mc l0"g se*ac 'posterior^'6

Pereopods 1 and 2 (Fig. 5J) tiny, close beside head on anterior margin of pereon; percopods 3-7 (Fi<- 5K) much

^

P°Tri°rf, SiX ple0meres- firsl onc Kr* shon' s“ond longer,

St taiy extfnd Z Z , SeC°"?' S‘X!h l0”geS‘' A" plCOmeres' indudi"B fi"d »”<=' produced into long

mrlit r extending, nontapering lateral plates: ends of lateral plates 2-5 making sides of pleon ncarlv

either long'L"^ P‘e0P°dS' S"lh ftam” ^ P«“"- ““■> ^tnre,

Male : Unknown.

,he cp,iMion °f ,his «-*■ * a ** -

1977, and the m„„„,yp,c genns Oareia described by Bourdon * Bruce, ?983),

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

235

key-shaped body outline caused by the peculiar extension of the plcon and by the arrangement and size differential

of its pereopods. For these reasons it is placed in a new genus.

This is the first record of bopyrid infestation of any species of the deep-water crangonid genus Prionocrangon.

Although the host has been identified as a new species, it is unknown when it may be described.

FlG. 5. — Eragia profunda gen. nov.. sp. nov., holotype female. A. Dorsal view. B. Right antenna 1. C, Right

antenna 2. D, Maxilliped. E, Palp of same. F. Plectron of same. G. Right side of barbula. H. Right oostegite 1,

external view. I. Same, internal view. J. Right pereopod 1. K. Right pereopod 7. L, Posterior pereopods and pleon in

ventral view. _ _

Scale : 2.0 mm for A. L; 1.0 mm for D. G-I; 0.35 mm for J, K; 0.18 mm for B, C, E. F.

SUBFAMILY ORBIONINAE Codrcanu. 1967

Orbione halipori Nierstrasz & Brendcr a Brandis, 1923

Fig. 6

Orbione halipori Nierstrasz & Brender a Brandis. 1923 : 64-66, fig. 2a-h [Paternoster and Kei Islands Illdonesi^

infesting Haliporus sibogae (de Man) = Haliporoides sibogae (de Man)]. — Dakin. 1931 : -70. — SHHNO. 1 19 9 . .. .

55- 1952 : 38- 1958 : 53-55, fig. 14 [off Owase, Mie Prefecture, Japan, infesting Parahaliporus sibogae (de Man)

Haliporoides’ sibogae]-, 1972 : 8. — BOURDON, 1979a : All-All. 480, figs 1-3 [Madagascar, infesting H. sibogae

Source : MNHN, Paris

236

J.C. MARKHAM

madagascariensis Crosnier and Hymenopenaeus halli Bruce]; 1979c : 431 [Reexamination of types; also Lorenzo

Marques, infesting Hymenopenaeus triarlhrus (Stebbing); and Sydney, Australia, infesting Aristaeomorpha foliacea

Risso; incorporation of O. halipori var. libera and O. natalensis into O. halipori]; 1982 : 195. _ Markham, 1982 :

362-365, 385, figs 21, 22 [Hong Kong, infesting Metapenaeus ensis (de Haan)]; 1986 : 144, 159, fig. II. _ Owens

& Glazebrook, 1985 ; 107, table 2 [Gulf of Carpentaria, Australia, infesting M. crisis], — Page 1985 • 208 _

Morton, 1988 : 94, 95, 118, pi. 31.

Crassione arisiaei Dakin. 1931 : 268-272, text-figs 1-9, pi. XIV [Off New South Wales. Australia, infesting Arisleus

foliaceus (Risso) = Aristaeomorpha foliacea (Risso)].

Orbione halipori var. libera Nierstrasz & Brender a Brandis, 1931 ; 155 [Off Wain, Moluccas, Indonesia, infesting host

^b^equenUy identified as Hymenopenaeus lucasii (Bate)]. — SmiNO, 1949 : 52, 55; 1965 : 463, fig. 1. — Bourdon,

Orbione natalensis Carton, 1970 : 47 [Nomen nudum. Mozambique Channel, infesting Hymenopenaeus triarlhrus

(Stebbing) = Haliporoides triarlhrus Stebbing].

Orbione natalensis Bourdon, 1972 : 104-107; figs.3-6 [Near Natal, South Africa, infesting H triarlhrus]

Non Orbione halipori libera Shiino, 1934 : 258-260, fig. 1 [Tanabe Bay. Japan, infesting Solenocera distincta (de Haan);

description]; 1972 : 8. — BOURDON, 1981 : 242 [excluded from species].

MATERIAL EXAMINED. — Philippines. Musorstom 2 : stn 83 : 13°55.2'N. 120°30.5'E, 320-318 m. 2 December

1980. Infesting Haliporoides sibogae (de Man), det. of host A. Crosnier : 4 9 , 4 <5 (MNHN-Ep 821).

Discussion. — Orbione Itaiipori has been described and rcdescribed in such thorough detail that it would be

superfluous to add anything here. The greatly enlarged and medially arching coxal plates on the first 2 pereomeres

ol the female, with their fringed edges, are very distinctive. In the present material, the other 3 females have more

prominent coxal plates on the long sides of pereomeres 3-7. and one also has a noticeably smaller maxilliped.

Three of the present females arc sinistral. only one dextral. This is the first record of O. halipori from the

Philippines, but the host is the same recorded for the types in Indonesia.

Orbione cf. kempi Chopra, 1923

Fig. 7

Orbione kempt Chopra 1923 . 416, 419. 446, 447-451. text-fig. 4. pi. XII, figs 1-5 [Bay of Bengal, infesting Sicyonia

btspmosa de Haan], — Shiino, 1949 : 52, 55. — Bourdon, 1981 : 243. — Devi, 1987 ; 23. 28, 31. table 1

[Kakinada, India, infesting Metapenaeus brevicornis H. Milne Edwards and M. lysianassa (de Man)|

«Orbione» kempi - BOURDON. 1981 : 243; table I [Questioning of generic slatus].

in ^ATER|AL EXAMINED. —New Caledonia. Smib 4 : stn DW 68. Norfolk Ridge, 22°55.0'S, 167°16.0'E. 440 m.

It) March 198 J. Infesting Sicyonia truncata (Kubo). host det. A. Crosnier : 1 9, 1 6 (MNHN-Ep 815).

DISCUSSION. — The present material differs from the type material of Orbione kempi of Chopra (1923) in a

lew respects, and it is not fully certain that it is assignable to it. To indicate that further material may prove that

it is actually an undescribcd species. I have reported it as Orbione cf. kempi. The original description was based on

a single pair, and that is all that is present here, so the range of variability in O. kempi is uncertain. This female

is slightly more distorted, the projections on its barbula less extended, the posterolateral projections on its first

oostegites slightly longer and narrower, and its uropods less extended. (The peculiar extension and division of the

lateral plate on the left side of pleomcrc 1 are probably an individual anomaly.) The body proportions, frontal

lamina, shape and palp of the maxilliped. coxal plates, sizes and basal enlargements of the pereopods and structure

o Ihe P eonal aPPcn(1ages all conform with those of the types. Although some details of the type male arc

uncertain, the present male seems to agree with it in most characters, except that its second antennae have four

articles, not three, and the dactyli of its pereopods become somewhat smaller posteriorly. Like the types, this

material infests a species of Sicyonia. assigned to 5. truncata. a new specific record. New Caledonia is a new

locality record for O. kempi.

Source :

BOPYRIDAE FROM THE TROPICAL INDO-PACIF1C

237

FIG. 6. - Orb, one haUpori NiersUasz & Brender a Brand, s. 1923. A-J, female. K-O. male. A. Dorsal view. B Ventral

view C Right antenna 1. I). Right antenna 2. E. right maxtlliped. F. Right side of barbula. G Right oostcgite

and coxal plate, external view. H. Same, internal view. I. Right pereopod 1. J. Right pereopod 7. K. Dorsal view.

L. Ventral view. M. Right antennae. N. Left pereopod 1. O. Left pereopod 7

Scale : 4.00 mm for A. B; 0.96 mm for C. D. 1. J; 1.92 mm for E-H. K, L; 0.15 mm for M. 0.30 mm for N. O.

Source : MNHN, Paris

238

J. C. MARKHAM

F,Gn ■-°rb‘one c^kZ'P‘ Ch°pra' 1923’ A’K' female; L-P- male- A. Dorsal view. B, Ventral view. C. Left maxilliped.

. 1 a p °J. same’ E' Plectron of same- F- Right side of barbula. G. Right oostegite 1. external view. H. Same, internal

view. 1, Right pereopod 1. J, Right pereopod 7. K. Picon in ventral view. L, Dorsal view. M, Ventral view N Left

antennae. O, Right pereopod 1. P, Right pereopod 7.

Scale : 1.00 mm for A, B, F-H, K; 0.50 mm for C. L, M; 0.31 mm for D, E, I, J, O, P; 0.15 mm for N.

Source :

BOPYRIDAE FROM THE TROPICAL INDO-PAC1EIC

239

Epipenaeoti fissurae Kcnsley, 1974

Figs 8-9

Epipenaeon fissurae Kcnsley, 1974 : 261-263, fig. 2a-j [Off Natal, South Africa, infesting Parapenaeus fissurus Bate];

1978 : 152, 153; fig. 67F-G. — Bourdon, 1979a : 498-501, figs 19-20 [Near Madagascar, infesting P. fissurus-,

redcscription]; 1979c : 428 [Bay of Bengal, infesting P. longipes Alcockl; 1981 : 239, 242. 255, 259-260

[unspecified locality in Philippines, infesting unidentified penaeid], — Nearhos & Lester, 1984 ; 258.

MATERIAL EXAMINED. — Indonesia. Strait of Makassar. CORINDON 2 : stn 205, 01°05.0'S, 117°45.2'H, 85-

79 m, fish trawl, 30 October 1980 : 7 9 , 6 6 (MNHN Ep-827). — Stn 295. 01°26.5'S, 1 17°02.1,E. 54-51 m, fish trawl,

11 November 1980 ; 4 2 , 3 6 (of which 1 2 and 1 6 drawn) (MNHN Ep-826). All infesting Parapenaeus longipes

Alcock, hosts det. A. CROSNIER.

Fig. 8. — Epipenaeon fissurae Kcnsley. 1974, female. A, Dorsal view. B. Ventral view. C. Right maxilliped. D. Right

side of barbula. E, Right oostegite 1. external view. F, Same, internal view. G, Left pereopod 1.

Scale ; 2.0 mm for A. B; 0.3 mm for C-F.

Discussion. — Diagnostic characters for E. fissurae, as described and illustrated by Kensley (1974) and

Bourdon (1979a). are, for the female, the shape of the head (Fig 8A). the angled nonarticulating maxilliped palp

Source

240

J.C. MARKHAM

(Fig. 8C), the deeply digitate processes on the barbula (Fig. 8D). the elaborate internal ridge of the first oostegitc

(Fig. 8F), the tuberculate pleopods (Fig. 8B) and the notched final pleomere (Fig. 8A); and, for the male, the

shape and proportions of the body regions (Fig. 9A) and pereopods 1 and 2 with larger dactyli and shorter meri and

carpi than in the other 5 pereopods. The specimens previously illustrated are quite similar to each other, but where

they differ, the present material is more similar to that of Bourdon (1979a). The female illustrated has a slightly

more slender posterolateral projection on the first oostegitc and a posterior pleon notch of length intermediate

between those previously illustrated. The male illustrated lacks eyes. Of the females examined, three are dextral and

eight simstral. Parapenaeon longipes is not a new host record for Epipenaeonfissurae , but Strait of Makassar is a

new locality.

F'GpeLT.dTeE:rf(pX"7K“Sley' 1974' *• D~1 ™W' B- V~- *"'• C- "W. Left

Scale : 2.0 mm for A, B; 0.3 mm for C-E.

Parapenaeon japonica (Thielemann, 1910)

Figs 10-11

EpiPZTnJTn‘Z Thiel“- 1910 : 7- ?9-81. 106-107. 108. text-figs 86, 87. table 8. pi. 2 fig. 31 [Okayama

hTraiwa Tq"^ SP‘ ■ _,C“0PRA- 1923 : 454- 458- - NiErstrasz & Brender a Brandi!, 1929 : 302. -

M aclivis (Rathbunl anii I“xt'flgs l'23- P11 [Hiroshima Bay, Japan; infesting Metapenaeopsis barbaius de Haan,

R , (Rathbun) and M. lamellate (de Haan); morphology and systematics); 1934 : 45-62, figs 1-14 (Hiroshima

Sevelopmentl " Mosod m3 W ‘"'T' 1936 : 101 '137- Pls ^ [Same locality and host;

NaSi 43~J[T i I - Rpverberi- 1942 : 60. - Reverber, & Pitottu 1942 : 116, 123. -

iqsq sa so aa’ ‘ 8 (Travancore, India, infesting unidentified penaeoid]. — Morris, 1948 : 4; 1949 • 21 1 _ oA7I

1959 : 56, 59-60. - Ivanov, 1982 ; 197. - Cash & Bauer, 1993 -123 ^

Ep.penaeonjaponicun, - Nierstrasz & Brender a Brandis, 1923 : 68; 1931 : 58. - Barnard 1925 • 408 IDelagoa Bav

Pr~'™ - ” : - <%r-

IK 8 DPR,lI,A^.N & Rosenfield, 1967 : 361. — Sindermann, 1970 : 171. — Devi, 1987 : 23, 26-27 31 3"> table I

[Kakinada, India; infesting Penaeus japonicus Bale], ’ ’

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

241

Apopenaeon japonicum var. hiraiwai Shiino, 1950 : 151-155, fig. 1 [Kulino, Izo, Japan; infesting M. aclivis ]; 1958 :

51-53, fig. 13, table I, plate III fig. 7 [Several Japanese localities; infesting M. aclivis and M. barbatus ].

?"Bopyrid parasites" [in part] - CHEUNG, 1963 : 428 [Hong Kong; infesting one or more species of five penaeid genera].

Apopenaeon japonicum hiraiwai - STROMBERG, 1971 : 6, 7, 29, 31, 33-34, 39, 42. — SHIINO. 1950 : 155 [Japanese

records], — PaLISOC, 1987 ; 286.

Apopenaeon japonica - SHIINO, 1950 : 155.

FIG. 10. — Parapenaeon japonica (Thielemann, 1910). A-L. female. M-R. male. A. Dorsal view B .Ventral view.

C Right antennae. D, Maxilliped. E, Palp of same. F. Plectron of same. G, Right side of barbula. H, Oostegite 1.

external view. I, Same, internal view. J. Right pereopod 1. K. Right pereopod 7. L. Pleon, ventral view M. Picon,

dorsal view. N. Right antennae. O. Right pereopod 1. P, Right pereopod 7. Q. Pleon in dorsal view. R, Same in

VCn'r ScIleT4.0 mm for A. B. G; 2.0 mm for C, D. H, I, L. M. Q; 0.6 mm for E. F. J. K; 0.15 mm for N.

242

J. C. MARKHAM

Parapenaeon japonicum - BOURDON, 1979a : 480-486. figs 6-7 [Madagascar; infesting M. monoceros (Fabricius) and

Penaeus semisulcatus de Haan; redescription and discussion]; 1979c ; 432 [Gulf of Martaban, Burma; infesting

Parapenaeopsis sculptilis (Heller)]. — BRANFORD, 1980 : 276, table 1 [Red Sea; infesting Penaeus latisulcatus

Kishinouye], — Ivanov, 1982 : 196 [Western Indian Ocean; infesting Melapenaeus monoceros], — MlQUEL, 1982 :

94. — Markham. 1986 : 159. — Anderson, 1990 : 290. — Courtney, 1991 : 615, 617, 620, table 2 [Central coast

of Queensland, Australia; infesting Penaeus longislylus Ktibo].

Parapenaeon japonica - MARKHAM, 1982 : 366-369, 385, fig. 23-24 [Pearl River Estuary, Hong Kong; infesting Penaeus

japonicus]; 1985b : 3, 50-51, 63, table [Chonburi, Thailand; infesting Trachypenaeus fulvus Dali].

Epipenaeon - Wilson, 1991 : 239. fig. 13.4.

Material EXAMINED. — Indonesia. Strait of Makassar. CORINDON 2 : stn 295, 0P26.5'S, 117°02.3'E, 54-

51 m, fish trawl. Infesting Metapenaeopsis sinica Liu & Zhong, del. of host A. CROSNIER : 1 2, 1 <3 (MNHN-Ep 817)

Philippines. MUSORSTOM 3 : stn CP 134, 12°01'N, 121°57'E. 92-95 in. 5 June 1985. Infesting Metapenaeopsis

velutmus (Dana), del. of host A. Crosnier : 2 9 , 2 <3 [of which 1 d drawn] (MNHN-Ep 832).

NW of Madagascar. Banc du Geyser. BENTIIEDI : stn 117, lagon. partie sud, 1 1 April 1977, 3-8 m on coralline rock

with Halimeda fragments. Infesting Metapenaeopsis hilarula (de Man), det. of host A. Crosnier : 1 9, 1 <3 [both drawnl

(MNHN-Ep 816).

Discussion. — Diagnostic characters for P . japonica

are, in the female, the general body shape and proportions,

the large coxal plates on the short side of the first two

pereomcres and on the long side of all pcreomeres, the

irregularly digitate internal ridge of the first oostegite, the

short foliate tuberculate pleonal appendages and the

posterior notch of the final pieomere; and, in the male, the

extended head, the hu ge anterior dactyli. the fringed propodi

and the semicircular plcon with an occasional posterior

indentation. One female has the nonarticulating extended

palp anteriorly rather than anterolaterally placed on the

maxillipcd, one male has its head fused with the pereon.

and one (Fig. 11) has eyes, in addition to the variable

pleonal notch. Some hosts of Parapenaeon japonica

previously reported have belonged to the genus

Metapenaeopsis , but all three of the present hosts tire new

species records. P arapenaeon japonica is a very widespread

species whose known range, from the Red Sea and

Madagascar to Japan, is essentially that of its subfamily as

a whole. Still, the finds in the Strait of Makassar and the

Philippines are new geographical records.

As the synonymy indicates, there has been much confusion about the gender of the generic name Parapenaeon.

It would seem to be a Greek neuter noun, but Richardson (1904). in describing its type-species. Parapenaeon

consolidata, indicated by the feminine ending that she considered her new generic name to be feminine. I have

I ol lowed that precedent in calling this species P. japonica and the following ones P. brevicoxalis , P. expansa and

P. secunda.

Fig. 11. — Parapenaeon japonica (Thielemann,

1910), male. A, Dorsal view. B, Ventral view.

Scale ; 1.0 mm.

Parapenaeon brevicoxalis Bourdon, 1981

Figs 12-13

Parapenaeon brevicoxale Bourdon, 1981 : 239. 249-251. figs 10-11, table II [between Bohol and Cebu Islands,

Philippines; infesting Penaeopsis rectacuta (Bate)].

Ma'I TRIAL EXAMINED. — Chesterfield Islands. MUSORSTOM 5 : stn 364: 19°45.30'S, 158°46.50'E. 675 m,

IV October 1986. Infesting Hymenopenaeus halli Bruce, det. of host A. Crosnier : 2 9 , 1 <3 (MNHN-Ep 830). —

Source : MNHN. Paris

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

243

Sin 365, 19°42,82'S, 158°48.00'E, 710 m, 19 October 1986. Infesting Hymenopenaeus halli Bruce, det. of host

A. CROSN1ER : 7 $ (1 infested by hyperparasitic Cabirops sp.; 1 drawn). 6 6 (1 drawn) (MNHN-Ep 831).

FIG 12. —Parapenaeon brevicoxalis Bourdon. 1981. A. Dorsal view. B. Ventral view. C. Right antenna 1. D, Right

antenna 2. E. Right maxilliped. F. Right side of barbula. G. Right pereopod 1. H. Right pereopod 7. I, Right

oostegite 1, external view. J, Same, internal view.

Scale : 2.0 mm for A, B. E, F; 1.0 mm for C, D. G, H.

Discussion. — The female of Parapenaeon brevicoxalis drawn (Fig. 12) is quite similar to the holotype,

except for some minor details. Its frontal lamina extends less to the sides, its head is longer and narrower

(Fig. 12A), its maxilliped palp is slightly broader (Fig. 12E), the posterolateral projection ol the first oostegite is

broader (Fig. 12I-J), the pereomeres are separated clear across, and the sixth pleomere is visible dorsally

(Fig. 12A). The male drawn (Fig. 13) differs in having a narrower head and slightly more extended plcon. Three

of the females are dextral and seven sinistral; all closely resemble the one illustrated. One of the males examined

has a markedly pointed pleon, but otherwise the males arc essentially alike. Although one of the temales bore a

hyperparasite, it was also accompanied by a male, an unusual occurrence. This is the first record ol P. brevicoxalis

since its initial discovery in the Philippines, so Chesterfield Islands is a new locality record for it; Hymenopenaeus

halli is a new host record.

244

J.C. MARKHAM

Fig. 13 . — Parapenaeon brevicoxalis Bourdon, 1981, male. A, Dorsal view. B. Ventral view. C, Left antenna 1 D Left

antenna 2. E, Lelt percopod 1. F, Left pereopod 2. G. End of pleon in ventral view.

Scale : 1.00 mm for A. B; 0.12 mm for C. D; 0.31 mm for E, F; 0.53 mm for G.

Parapenaeon expansa Bourdon, 1979

Fig. 14

Pa71T-Op CXPanTu ,B0Urd°"’ 19193 ' 494' 495-498’ fiSs 15‘17- l8b'c I Near Madagascar, infesting Penaeus teraoi

Kubo -Penaeus (Melt certus) marginal us Randall], — Nearhos & Lester, 1984 : 257, 258 [Moreton Bay, Queensland

Australia, infesting Penaeus (Met, certus) plebejus Hess; and Karumbu, Gulf of Carpentaria, Australia, infesting

pZTr SPH w vT &,GLnZEBROOK' 1985 : 105-112- ,ables 2-4 Several localities, northern Australia, infesting

r. indicus H. Milne Edwards, P. merguiensis de Man and P. longislylus KuboJ. — ANDERSON, 1990 : 290. — Owens

& KOTHLISBERG, 1991 779.

7 Parapenaeon pro x. expansus Bourdon, 1979c : 435 [North Darwin. Northern Territory, Australia, infesting "tiger prawn"

• = Penaeus esculentus Haswell or P. semisulcatus de Haan], F

MAIERIAL EXAMINED. — New Caledonia. Baie Ire, night dive. P. Laboute coll., 15 m, 29 May 1990. Infesting

Metapenaeopsis gaillardi Crosmer. del. of host A. CROSNIER : 1 9 . 1 6 [Both drawn] (MNHN-Ep 818)

a" Makass,ar- CORINDON 2 : stn 201, 01-10.2-S. 117°06.rE. 21 m. fish trawl. 30 October 1980.

Inlesting Metapenaeopsis sinica Liu & Zhong, del. of host A. CROSNIER : 1 9 (MNHN-Ep

«j2Che,"r-flEWESs I' Stn‘ 4' 05:07-7'S- 56°34.0'E, 32 m, 2 September 1980. Infesting Metapenaeopsis faouzii

Ramadan, det. of host A. CROSNIER : 1 9 (MNHN-Ep 823). juvuzu

1 dTMNHNSEpr833)W C°aSt' lnfeSl'ng MeIaPe™eopsis mogiensis consobrina (Nobili), det. of host A. CROSNIER ; 1 9,

Discussion. - Although the present material differs from the types of Parapenaeon expansa in some details

it appears assignable to that species because of the female's body shape, great extension of frontal lamina and coxal

BOPYRIDAE FROM THE TROPICAL 1NDO-PACIFIC

245

plates, deeply digitate barbula and internal ridge, pointed lateral plates and cleft last pleomerc. The males are less

similar, in that the type had a smaller head, no eyes, larger seventh dacytli and a longer pleon. All of the present

material infested species of Metapenaeopsis, while all previous records were of parasites of Penaeus spp., so all of

the present hosts are new records. Parapenaeon expansa is previously known from Madagascar, its type-locality,

but the records from the Strait of Makassar. New Caledonia and the Seychelles are new.

Fig 14 - Parapenaeon expansa Bourdon. 1979. A-K. female; L-Q. male. A, Dorsal view. B, Right side of head

8C Right maxilliped. D. Palp of same. E, Plectron of same. F. Right side of barbula. G. Right oostegite 1. external

vfew H, Same, internal view. I. Right pereopod 1. J. Right pereopod 7. K Pleon in ventral v^iew. L Dorsal v,ew.

M. Ventral view. N, Right antennae. O, Right pereopod 1. P. Right pereopod 7. Q. End of pleon pin ventral vies ^

Scale : 2.00 mm for A-C. F-H, K; 1.00 mm for L, M; 0.52 mm for D. E; 0.31 mm for I, J. O, P. 0.15 mm .or N. Q.

246

J. C. MARKHAM

Parapenaeon secunda Nierstrasz & Brender a Brandis, 1923

Figs 15-16

ParZZZTP SeC,Wdum Nicrstrasz & Brender a Brand.s, 1923 : 67-68, fig. 4a-e [South of Flores Island. Indonesia

infesting Parapenaeus fissurus (Bate)]. — Nierstrasz & Brender a Brandis, 1932 • 93-94 — Piliai 1954 -^1

Bourdon, 1979c : 435; 1981 : 245, 255. rlLLAI’ ■ n- ~

Non Parapenaeon prox. secundum Bourdon. 1979a : 492-495. figs 13-14 = Parapenaeon coarc, alum Bourdon. 1981.

Parapenaeon secunda Nierstrasz & Brender a Brandis, 19^3 A-H female- IN ma|p An . ■

mJS l: g; I fObarbra' ^ ft ~ ^ ^

pereopod 1. M. Left peL,X J' V"W* K-

Scale : 4.0 mm for A-F. I, J; 0.5 mm for G. H; 0.3 mm for K-N.

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

247

MATERIAL EXAMINED. — Philippines. Musorstom 2 : sin 86. near Manila. 50 m, 02 December 1980. Infesting

Metapenaeopsis palmensis (Haswell), del. of hosl A. Crosnier : 1 2 immature [Drawn) (MNHN-Ep 822).

Indonesia. Strait of Makassar. CORINDON 2 : stn 295, 01°26.5S, 117°02.1'E, 54-51 m. Infesting Metapenaeopsis

sinica Liu & Zhong, det. of host A. Crosnier : 1 9. 1 6 [Both drawn) (MNHN).

Discussion. — Although some doubl remains whether all described species of Parapenaeot i are valid and thus

distinguishable, this material seems assignable to P. secunda. The adult female (Fig. 15A-H) shows the body

outline, large coxal plates (two of them serrated), slender and laterally placed maxilliped palp, deeply digitate

barbula, slender posterolateral projection of the first oostegite. and long narrow tubcrculate pleonal appendages of

the holotype. The present male (Fig. 15I-M) is rather more different from the allotype, which had ihe head separate

from the pereon and a pointed pleon. The small immature female (Fig. 16) differs from the holotype in some

details, such as the more nearly circular body outline, the proportionately larger coxal plates, the flat nearly closed

brood pouch and the less extended pleon. Because of these differences, to be expected in an immature female, it is

somewhat difficult to assign with certainty to any species, though it docs appear to belong here. Both the

Philippines and the Strait of Makassar are new localities for P. secunda, and the hosts in the genus

Metapenaeopsis are also new records.

Fig. 16. — Parapenaeon secunda Nierstrasz & Brendcr a Brandis, 1923. female. A. Dorsal view. B. Ventral view.

Scale : 2.0 mm.

SUBFAMILY ATHELGINAE Codreanu, 1956

Pseudostegias setoensis Shiino, 1933

Fig. 17

Pseudostegias setoensis Shiino, 1933 : 290-293. fig. 16 [Seto. Japan, infesting Clibanarius bimaculatus de Haan);

1950: 161-162; 1952 : 35-36; 1958 : 68 |Seto, Wakayama Prefecture, Japan, infesting C. bimaculatus-, and Taiwan,

infesting C. striolatus Dana); 1972 : 9. — LEMOS DE Castro, 1965 : 105-108. — Markham. 1982 : 369-370, 372-

373, 385 [Hong Kong, infesting C. bimaculatus and C. ransoni Forest). — MORTON & MORTON, 1983 : 96. 98, -01,

fig. 7, 5 (6), table 10.2 [Hong Kong, infesting C. bimaculatus and C. striolatus]. — Page. 1985 ; 201. 203.

248

J. C. MARKHAM

Material EXAMINED. — Chesterfield Islands. MUSORSTOM 5 : stn 361, 19°53.50’S, 158.38'10'E, 400 m,

19 October 1986. Infesting "Trizopagurus" sp. (to be described as gen. nov., sp. nov. by J. FOREST), det. of host

J. Forest : 1 2. 1 6 (MNHN-Ep 819).

New Caledonia. Smib 4 : stn DW 67, Ride dc Norfolk. 22°55.1'S, 167°15.6'E, 460 m. 10 March 1989. Infesting

"Trizopagurus" sp. (to be described as gen. nov., sp. nov. by J. FOREST), det. of host J. Forest : 1 $ , 1 d [Both drawn],

(MNHN-Ep 834).

Fig. 17. Pseudoslegias seloensis Shiino, 1933. A-H, female; I-N, male. A. Dorsal view. B. Ventral view. C, Right

maxilliped. D, Lelt side of barbula. E. Right oostegite 1 in external view. F, Same in internal view. G, Right

pereopod 1. H, Right pereopod 7. I, Dorsal view. J, Ventral view. K. Right antennae. L, Left pereopod I. M, Left

pereopod 7. N. End of pleon in ventral view.

Scale ; 4.0 mm for A, B, E. F; 2.0 mm for C. D, 1, J; 1.0 mm for G, H; 0.3 mm for K-N.

Discussion. — In contrast to previously recorded specimens, the female drawn is relatively slightly broader

anleriorly, and ils pleonal lateral plates less extended: and ihc illustrated male has head and pleon both slightly

shorter and no eyes. The other two specimens examined are nearly identical with those illustrated. Chesterfield

BOPYRIDAE FROM THE TROPICAL INDO-PACIFIC

249

Islands and New Caledonia are new localities for Pseiidostegias setoensis. and the host hermit crab, the first not in

the genus Clibanariiis , is also a new record.

SUBFAMILY ENTOPHILINAE Richardson. 1903

Entophilus omnitectus Richardson, 1903

Entophilus omnitectus Richardson, 1903 : 824-826, figs 6-8 [Between islands of Hawaii and Molokai, Hawaii, infesting

Munida normanni Henderson; made type of Subfamily Entophilinae nov.]. — Richardson, 1904 ; 679-681, figs 34-

39 [Reprint of above article). — NlERSTRASZ & Brender a Brandis. 1923 ; 63, 66. — Danforth, 1963 ; 847, 849;

1970a : 27; 1970b ; 462. — Bourdon, 1976 ; 385-391. figs 21-23 [Madagascar, infesting M. incerla Henderson;

redescriptionj; 1979b : 511 [Azores, infesting M. sanctipauli Henderson; mention of unpublished records from

Australia and Philippines). — BOURDON el at.. 1981 : 498-500, 502, fig. 611. — MaRKHAM, 1986 : 144, 148, 156,

fig. 1H. — ADKISON & COLI.ARD, 1990 : 649-654, figs lb-1, 2 [Northern Gulf of Mexico, infesting M. valida Smith

and M. niicroplitlialma A. Milne Edwards).

MATERIAL EXAMINED. — Chesterfield Islands. MUSORSTOM 5 ; stn 365, 19°42.82'S. 150°48.00'E, 710 m,

19 October 1986. Infesting Munida "incerla" Henderson, del. of host K. Baba : 2 9 , 2 6 , (MNHN).

Discussion. — Chesterfield Islands is an important extension of the range of Entophilus omnitectus. which is

now known to infest several species of the galatheid genus Munida in deep waters around the world. The present

material falls within the range of characters reported for this enigmatic species. Upon identifying the host on my

behalf. Dr Baba pointed out that Munida incerla is a name applied to a complex of species yet to be properly

described and separated. Thus it is uncertain whether the present host is the same as that reported for E. omnitectus

at Madagascar by Bourdon (1979b). or what its true identity may ultimately prove to be.

ACKNOWLEDGMENTS

Sinccrest thanks arc extended to Alain CROSNIFR for sorting out the ORSTOM collections and putting them at

my disposal, for arranging for and hosting my visit to the Museum national d'Histoire naturelle, lor identifying

the penaeoid hosts, for helping me find elusive library references, for cataloguing the material and providing other

essential curatorial services and for editing this report. The financial support ol the Institut Fran^ais dc Recherche

Scicntifique pour le Developpcment cn Cooperation is gratefully acknowledged. Other identifications ol hosts were

made by Drs Keiji Baba, Michelle DE SAINT LAURENT. Jacques FOREST, and Lipke HoLTHUIS, the latter also

providing advice on nomenclature. Dr Robert WaSMER passed on the infested Prionocrangon , which had come into

his possession. Dr R. Bourdon and an anonymous reviewer made valuable comments on the manuscript,

including suggestions for revisions of some identifications and more extensive discussions. Mrs W. A. Markham

provided facilities at the Arch Cape Marine Laboratory, of which this is publication number 23.

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JLTATS DES CAMP AGNES MUSORSTOM, VOLUME 12 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 12 — RESULT ATS D

Crustacea Decapoda : Les Metapenaeopsis

indo-ouest-pacifiques

avec un appareil stridulant (Penaeidae)

Alain CROSNIER

Chercheur ORSTOM

Museum national d'Histoire naturelle

Laboratoire de Zoologie (Arthropodes)

61 rue Buffon, 75231 Paris Cedex 05

RESUME

Ce travail, qui est la troisieme et dernicre partie de la revision du genre Metapenaeopsis que nous avons entreprise en

1987, traite des cspcces pourvues d'un appareil stridulant.

De meme que nos precedents travaux, celui-ci a ete base, au depart, sur les nombreuses recoltes faites par les chercheurs

de l'ORSTOM dans l'lndo-Ouest-Pacifique et sur celles faites dans le cadre du programme MUSORSTOM, mene conjointemcnt

par l'ORSTOM et lc Museum national d'Histoire naturelle de Paris, essentiellement aux Philippines, en Indonesie et en

Nouvelle-Caledonie. II s'y est ajoute un materiel important fourni par divers Museums et. en particulier, le National

Museum of Natural History de Washington, qui a mis a notre disposition les recoltes de \"‘ Albatross" faites principale-

ment aux Philippines.

Les specimens, cites dans des publications, out etc reexamines dans toute la inesure du possible.

Parmi les cspcces deja dccritcs, 14 sont reconnues comme valides : M. acclivis (Rathbun, 1902), M. aegyptia Galil &

Golani. 1990. M. barbata (de Haan, 1844). M. crassissima Racek & Dali. 1965. M. dura Kubo, 1949, M.fusca R. J. G.

Manning, 1988, M. Undue R. J . G. Manning, 1988, M. novaeguineae (Haswell, 1879), M. palmensis (Haswell, 1879),

M. rosea Racek & Dali, 1965, M. sinica Liu & Zhong, 1988. M. sinuosa Dali, 1957, M. stridulans (Alcock, 1905).

M. toloensis Hall, 1962.

Les synonymies de M. akayebi (Rathbun. 1902) avec M. barbata et de M. barbeensis Hall, 1962, avec M. palmensis

sont confirmees.

La validite de M. tcliekunovae Starobogatov, 1972, reste en suspens.

Une espece nouvclle, M. parapalniensis. est decrite.

Ce ne sont finalement que 15 especes qui sont etudiees ici. dont une settle est nouvelle. L’originalite de ce travail a en

fait consiste a mettre de l'ordre dans la Literature et a dotiner, esperons nous, une idee claire des differentes especes. II s'est

en effet revelc que, par lc passe, les confusions entre especes ont ete tres nombreuses.

Deux cles d'identification, 1'une pour les males, l'autre pour les femelles sont proposees. Les caractcres facilement

utilisables retenus les rendent, malheureusement, entierement artificiel les et ne font pas ressortir les groupements

CROSNIER. A., 1994. — Crustacea Decapoda : Les Metapenaeopsis indo-ouest-pacifiques avec un appareil stridulant

(Penaeidae). In : A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM, Volume 12. Mem. Mas. naln. Hist, not..

161 : 255-337. Paris ISBN 2-85653-212-8.

256

A. CROSNIER

d'especes. Ces cles sont surtout destinees S guider le lecteur vers l'illustration qui est abondante, etant donne la

complexite des structures des pieces genitales largement utilisees pour distinguer les especes. Les dessins, au trait et

lavis, sont au nombre de 192, repartis en 45 figures.

Des tableaux montrent, de mani&re synoptique, les repartitions bathymetrique et geographique des diverses especes.

L'Australie se revele particuliferement riche en especes, l'oclan Indien pauvre. la Polynesie et les lies Hawaii totalement

ddpourvues.

Dans le cas de l'Australie, on observe deux groupes d'especes, l'un semble purement australien, l'autre composd

d'esp&ces tropicales deferlant par le nord. Un groupe d'origine japonaise, migrant vers le sud, et un groupe indo-

philippin, migrant dans toutes les directions, s'observent egalement. Quelques repartitions geographiques demeurent

difficiles a expliquer, de maniere satisfaisante, avec les donnees actuelles.

Apres cette revision, le genre Melapenaeopsis compte 71 especes ou sous-especes, dont 46 sont indo-ouest-

pacifiques, quatre est-pacifiques, cinq ouest-atlantiques et une est-atlantique.

ABSTRACT

Crustacea Decapoda : The Indo-West Pacific species of Melapenaeopsis with stridulating

organs (Penaeidae).

This is the final work in a trilogy of papers and concludes the revision of the genus Melapenaeopsis which

commenced in 1987. The present study refers to those species of Melapenaeopsis with stridulating organs.

As with the preceding papers, this study is primarily based on numerous specimens collected by ORSTOM scientists

working in the Indo-West-Pacific. Further collections were made in the Philippines, Indonesia and New Caledonia as part

of the Musorstom programme, a joint project established between ORSTOM and the Museum national d'Histoire

naturelle. Paris. This material was supplemented by specimens from "Albatross" stations, principally from the

Philippines, generously made available by the National Museum of Natural History, Washington. Numerous requests were

also made to those institutions holding material cited in publications.

Of the species previously described, 14 are considered valid : M. acclivis (Rathbun, 1902), M. aegyptia Galil &

Golani, 1990, M. barbaia (de Haan, 1844), M. crassissima Racek & Dali, 1965, M. dura Kubo, 1949, M.fusca R. J. G.

Manning, 1988, M. lindae R. J . G. Manning, 1988. M. novaeguineae (Haswell, 1879), M. palmensis (Haswell. 1879),

M. rosea Racek & Dali, 1965, M. sinica Liu & Zhong, 1988, M. sinuosa Dali, 1957, M. stridulans (Alcock. 1905), and

M. toloensis Hall, 1962.

The synonymies of M. akayebi (Rathbun, 1902) with M. barbaia and of M. barbeensis Hall. 1962, with M. palmensis

are confirmed.

The validity of M. ichekunovae Starobogatov, 1972. remains uncertain.

One new species. M. parapalmensis, is described.

Thus 15 valid species are studied here, of which one is new. The aim of this work has been to clarify the taxonomy of

the group and, it is hoped, to give a clear idea of the different species. It has been shown that confusions between species

have actually been very common in the past.

Two identification keys, one for females and another for males, are provided. The keys are supplemented by

192 illustrations arranged in 45 figures. The morphology of the genitalia is highly complex and for this reason these

organs are drawn in various aspects.

Two synoptic tables are given showing the bathymetric and geographical distribution of species. Australia is

particularly rich in species, in contrast to the Indian Ocean, which is poor, and Polynesia and Hawaii which are

completely devoid of Melapenaeopsis species with stridulating organs.

The characters used in both keys are easy to use, but lead to artificial assemblages which do not reflect the natural

groups of species. Melapenaeopsis species with stridulating organs seem to be divided into three natural groups, one

originating from Japan and migrating southwards, a second originating from the Philippines and Indonesia and

dispersing in all directions, and a third, endemic to Australia where members of the other two groups are also found. Some

distributions, however, remain difficult to explain.

Now that the revision of Melapenaeopsis is completed, 71 species and subspecies have been recognized : 46 species

from the Indo-West Pacific; 4 from the East Pacific; 5 from the West Atlantic; and 1 from the East Atlantic.

INTRODUCTION

Ce travail I ait suite a ceux que nous avons publics en 1987 [Les especes indo-ouest-pacifiques d'eau profonde du

genre Melapenaeopsis (Crustacea Decapoda Penaeidae), Bull. Mus. natn. Hist, nat., Paris, sdr. 4, 9, sect. A (2) :

409-453] et en 1991 [Crustacea Decapoda : Les Melapenaeopsis indo-ouest-pacifiques sans appareil stridulant

METAPENAEOPSIS IND0-0UEST-PAC1FIQUES

257

(Penaeidae). Deuxieme partie. In : A. Crosnier (ed.). R6sultats des campagncs Musorstom. Volume 9. Mini.

Mus. natn. Hist, nat ., (A), 152 : 155-297]. Ces deux articles ont etudic l'enscmblc dcs Metapenaeopsis indo-

oucst-pacifiques sans appareil stridulant.

Lc present travail passe en revue les Metapenaeopsis indo-oucst-paciliques munies d'un appareil stridulant. Cet

appareil consiste en une rangee de cretes transversales. situees sur chacunc dcs faces latcralcs dc la carapace, a lcur

tiers infcricur environ, juste en avant de leur bord posterieur. Ces cretes peuvent s'engrener avee lc bord lateral

anterieur correspondant du premier segment abdominal. Lc nombre ct la taille des cretes peuvent varier beaucoup

d'une espbee h l'autre. fournissant ainsi de bons caracteres systematiques. temperes par le fait quTt l'interieur dune

mcme espece on observe des variations non ncgligeables de ces caracteres.

Le materiel 6tudi6. de meme que lors de nos travaux precedents cites plus haut, a etc fourni. au depart, par les

recoltes faites par les chercheurs de l'ORSTOM dans rindo-Oucst-Pacifique. a Madagascar (recoltes du "Vauban"),

aux Seychelles (campagne Reves), en Nouvelle-Caledonie (programme Lagon principalement) et par certaines de

celles faites conjointemcnt par l'ORSTOM et lc Museum national d'Histoire naturelle (campagnes Musorstom 1-

3 aux Philippines, Corindon en Indonesie. en particulier). Nous avons donne dans l'introduction de notre travail

dc 1991 . les references bibliographiques pennettant d'obtenir des details sur ces recoltes et nous n'y reviendrons pas

ici. Nous mentionnerons toutefois qu'en 1991, B. Richer de Forges a publie un travail tres complct sur les

echantillonnages effcctues dans le cadre du programme Lagon auquel le lecteur pourra se reporter utilement.

Du materiel complcmcntairc nous a ete fourni par de nombreux musdums et instituts dont nous donnons ci-

apits la liste avec les abreviations correspondantes, utilises pour indiquer, dans cet ouvrage, les origincs du

materiel examine :

AMS : Australian Museum, Sydney.

BMNH : The Natural History Museum (anciennement British Museum, Natural History), Londres.

MNHN : Museum national d'Histoire naturelle, Paris.

SMF : Natur-Muscum Senckenberg, Francfort/Main.

NMW : Nalurhistorisches Museum. Vienne.

MSNTO : Museo Civico di Storia Naturalc di Torino.

NTM : Northern Territory Museum. Darwin.

NTOU : National Taiwan Ocean University, Keelung.

POLIPI : Pusat Penelitian dan Pengembangan Oseanologi -L1PI. Djakarta.

QM : Queensland Museum. Brisbane.

RMNH : Nationaal Natuurhistorisch Museum (anciennement Rijksmuseum van Natuurlijke Historic), Leiden.

SAM : South African Museum. Le Cap.

UMZC : University Museum of Zoology, Cambridge.

VM : Victoria Museum, Melbourne.

USNM : National Museum of Natural History, Washington.

WAM : Western Australian Museum, Perth.

ZMA : Zoologisch Museum, Amsterdam.

ZMUC : Zoologisk Museum, Copenhague.

MMSU : Musdum dc Zoologie. University de Moscou.

ZIL : Zoological Institute of Russia, St Petersbourg.

Outre les organismes ci-dessus, plusieurs collbgues, dont on trouvera la liste en fin de travail dans le chapitre

remerciemcnts, nous ont envoye des dons de materiel.

L'absence de tout tcchnicien pour aider a l'cnregistrement du materiel examine, nous a conduit & ne plus

enregistrer cc mat6riel, sauf lorsqu'il s'agit de types ou dc specimens figures. Lorsque le materiel dtait deja

enregistre. avant notre etude, ces numeros sont indiques.

L'ensemble de notre materiel est depose au Museum national d'Histoire naturelle. a Paris, It l'exccption de

doubles, envoy^s. dans toute la mesurc du possible, au National Museum of Natural History a Washington.

258

A. CROSNIER

Les references fournies pour chaque cspece ne sauraient etre considerbes commc completes, surtout pour les

especes a large repartititon, souvent abondamment cit6es, mais nous espbrons que toutes les importantes s'y

trouvent.

A quelques rares exceptions prbs (clairement indiqubes), aucune des references citees n'a ete copiee d'un autre

auteur. Comme a noire habitude, nous les avons toutes verifiees d'apres les textes originaux.

Comme dans nos travaux anterieurs, les dimensions des specimens correspondent a la longueur de la carapace

(Lc). mesuree du fond de l'orbite a la panie dorsale du bord postbrieur de la carapace. Lorsque nous mentionnons la

longueur totale (Lt), celle-ci correspond a la distance separant la pointe du rostre de 1'extremite du telson.

Mentionnons, enfin. que les ooms des navires ayant effectues les rbcoltes sont en italiques et entre guillemets.

ETUDE SYSTEMATIQUE

Genre METAPENAEOPSIS Bouvier. 1905

Nous ne reviendrons pas ici sur la definition du genre Meiapenaeopsis. Nous l'avons donnbe dans noire travail

de 1987 et nous y renvoyons le lecteur, afin d'evitcr des repetitions fastidicuses.

De meme. nous ne redonnons pas ici le tableau indiquant la repartition des branchies, epipodites et exopodites

dans lc genre. Nous l'avons publie en 1987 et 1991 et renvoyons le lecteur it l'un de ces deux travaux. Nous

mentionnerons simplement que toutes les espbees traitees ici ont la formule type des Meiapenaeopsis que nous

avons publibe La plcurobranchic antcrovcntrale du segment VII, commc cela est la rbgle dans le genre, cst

toujours reduii: a unc lamcllc plus ou moins developpcc. non divisce et souvent difficile a discerner.

Les diverses espbees de Meiapenaeopsis etudiees ici presentent un aspect tres homogcnc (de meme d'ailleurs que

l'ensemble des Meiapenaeopsis). Afin d'evitcr de continuelles redites dont la monotonie emousse vitc l'attention,

nous avons renoneb it donner des descriptions completes des diverses especes. De meme que dans noire travail de

1991. nous nous contenterons d'insister sur quelques bons caracteres permettant de sbparer les especes :

— le rostre (forme, longueur, nombre de dents);

— la taille de 1'bpine pterygostomienne:

— l'appareil stridulant;

Sierniles ihoraciques

Fig. 1. — Temies utilises pour la description des demiers sternites Ihoraciques chez la femelle.

Source :