zoosystema fait suite,

avec la même tomaison, au Bulletin du

Muséum national d’Histoire naturelle,

4» série, section A, Zoologie.

Rédacteur en chef :

D. Defaye

Conseil éditorial :

S. Gofas (Illustration)

A. Ohler (Nomenclature)

Assistante de rédaction :

F. Kerdoncuff

Corrections-relecture :

H. Bertini

S. Hoffart-Muller

J. Thomas

Comité scientifique :

G. Balvay, INRA, Thonon-les-bains

C. Combes, CNRS, Perpignan

J. Génermont, UPS XI, Orsay

L. Laubier, Aix-Marseille II, Endoume

J. Lebbe, UPMC Paris VI

C. Lévêque, ORSTOM, Paris

B. Salvat, EPHE, Perpignan

M. Sibuet, IFREMER, Brest

A. Thiéry, UAPV, Avignon

J. Vacelet, Aix-Marseille II, Endoume

A. Matsukuma, Kyushu University, Japan

A. Minelli, University of Padova, Italy

P. Ng, University of Singapore

N. I. Platnick, AMNH New York, USA

J. M. Ramos, Universidade Santa Ursula, RJ, Brazil

F. Vuilleumier, AMNH New York, USA

Abonnements pour l’année 1998 (prix HT)

Annual subscription rates 1998 (excluding VAT)

Abonnement général / General subscription : 1 800 FF

zoosystema : 800 FF

adansonia : 500 FF

geodiversitas : 800 FF

zoosystema peut être obtenu par voie d’échange.

Pour toutes informations s’adresser à :

zoosystema may be obtained on an exchange basis.

For further information please Write to:

Service des périodiques et des échanges de la

Bibliothèque centrale du Muséum national

d'Histoire naturelle

38 rue Geoffroy Saint-Hilaire

75005 Paris

Tél. : (33) (0)1 40 79 36 41

Fax : (33) (0)1 40 79 36 56

zoosystema

Éditions scientifiques du Muséum, Paris

Photocopies :

Les Publications Scientifiques du Muséum adhèrent au Centre

Français d’Exploitation du Droit de Copie (CFC), 20 rue des

Grands Augustins, 75006 Paris. Le CFC est membre de

l’International Fédération of Reproduction Rights Organisations

(IFRRO). Aux États-Unis d’Amérique, contacter le Copyright

Clearance Center, 27 Congress Street, Salem, Massachusetts 01970.

Photocopies:

The Scientific Publications of the Muséum adhéré to the Centre

Français d'Exploitation du Droit de Copie (CFC), 20 rue des Grands

Augustins, 75006 Paris. The CFC is a member of International

Fédération of Reproduction Rights Organisations (IFRRO).

In USA, contact the Copyright Clearance Center, 27 Congress Street,

Salem, Massachusetts 01970.

An enigmatic filiform organism in the epicardium

cavity of a new polycitorid ascidian

KEYWORDS

ascidians,

systematics,

parasitology.

MOTS CLÉS

ascidies,

systématique,

parasitologie.

Françoise MONNIOT

Laboratoire de biologie des invertébrés marins et malacologie,

CNRS D 0699, Muséum national d’Histoire naturelle,

55 rue de Buffon, F-75231 Paris cedex 05 (France)

monniot@mnhn.fr

Monniot F. 1998. — An enigmatic filiform organism in the epicardium cavity of a new polyci¬

torid ascidian. Zoosyslema 20 (3) : 429-438.

ABSTRACT

In several samples of the ascidian Eudistoma hospitale n.sp. collected in the

western Pacific Océan and in South Africa, a filiform organism was found

implanted on the heart tissue. It is rolled in balls, occupies a large part of the

ascidian abdominal cavity, and reaches such a length that it bursts through

the branchial sac and finally cornes out at the colony surface with a free tip.

Présent in ail zooids and in larvae, always single, this filament remains of an

unknown nature, despite ail cytochemical and microanalytical methods used

to characterize it. This kind of organism has never been recorded in any

other ascidian.

RÉSUMÉ

Un énigmatique organisme filiforme de la cavité épicardiale d’une nouvelle espè¬

ce d'ascidie polycitoride. Dans tous les spécimens de l’ascidie Eudistoma hospi¬

tale n.sp. récoltés dans le Pacifique occidental et en Afrique du Sud, un

organisme filiforme est implanté dans les tissus du cœur. Le filament est

roulé en pelote et occupe la plus grande partie de la cavité abdominale de son

hôte. 11 atteint une longueur telle qu'il est expulsé dans la cavité branchiale et

que, finalement, son extrémité libre sort de la colonie. Présent dans tous les

zoïdes et les larves, le filament reste d'une nature inconnue malgré les

méthodes de cytologie et de microanalyse utilisées pour le caractériser. Ce

type d’organisme n’avait jamais été rencontré dans une ascidie.

ZOOSYSTEMA • 1998 • 20(3)

429

Monniot F.

INTRODUCTION

An encrusting colonial ascidian shows curious

thin threads Iocatcd inside the zooids (Fig. 1E)

and sometimes coming out to th.e surface

through the oral apertures. This species was col-

lected for the First time at the Chesterfield

Islands (southwest Pacific) in July 1988, and

later in September 1996; it was also collected on

Fiji in 1991 and October 1996, and in February

1994 and 1996, it was found in two widely sepa-

rated stations in South Africa.

Material

Chesterfield Islands (Laboute coll.). Skeleton,

30 m; île Longue, 22 m.

Fiji Islands. Nukubalavu, 10 m, I6°45’06S -

179°47’83W (Coral Reef Reseach Foundation

coll).

South Africa. Sodwana Bay, 19 m (Schleyer

coll.); South Cape coast, intertidal (Glassom

coll.).

Methods

Specimens fixed in 4% formaldéhyde in sea

water were dissected, stained and mounted on

slides for general anatomical observations.

Sections of 7 |im werc stained with hemalum-

picroindigocarmine or toluidine bluc to reveal

the morphology of the tissues. Ultrathin sections

were observed with an Hitachi 600 électron

microscope Dissected filaments were critical-

point dried, coated with gold and examined with

a Jeol JSM 840 microscope.

Several cytochemical réactions werc used (Pearse

1968; Gabe 1968; Ganter & Jolies 1968-1970).

They are:

- the periodic acid Schiff réaction (PAS) for glu-

cids and glycopnneins;

- the Lugol’s iodine reactive for glycogen;

- alcian blue ar PH 0.5 and 2 5 for acidic glucids

- the alloxane-Schifl reaction for proteins;

- the ferrie ferricyanide method (Schmorl’s and

Lillie processes) for thiols and other reducing

groups especially purines and sulphides;

- the Massons argentaffin reaction for phénols

(tunichrome and analogues);

- the coupled tetrazonium reaction of Danielli,

using the fast blue B sait for proteins and phénols;

- the l-(4-chloromercuriphenyldiazo)-2-naphtol

(mercury orange, red sulphydril reagent) and the

2.2'-dihydroxy-6.6dinaphtyl disulphide (DDD)

methods for thiol groups;

- the Lugols sulfuric acid for cellulose;

rhe chlor-zinc-iodine for both chitin and cellu¬

lose;

- the ferrie chlorure-acetic acid for porphyrins.

Elementary microanalysis was performed on

paraffln sections spread on carbon-coated mylar

slides, using a CAMECA MS46 électron micro¬

probe equipped with wavelengrh dispersing spec-

trometers. The diameter ol rhe probe was 1 pm.

Molecular analysis was carried out on similar sec¬

tions using a Raman laser microprobe Dilor

Microdil 28.

RESULTS

The ascidian

Family POLYCITORIDAE Michaelsen, 1904

Genus Eudistoma Caullery, 1909

Eudistoma hospitale n.sp.

Type material. — MNHN A3 Eud 188,

Chesterfield Islands.

Description

Ail samples hâve the same colonial structure:

cash ions or crusts reaching several decimeters

across and up to threc centimeters thick. The

colour in life is uniform through cach colony,

but in only one station varies among black, light

brown, pink or purple (Fig 1A-D), The colony

surface is smooth, shiny, sometimes slightly

undulated The zooid oral openings appear as

small holes arranged in çircles in the center of

which the cloacal apertures open, grouped on

small protrusions (Fig. 1A-D). The tunic is

opaque without any embedded sédiment. The

general consistency is firm with an even more

résistant surface layer. When fixed, the tunic

becomes more translucent and more or less dark

grey, It contains abondant vesicular and pigment

cells, and vanadocytes. The zooids lie perpendi-

cular to the colony surface; the less contracted

430

ZOOSYSTEMA • 1998 • 20(3)

An enigmatic organism in ascidians

Fig. 1. — Eudistoma hospitale n.sp.; A, B, C, three different colours of colonies from the Chesterfield Islands; D, colony from

Sodwana Bay, South Africa; E, a zooid, 1 cm long, with ifs thread stained in red with acidic fuschin.

ZOOSYSTEMA • 1998 • 20(3)

431

Monniot F.

ones easily reach 1 cm in length. The thoracic

body wall is black, especially in the anterior part.

The abdomens are yellowish, the embryos bright

orange, the larvae paler orange. These colours

disappear after fixation.

The zooids ail hâve the same characters, whatever

their geographical origin. Borh siphons are tubu-

lar, edged with six rounded lobes, the cloacal

siphon longer (Fig. 2A, C). The cndostyle is

thick. The thoracic muscles are dense with

numerous longitudinal and transverse fibers.

Generally, sixteen oral tentacles in two orders of

size alternate along two circles. The branchial sac

has anteriorly an imperforated area only seen in

relaxed zooids. An average ot twenty-five elonga-

ted stigmata were counted in the first row.

The abdomen lies below a constriction of the

body. The isodiametric œsophagus is very long

(Fig. 2A). The stomach has a generally round

shape with sometimes, in the largest individuels,

two latéral crests when the gut is empty. The

post stomach is cylindrical, separated from the

mid-intestine by a constriction. The rectum

begins with an annular widening (Fig. 2E, F)

which sometimes forms two crescent-shaped

caeca. After the curve of the gut loop, the intesti¬

ne remains parallel to the œsophagus up to the

level of the second stigmata row, where rhe rec¬

tum opens via a bilobed anus. Beyond the mid-

intestine the rectum has a green colour in life, At

the stomach level, the rectum is covered with

parallel pyloric tubules converging into a short

duct that connects with the posterior part of the

stomach. One or two vascular processes prolong

the abdomen into the tunic.

The hearr forms a J-shaped tube, with unequal

sides, more or less inflated (Fig. 2D-F). The

endothélium of the vessels dtsappears at a short

distance from the heart. On the cardiac tissue,

between the ovary and the base of the gut loop,

there is an extremely long filiform structure, of

unknown nature. This thrrad begins with a coni-

cal root (Figs 2D, E, 3C, D, 4A), then remains

the same diameter along its whole length. Ir is

coiled on itself (Figs 2D-E 4A), and invades the

epicardium cavity (Fig. 1E). Reaching the thorax,

it becomes so voluminous and tightly wound

that it pierces the wall, enters the branchial sac

and finally extends out through the oral siphon.

I 432

The gonads hâve the common structure of the

genus. Fhe testis is made of numerous round

vesicles (Fig. 2A, B); the ovary is located in the

center of the gut loop, below the stomach

(Fig. 2F). In colonies from the Chesterfield

lslands, the zooids are ail male or female; but in

ones front Fiji there is a young ovary amidst the

testis vesicles and embryos are présent in the

cloacal cavity (Fig. 2C). The South African colo¬

nies are simultaneously hermaphroditic.

One or sometimes two embryos are incubated at

the same tinte. Well-deveJoped larvae were found

only in Chesterfield specimens (Fig. 2G, H).

They remain inside the tip of the oviduct until

their release into the sea. The larval trunk is oval,

1 mm long. The tail makes a hall turn around it.

When younger embryos are orange and translu-

cent, the tadpole body wall contains numerous

pale opaque cells. The sensory vesicle has the

usual otolith and ocellus. Three rows of about fif-

teen stigmata apiece are already visible in an incu-

bating larva (Fig. 2H), and an embryonic gut lies

in the posterior part of the body (Fig. 2H).

Anteriorly, the three adhesive papillae, ail in a

line, hâve oval tips. Their thick bases alternate

with four foliose vesicles. In addition, there is a

vesicle applied on the side of the most dorsal

vesicle on each side (Fig. 2FI) In aged larvae, a

short thnead can be seen in transparency amidst

the virellus near the embryonic gut (Fig. 2G).

The presence of this filamentous structure has

been ascertained in histological .sections.

Within the tunic, between functional zooids,

traces of degencrated zooids persist with pièces of

filaments which are only empty tubes, some with

lysed cells. No filaments were found in the tunic

outside the zooids.

Eudistoma hospiiale difters from other species of

the genus only by its larval structure and the

constant presence in ail colonies and ail zooids of

a filiform organism whence the species name.

The fiiamentous structure

A single thread winds about inside the pericardial

cavity' of each zooid (Figs 1 E, 4C). Its conical

basis ralces root on the cardiac lining of the asci-

dian and is exrernally covered by a cellular layer

of the same constitution as the hearts endothé¬

lium (Fig. 3C, D). An aperture allowing a com-

ZOOSYSTEMA • 1998 • 20(3)

Monniot F.

munication between the cardiac cavity and the

inside of the filament is clearly visible in scan-

ning électron microscopy after dissection and cri-

tical point déhydration. Following the basal part

which measures approximatcly 100 pm in dia-

meter, the thread rapidly narrows, and then

remains of a constant dia meter of 20 pm along

its whole length (Figs 3A, 4A, C). At First only

slightly coiled, the thread lengthens during the

zooid’s growth, and becomes a more and more

tightly coiled hall (Figs 1E, 2D, E, 4C) that

invades the whole abdomen and reaches the tho¬

rax. There the filamentous structure is so large

that it breaks the thorax fioor, spreads freely into

the branchial sac, and finally cornes out through

the oral siphon. On the colony surface, we hâve

found unrolied flexible threads reaching 1 cm in

length. The length of the thread of an uncoiled

bail measures about 10 cm.

The structure of the thread comprises three

parts.

The external sheath of pariétal cells

The external sheath of the filament seems to be

made of cells originating in the cardiac endo¬

thélium of the ascidian with which it is in conti-

nuity at the levd of its conical root (Figs 3C, D,

4A). There, the cells are rhick, sphetical, covered

with villosities, with a sphetical nucléus. These

cells progressively Hatten along the internai fila¬

ment the farther they lie from the heart (Fig. 3C,

D). Their nucléus become kidney-shaped

(Fig. 4D, E), long villosities persist (Fig. 4B, D);

and the basal layet is thin. In light microscopy,

the nucléus is strongly stâined, and the reduced

cytopiasm is constituted almost entirely of

microvillosities. In électron microscopy, we esta-

blished that a clear limil exists between this cel¬

lular envelope and the filament linïng itself,

without interpénétrations (Fig. 4E). As the

thread lengthens, its external cells become pro¬

gressively very much flatter and less tightly pac-

ked and seem to be only linked by their

villosities, which cover the whole filament.

The filament lining

The filament has a thick wall which is not seg-

mented and neither interiorly nor exteriorly

ornamented. The external surface is totally

smooth when observed in SEM or TEM

(Figs 3B, 4D-F). The internai surface is granular,

in close contact with the cells inside which seems

responsible for its sécrétion. No stratified struc¬

ture is revealed by TEM in this 2 pm rhick wall

(Fig. 4F). The tube is devoid of cellulose: the

Lugol-sulfuric acid mixing reaction is négative,

while on the contrary, it is positive for the asci¬

dian host-tunic. The presence of chitin seems

ruled ont for the reaction with chlor-zinc-iodine

(also revealing cellulose) is négative. A micro-

analysis in Raman spectroseopy présents a single

peak centered on 1450 cm then the absence of

chitin The hypothèses that the filament is a fun-

gus closely allied to the Trichomycetes is cxcluded

by these resuit.

The internai cells of the filament

The cells of the filament hâve two different

aspects (Fig. 4D); rhey may belong to two diffe¬

rent kinds of cells or represent two States ol évo¬

lution of a same kind of cell, as rhe ratio of the

chromophile cells increases in die distal part of

the filament. However, no transition State was

observed between these two celluiar aspects. In

both cases, the cells hâve a round outline and a

large central nucléus (Fig. 4D, E), thus hclong-

Lng to an eucaryote. The cells are pressed against

each other in the whole length of the filament.

They seem to be bound together and to the wall

by a substance containing microgranules.

Histological srains (Massons trichrome, roiuidine

blue) show a différence between two cellular

aspects (Fig. 4B): cells with a clear homogeneous

cytopiasm with a central nucléus rich in chroma-

ttn, and chromophile cells, slightly larger, whose

cytopiasm contains multiple vacuoles with a

dense content (Fig. 4D, E).

The vacuoles’ contents hâve two peculiar charac-

ters: a strong reactivity to the Lugol treatment

and the coexistence of sulphur and vanadium as

revealed by X-ray microanalysis, The coloration

obtained with Lugol’s reagent is intense and

immédiate; it produces a brown-red tint. So it

difiers from the purple color of animal and végé¬

tal glucids, which, anteriorly the chemically

based techniques, were considered as spécifie of

these compounds. Such anomalies in the results

of the reaction were recorded in the search for

434

ZOOSYSTEMA • 1998 ■ 20(3)

An enigmatic organism in ascidians

glucids in sporozoans and had led to the notion

of “paraglycogen” Grasse (1953). The absence of

reactivity to PAS as Pearse (1968) noted in criti-

cism of this old technique (proteins being able

positively react as well), let us conclude that

vacuole content is not glucidic nor glycoproteic.

Fig. 3. — A. part of a thread bail; B, pariétal sheath disrupted, showing the filament; C, thread root inserted on the heart tissue;

D, detail of the root showing the progressive flattening of the pariétal cells. Scale bars: A, 100 pm; B, D, 10 pm; C, 50 pm.

ZOOSYSTEMA • 1998 • 20(3)

435

Monniot F.

The coexistence of sulphur and vanadium has

often been demonstrated in ascidian vanado-

cytes. We hâve previously shown (Martoja et al

1994) that only a part of the sulphur is linked to

the vanadium; rhis occurs in only two kinds of

cells, compartment cells and signet ring cells,

while in morula cells the vanadium is linked to

oxygen and coexists with a tunichrome. The éli¬

mination ol' vanadium by normal hydrochloric

acid or Schmorl’s reagent, though maintaining

the sulphur, does not unmask thiol groups (néga¬

tives reactions with ferrie ferricyanure or red-sulf-

hydril reagent). So it is not possible to equate the

vacuole content of the filament cells with that of

the vacuoles of the signet ring cells and compart¬

ment cells. Massons argentaffm reaction being

also négative, the presence of a tunichrome is

also excluded, which rules out a similarity with

the host’s morula cells. The vanadium ligand

seems very different from that of the host. The

hypothesis of a porphyrin-vanadium compound

is also excluded because protracted treatment

with chloroform (24 h), the best solvent of por-

phyrins, has no effect on the composition of the

vacuolar content, and the ferrie chlorure-acetic

acid reaction is négative. The problem of the

vanadium ligand remains unsolved.

The samples were fixed in the field with seawa-

ter-formalin. which does not allow for a more

précisé or advaneed histological examination

than what we hâve doue.

DISCUSSION

It is difficult to détermine whether this curious

filament represents a parasite or a symbiont. At

first, the hypothesis of a peculiar organ, excretory

for example, in a particular ascidian species was

considered but rapidly abandoned; the filament

develops in a natural cavity without any link

with its wall, then bursts through the branchial

sac and even extends a lengthy free extremity

outside the ascidian. The cells of the filament

hâve not the usual chatucters of excretory cells.

The coating of ascidian cells along the thread

more likely suggests a hosfs reaction against a

foreign organism, analogous to the cysts that

ascidians make around endovascular parasitic

copepods (Dudley 1968). The absence of any

differentiated tissues that compose the filament

excludes the possibility of a parasitic copepod,

even a very modified one. Assignaient to a parti¬

cular metazoaii phylum remains unseraed. The

large central nucléus and the thin cell membrane

exclude, as well, bacteria, procyanophytes and

algae. No known fungi correspond to this struc¬

ture, nor does the Chemical composition of the

filament support such an affiliation.

Influence of the filament on its host

The zooids of Eudistoma hospitale do not show

any peculiarity which may be due to the influen¬

ce of the filament, for example no morphological

deformation, no castration. The zooids feed nor-

mally, their gonads are well-developed, their

embryos reach full maturation without monstro-

sities. Asexual reproduction is not affected either,

as the colonies teaçh very large sizes compared to

other species of the sanie genus.

Transmission mode

We do not know if the organism which forms

the filament goes through its life cycle a free

living stage or obligatorily housed at ail times in

a host. We only notice that the association begins

from the larval stage of the ascidian, as a short

filament is already présent in the larva that is

being incubated in the oviduct, before its release

into the sca. "Infection" thus may take place in

the mother zooid before the young tadpole

begins to elaborate its tunic, perhaps at the level

of the cells of the egg envelope. This would

imply that, in addition to embedded cells inside

a filament, isolated cells or gamètes would exist

and circulate widi die hosfs blond, equallv able

to infect eggs and the regenerating buds produ-

ced by strobilation responsible for the asexual

growth of the colony. Another hypothesis would

take into account the émission of the filament

extremity at the colony surface, allowing cells to

be iiberated in sea water and ingested to infest

newly formed zooids. A direct transmission

through the colonial tunic from one zooid to

another is also a possibility.

Presently nothing allows us to explain why there

is always just a single thread per zooid and we

hâve found no gametes.

436

ZOOSYSTEMA • 1998 • 20(3)

An enigmatic organism in ascidians

Fig. 4. — A, B. cardiac extremity of a thread in light microscopy (toluidine blue). C, sections of a thread bail contained in the pericar-

dium cavity occupying the most part ot the abdominal section of a zooid. D, E, sections of a thread in TEM showing pariétal cells with

elongated nucléus and microvillies, the tube wall, the internai granular and vacuolated cells. F, detail of the filament wall. Scale bars:

A, 50 pm; B, 10 pm; C, 100 pm; D, 5 pm; E, 2 pm; F, 1 pm.

ZOOSYSTEMA • 1998 • 20(3)

Monniot F.

Other commensals, symbionts or parasites

KNOWN IN ASCIDIANS

Organisms living in association with ascidians

were reviewed by Monniot (1990). They are very

numerous: Virales, Bacteria, Flagellata,

Trichontonadina, Rhizopoda, Gregarina,

Coccidia, Haplosporidia, Ciliophora, Nephro-

myces, Algae, Cnidaria, Ctenaria, Turbellaria,

Annelida, Bryozoa, Nemertina, Mollusca, many

Crustacea and especially copepods, and also

Pisces.

Aknowledgements

I hâve greatly benefited from the precious know¬

ledge of R. Martoja in cytology, and I thank him

as well for the cytochemistry and microanalyses

he has performed as for the thoughtful discus¬

sions we had during this study. I am much

indebted to T. Newberry for the english language

corrections.

REFERENCES

Dudley P. 1968. — A light and électron microscopie

study of tissue interactions between a parasitic

copepod Scolecod.es buntsmani (Henderson) and its

host ascidian, Styela gibsii (Stimpson). Journal of

Morphology 124 (3): 263-281.

Gabe M. 1968. — Techniques histologiques. Masson

ed., Paris, 1113 p.

Ganter P. & Jolies G. 1969-1970. —- Histochimie

normale et pathologique. Gauthier-Villars, Paris,

1904 p.

Grasse P. P. 1953. — Sous-embranchement des

Sporozoaires, volume 1 (2), in Grassé P. P. (ed.).

Traité de zoologie. Masson ed., Paris, 545 p.

Martoja R., Gouzerh P. & Monniot F. 1994. —

Cytochemical studies of vanadium, tunichromes

and related substances in ascidians: possible biolo-

gical signifïcance. Oceanography and Marine Biology

32:531-556.

Monniot C, 1990. — Diseases of Urochordata, volu¬

me 3: 569-636, in Kinne O. (ed.), Diseases of ma¬

rine animais. Biologische Anstalt Helgoland,

Hamburg.

Pearse A. G. E. 1968. — Histochemistry, theoretical

andapplied. Churchill, London, 1518 p.

Submitted on 10 February 1998;

accepted on 6 April 1998.

438

ZOOSYSTEMA • 1998 • 20(3)

Two new species of the genus Ophiuraster

(Ophiurinae, Ophiuroidea, Echinodermata)

from French collections and some remarks

on the genus

Nina M. LITVINOVA

Institute of Oceanology P. P. Shirshov, Russian Academy of Sciences,

Nakhimovsky Prospect 36, 117851 Moscow (Russia)

stal@glas.apc.org

Litvinova N. M. 1998 — Two new species of the genus Ophiuraster (Ophiurinae,

Ophiuroidea, Echinodermata) from French collections and some remarks on the genus.

Zoosystema 20 (3) : 439-444.

KEYWORDS

Ophiuroidea,

Ophiurinae,

Ophiuraster,

O. belyaevi n.sp.,

O. patersoni n.sp.,

systematics.

ABSTRACT

Two new species of the poorly known genus Ophiuraster (Ophiurinae,

Ophiuroidea, Echinodermata), O. belyaevi n.sp. from Kerguelen Islands and

O. patersoni n.sp. from Bay of Biscay are described on the basis of the collec¬

tion of the Muséum national d’Histoire naturelle, Paris. The genus is charac-

terized by a combination of plesiomorphic and apomorphic fearures. The

distribution of these species is shown on a map.

MOTS CLÉS

Ophiuropidea,

Ophiurinae,

Ophiuraster,

O. belyaevi n.sp.,

O. patersoni n.sp.,

systématique.

RÉSUMÉ

Deux nouvelles espèces du genre Ophiuraster (Ophiurinae, Ophiuroidea,

Echinodermata) de Li collection du Muséum natioml d’Histoire naturelle , Paris

et quelques remarques à propos du genre. Deux espèces nouvelles du genre

O phi u ras ter (Ophiurinae, Ophiuroidea, Echinodermata) sont décrites à partir

des collections du Muséum national d’Histoire naturelle, Paris : O belyaevi

n.sp. des îles Kerguelen et O. patersoni n.sp. du golfe de Gascogne. Le genre

est caractérisé par la combinaison de caractères plésiomorphes et apo-

morphes. La carte de la répartition de ce genre, très peu connu, est donnée.

ZOOSYSTEMA • 1998 • 20(3)

Litvinova N. M.

INTRODUCTION

The genus Ophiuraster was established by H. L.

Clark (1939). It includes very small britde stars,

having disks thac are poorly differentiated from

the short and wide arms. Large, closely connec-

ted plates border the dorsal and venrral margins

of the disk As a resuit the brittle stars of this

genus resemble some sea stars of the Family

Goniasteridae. Only two species of this genus

hâve previously been described each represented

by only one specimen. I found two undescribed

species of this genus in the collection of the

Laboratoire de Biologie des Invertébrés marins et

Malacologie of the Muséum national d’Histoire

naturelle, Paris. The unusual characters of these

species broaden our understanding of this genus.

The genus Ophiuraster includes now four species:

O. perissus Clark, 1939; O symmetricus Fell,

1958; O. belyaevi n.sp.; O patersoni n.sp. The

map of distribution of the genus is given (Fig. 1).

Order OPHIURIDAE

Suborder CHILOPHIURINA

Family OPHIURIDAE

Subfamily OPHIURINAE

Ophiuraster belyaevi n.sp.

(Fig. 2)

Holotype. — One specimen, (MNHN EcOs 5839)

collected off Kerguelen Isiands, Marion-Dufresne MD-

04 cruise, stn 1-106, CP 258, 48°43’5"S - 71°06’5”E,

925 m, 13.III.1975.

Etymolggy. — The species is named to honour my

teacher, late Proféssor Georgy Michailovich Belyaev, a

specialist of Echinodermatae.

DlFFEREN MAL DIAGNOStS — The new species is dis-

tinguished by the absence of dorsal-arm plates and

arm spines, vertebrae covered by transparent sldn, and

a large terminal plate présent at the tip of each arm.

The new species is more closely related to O. patersoni

described below than to O. perissus and O. sym¬

metricus.

Description

The disk diameter is 3 mm, and the arms are

short and wide, 2.5 mm in length with a basal

width of 1.3 mm. The center of the dorsal surfa¬

ce of the disk is elevated and covered by primary

plates: the centrodorsal plate is divided into three

small irregularly shaped plates, which are in

contact with each other The radial plates are

large, quadrangular and irregularly shaped; one is

divided into several pièces by radial fissures. A

small interradial plate lies distal to each pair of

radial plates, Ail plates hâve a porous surface and

irregular edges.

The marginal part of the dorsal surface disk is

depressed and covered with skin in which the

radial shields are embedded, The radial shields

are longer than broad and separated by a srrip of

integumenr In each interradius, the edge of the

disk is occupied by two quadrangular plates,

which meet interradially. The same plates extend

to the ventral surface of the disk. They are very

large, quadrangular, closely connected and occupy

the entire périphéries of the ventral interradius.

The arms appear to be shorter than diameter of

the disk. They are wide at their base and connec¬

ted tightly with the disk There are no dorsal arm

plates. There are six arm joints beyond the edge

of the disk, each of which bears protruding laté¬

ral arm plates. The latéral plates of each segment

are widely separated from each other by a srrip of

thin tntegument through which the vertebrae

can be seen. The latéral plates of successive seg¬

ments contact each other. There are no spines in

the arms. The end of each arm is occupied by a

large widened heart-shaped terminal plate, There

are two very small pointed protrusions in the dis¬

tal end of it that are rather hard to notice. The

ventral arm plates are présent only in four seg¬

ments of the arm. The first ventral arm plate is

large and rounded. Three distal ones are minute

and deeply depressed. The ventral arm plates are

widely separated from each other The second

oral tentacle pore opens outside the mouth, and

tentacle pores are présent throughout the length

of the arm. They are small and covered with thin

skin that lias a central hole for the tentacles.

There are neither tentacle scales nor oral shields.

The distal part of adorai shield of one of interra¬

dius is elevated and bears a large water-pore. The

adorai shields are large and irregularly quadran¬

gular. The oral plates are large and hâve one to

three ill-defined oral papillae, of which the apical

ZOOSYSTEMA • 1998 • 20(3)

Two new Ophiuroidea species

papillae are larger. In the deprh of the mouth a

tooth-like apical papilla can be seen. There is a

minute bursal slit in one of interradius.

Ophiuraster patersoni n.sp.

(Fig- 3)

Holotype. — One specimen (MNHN EcOs4895)

collected in Bay of Biscay, /(un Charcot BJOGAS VI,

Stn 6, DS86, 44°05’N - 04°19'W, 1950 m.

Etymoi.OGY. — The species is named to honour Dr

Gordon Paterson who originally identified this speci¬

men up to genus.

Differential DIAGNOSIS. — The new species closely

resembles O. belyaevi n.sp., differing from it by die

absence radial shields ancl adorai plates, by the small

terminal plate, and in the presence of onlv primary

plates on the disk, lt is possible that the only specimen

available is not adult and additional material could

make the diagnosis more précisé.

Description

Disk is fiat, diameter 1.9 mm. Arms and disk are

not well-differentiated and the animal at first

sight resembles a sea-star. The dorsal surface of

the disk is covered wirh transparent skin and

large, centrally situated, primary plates wirh a

porous appearance. The centrodorsal plate is

dividcd transversely in two unequal parts. The

five radial plates are larger and irregularly roun-

ded. There are no radial shields and the latéral

plates of the first segments of arms take their

place. Two large, closely connected marginal

plates make contact interradially and extend to

the ventral side of the disk.

The arms are short and consist of five segments.

The arm length is 1.4 mm; the basal width

0.6 mm. There arc no dorsal plates in the arms

but only large and swolien latéral ones. The laté¬

ral plates of each segment are separated on dorsal

side of arm by a strip of transparent integument

through which the vertebrae are seen. The first

pair of latéral plates is small and situated within a

range of the disk close to its marginal plates.

They could be easily mistaken for radial shields

but there are the vertebrae, which can be seen

through the skin. The distal latéral places are lar¬

ger and more swoilen, meeting broadly above

and below the arm.

The terminal plates are large; each bears two to

Fig. 1 — Distribution of the species of the genus Ophiuraster, ■, O. perissus (2312 m); ♦, O. symmetricus (720 m); ▲, O. belyaevi

(925 m); •, O. patersoni (1950 m).

ZOOSYSTEMA • 1998 • 20(3)

441

Litvinova N. M.

four very small pointed protrusions in the distal The long oral plates hâve small and ill-defined

end. There are no arm spines. The first ventral oral papillae. The apical papillae are largest and

plate is large and rounded, the two distal ones are similar to the teeth situated above them. The

very small and deeply depressed. There are no second oral teutacie pore opens outside the

oral shields or adorai shields. In the place of the mouth. The small tentacle pores are présent on

oral plates, there are diamond-shaped patches of the full length of the arm. There are no tentacle

thin integument. scales. The ventral interradial side of the disk is

Two new Ophiuroidea species

occupied by two large marginal plates, which But, because there only was one specimen of this

meet interradially. No génital plates and bursal species, it was not possible to extract and identify

slits were évident. this animal.

Remarks Discussion

A well-preserved crustacea, occupying the whole There are some contradictions in the interpreta-

stomach, is seen through the semi-open mouth. tion by different authors of some morphological

Litvinova N. M.

features of species of this genus. In O. perissus,

Clark (1939) considered the large marginal

plates occupying the whole margin of the disk to

be radial shields. Fell (1958) supposed that these

plates in O. symmetricus were the broadened laté¬

ral arm plates. On the basis of these two new

species both described from only one specimen, I

propose that the plates occupying dorsal and

ventral interradial surface are considered as mar¬

ginal intcrradial plates of the disk.

H. L. Clark (1939) had noticed that the genus

Ophiuraster is probably closely relutcd to the

genus Ophiomastuv, Matsumoto (1917) and

Vadon (1991) considered that généra similar to

Ophiomastus are paedomoiphtc. The species of

the genus display a combination, of both paedo-

morphic and specialized characters. On the one

hand the new species hâve some characters that

are typical for juvénile ophiuroids: (T) die arms

are extremely short and consist of fevv segments;

(2) the arm spines are absent; (3) the oral shields

and adorai plates (O. patersonî) and tentacle

scales are also absent; (4) the structure of oral

papillae is primitive; (5) the génital cleft is not

noticeable; (6) the body size is small.

On the other hand, the new species hâve charac¬

ters that constitute a specialization among

Ophiurinae: (1) the presence of skin covered

patches on the disk and arms; (2) the presence of

a large terminal plate; (3) the wide séparation bet-

ween the latéral plates of each arm segment on

the dorsal side; (4) the absence of dorsal plates on

the arms; (5) the presence of thin skin in the

place of the dorsal plates through which the ver-

tebrae can be seen; (6) the presence of widened

cap-like terminal plate resemblîng those plates

that were found in Ophiambix meteoris Bartsh,

1983, a représentative of an extremely specialized

genus. The absence of dorsal plates and their

replacement by the skin are interpretated as a

resorbtion of the plates (Hotchkiss 1993).

On rhe basis of their plesiomorphit and apomor-

phic characteristics the Ophiuraster species hâve

clear affiniries to the Ophiurinae such as

Ophiambix , Astrophiura and Ophiophycis , which

superficially resemble sea-stars. But this similari-

ty with asteroids has been reached by different

ways in different généra.

Acknowledgements

I would like to thanks the members of the MD-04

and BIOGAS IV expéditions, especially Alain

Guille and Lucien Laubier, who collecfed this

material. I am very grateful to Nadia Ameziane,

Alain Crosnier, Bernard Métivier, Danièle

Dondon for their friendliness, hclp and for giving

me an opportunity to work in the Muséum natio¬

nal d’Histoire naturelle, Paris and to Kir Nesis for

reading and commenung on draft of this manus-

cript.

REFERENCES

Clark H. L. 1914. — Growth-changes in brittle-stars.

Papers front the Tortugas Laboratory of the Carnegie

Institution of Washington 5: 91-126.

— 1939. — Ophiuroidea. Brhish Muséum (Natural

History). John Murray Expédition 6 (9): 1 -136.

Fell H. B. 1958 — Dcep-Sea Echinoderms of New

Zealand. Zoology Publication front Victoria

Universiry of Wellington 24; 1-40.

Huichkiss F. H. 1993. — A new devonian Ophiuroid

(Echinodermata: Oegophiuridae) from New York

State and its bearing on the Origin of Ophiuroid

Upper arm plates. Proceedings of the Biological

Society of Washington 106 (1); 63-84.

Matsumoto I L 1917, — A monograph of Japanese

Ophiuroidea arranged accotding to a new classifica¬

tion. Journal of the College of Science, Impérial

Universiry, Tokyo 38, 408 p.

Vadon C, 1990, — Ophiozonella tiovaecaledoniae n.sp.

(Ophiuroidea, Echinodermata); description, onto-

geny and phyletic position. Journal of Natural

Histoty 24: 165-179.

Submitted on 26 May 1997;

accepted on 8 December 1997.

444

ZOOSYSTEMA • 1998 • 20(3)

Une nouvelle espèce de Molineus

(Nematoda, Trichostrongylina, Molineoidea),

parasite d’un Primate sud-américain

Marie-Claude DURETTE-DESSET & Matthieu CORVIONE

Laboratoire de Biologie parasitaire, Protozoologie, Helminthologie,

CNRS et Laboratoire de Protozoologie et Parasitologie comparée,

École Pratique des Hautes Études, Muséum national d'Histoire naturelle,

61, rue de Buffon, F-75231 Paris cedex 05 (France)

mcdd@cimrs1 .mnhn.fr

Durette-Desset M.-C. & Corvione M. 1998 — Une nouvelle espèce de Molineus

(Nematoda, Trichostrongylina, Molineoidea), parasite d’un Primate sud-américain.

Zoosystema 20 (3) : 445-450.

RÉSUMÉ

Molineus midas n.sp., parasite de l’intestin grêle de Saguinus midas (Primate,

Hapalidae) en Guyane française est décrite ici. Les nombreuses espèces du

genre, décrites chez les Carnivores dans le monde entier, ont trois caractères

primitifs fréquents chez les Strongylina, mais rares chez les Trichostron¬

gylina, à savoir : une bourse caudale de type 2-1-2, des côtes 4 courtes et un

sillon transversal au niveau du pore excréteur. L un ou l’autre de ces carac¬

tères manque chez les espèces parasites de Singes américains ou chez les

espèces géographiquement proches de la Guyane. Ces trois caractères sont

absents chez la nouvelle espèce.

ABSTRACT

A new Molineus species (Nematoda , Trichostrongylina, Molineoidea), parasite

ofa South American primate. Description of Molineus midas n.sp., a parasite

of the small intestine of Saguinus midas (Primate, Hapalidae) from French

Guyana. The many species of the genus, parasites of Carnivora throughout

the world, hâve three primitive characlers frequently occurting in

Strongylina but rarely in Trichostrongylina. Thèse characters are the follo-

wing: a pattern bursa of 2-1 -2, ravs 4 short and a cervical groove at the level

of the excretorv pore. One or other of thèse characters is lacking in the spe¬

cies parasitic in American primates or in the species geographically close to

Guiana. These three characters are absent in the new species.

KEYWORDS

Molineus midas n.sp.

Nematoda,

T richostrongylina,

Molineoidea,

Saguinus,

French Guiana.

MOTS CLÉS

Molineus midas n.sp.,

Nematoda,

Trichostrongylina,

Molineoidea,

Saguinus,

Guyane française.

ZOOSYSTEMA • 1998 • 20(3)

445

Durette-Desset M.-C. & Corvione M.

INTRODUCTION

Le genre Molineus, créé par Cameron (1923) et

redéfini par Durette-Desset & Chabaud

(1981b), se rencontre chez des Carnivores dans le

monde entier, saul l’Australie, et chez des

Primates néotropicaux. Il est intéressant car il est

morphologiquement très proche du genre

Osuialdocruzia Travassos, 1937, parasite

d’Amphibiens et de Reptiles. La plupart des

espèces montrent en effet des caractères

archaïques qui se rencontrent chez de nombreux

Strongylida, mais très rarement chez les Tricho-

strongylina.

L’espèce trouvée chez un Primate Hapalidae

Saguinus midis (Linné. 1758) en Guyane française

et décrite ci-dessous, est originale du fait qu’elle

ne possède aucun de ces caractères primitifs.

MATÉRIEL ET MÉTHODES

Le matériel a été étudié selon les méthodes clas¬

siques et éclairci dans le lactophénol si nécessaire.

La nomenclature utilisée au-dessus du groupe

famille est celle de Durette-Desset & Chabaud

(1995). Le synlophe est étudié selon la méthode

de Durette-Desset (1985). La nomenclature uti¬

lisée pour l’étude de la bourse caudale est celle de

Durette-Desset & Chabaud (1981a).

Les spécimens-types ont été déposés dans les col¬

lections du Muséum national d’Histoire natu¬

relle, Paris (MNHN).

SYSTÉMATIQUE

Genre Molineus Cameron, 1923

Molineus midas n.sp.

(Fig. 1)

Matériel-type. — â holotype, 9 allotype

(MNHN1 95KHa), 3 d Ô et 3 2 9 paratypes

(MNHN195KHb).

LOCALITÉ-TYPE. — Guyane française, rivière Aratay, à

80 kms de la côte, 19.XI.1980.

Hôte. — Saguinus midas (Linné, 1758) (Primate,

Simioidea).

Localisation. — Intestin grêle.

Autre MATERIEL étudié. — 7 d d, 16 9 9, 6 parties

postérieures et 2 parties antérieures, l’ensemble

MNHN196KH. Même espèce-hôte, même région,

meme date de récolte.

Description

Trichostrongle de taille moyenne dont le corps

est déroulé. Pore excréteur et deirides situés juste

en arrière de l’anneau nerveux. Absence de sillon

excréteur. Vésicule céphalique présente.

Synlophe (étudié chez un mâle et une femelle)

Chez les deux sexes, le corps est parcouru longi¬

tudinalement par quatorze crêtes cuticulaires

longitudinales dépourvues de soutien chicinoïde.

En coupe transversale, au milieu du corps, on

compte cinq crêtes ventrales, cinq dorsales, deux

latérales gauches et deux latérales droites

(Fig, 1D, E). Les crêtes sont de petite taille et

orientées perpendiculairement à la paroi du

corps. Elles prennent naissance en arrière de la

vésicule céphalique et se terminent en avant de la

bourse caudale chez le mâle et à environ 60 pm

en avant de la pointe caudale chez la femelle.

Mâle holotype

Long de 3200 pm sur 65 pm de large dans sa

parrie moyenne. Vésicule céphalique haute de

45 pm sur 25 pm de large. Anneau nerveux, pore

excréteur et deirides situés respectivement à

140 pm, 160 pm et 170 pm de l’apex. Œso¬

phage long de 340 pm (Fig. 1 A).

Bourse caudale symétrique de type 2-1-2 avec

une ornementation épineuse sur les lobes laté¬

raux, principalement entre la paroi du corps et

les côtes 6. Papilles prébursales non observées.

Côtes 4 très courtes. Leurs extrémités sont plus

proches de celles des côtes 3 que de celles des

côtes 5. Côtes 8 naissant presqu'à la racine de la

côte dorsale ; côte dorsale divisée dans son tiers

distal en deux branches, une branche externe

(côte 9) et une branche interne qui se divise dis-

talement en deux rameaux (côte 10 et plias-

rnides). Cône génital peu développé, portant sur

sa lèvre antérieure la papille zéro arrondie et sur

sa lèvre postérieure deux minuscules papilles 7

(Fig- IJ).

446

ZOOSYSTEMA • 1998 • 20(3)

Un nouveau Molineus (Nematoda) chez un Primate de Guyane

|jp T*

Fig. 1. — Molineus midas n.sp. ; A, <5, extrémité antérieure, vue latérale droite ; B, 9, extrémité postérieure, vue latérale droite ;

C, 9, queue, vue latérale droite ; D, E, synlophe ; D, 9, au milieu du corps ; E, <5, au milieu du corps ; F, d, gubemaculum, vue laté¬

rale gauche ; G-l, d, spiculé gauche disséqué ; G, vue dorsale ; H, vue externo-latérale ; I, vue ventrale ; J, K, bourse caudale ;

J, vue ventrale ; K, vue latérale droite. Les coupes sont orientées comme la figure D : d, dos ; v, ventre ; g, gauche ; dr, droite.

Échelles : A, C, 50 pm ; B, 60 pm ; D-K, 30 pm.

Durette-Desset M.-C. & Corvione M.

Spiculés longs de 120 pm, divisés en trois

branches : une branche externo-latérale et deux

branches internes. Les extrémités des branches

externo-latérale et interno-ventrale sont pointues,

celle de la branche interno-dorsale est en forme

de marteau (Fig. 1G-I). Gubernaculum inctfrvé,

haut de 70 pm en vue latérale (Fig. 1F).

Dimensions des trois mâles paratypes, longueur :

3250-3450-3600 pm, largeur 70-70-60 pm ;

vésicule céphalique haute de 45-50-55 sur

30-30-30 pm de large ; anneau nerveux, pore

excréteur et deirides situés respectivement à

130-165-140 pm, 140-175-150 pm et 145-190-

160-pm de l’apex ; oesophage long de 340-370-

340-pm ; spiculés longs de 135-135-120 pm et

gubernaculum haut de 70-70-70 pm.

Femelle allotype

Longue de 3700 pm. large de 70 pm dans sa par¬

tie moyenne. Vésicule céphalique haute de

50 pm sur 30 pm de large. Anneau nerveux, pore

excréteur et deirides situés respectivement à 150,

170 et 190 pm de l’apex. Œsophage long de

410 pm.

Didelphie : vulve s’ouvrant â 660 pm de la poin¬

te caudale, soit au sixième postérieur du corps.

Vagina vera de 20 pm partageant le vestibule

long de 270 pm en deux parties égales. Branche

génitale antérieure : sphincter, trompe et branche

utérine longs respectivement de 30, 20 et

480 pm. Branche génitale postérieure : sphincter,

trompe et branche utérine longs respectivement

de 23, 20 et 300 pm. Branche utérine antérieure

contenant six œufs et postérieure quatre

(Fig. IB). Œuts au stade morula, hauts de 50 pm

sur 30 pm de large. Queue longue de 60 pm avec

une pointe caudale de 12 pm (Fig. IC).

Dimensions des trois femelles paratypes, lon¬

gueur : 3750-3950-3900 pm ; largeur 70-70-

70 pm ; vésicule céphalique haute de 50-45-50

sur 30-30-30 pm de large ; anneau nerveux, pore

excréteur et deirides situés respectivement à

140-140-135 pm, 160-165-150 pm et

160-170-165 pm de l’apex ; œsophage long de

400-370-350 pm ; vulve s’ouvrant à 660-770-

750 pm de la pointe caudale ; ovéjecteur :

321-357-346 pm ; branche utérine antérieure :

470-410-550 pm avec 6-3-3 œufs ; branche uté¬

rine postérieure : 330-350-330 pm avec

4-1-3 œufs ; queue longue de 70-80-80 pm avec

une pointe caudale de 12-12-12 pm.

Discussion

Les spécimens décrits ci-dessus appartiennent au

genre Molineus Cameron, 1923 (Molineoidea)

dont ils possèdent les principaux caractères : syn-

lophe avec crêtes orientées perpendiculairement à

la paroi du corps, bourse caudale avec côtes 4

courtes, côtes 2 et 3 d’une part, côtes 5 et 6 de

l’autre, proches et parallèles, spiculés courts et

épais, femelle didelphe avec vestibule allongé et

trompe très courte.

Les spécimens du Saguinus se distinguent immé¬

diatement de toutes les autres espèces du genre

pat des côtes 8 longues, atteignant le bord de la

bourse caudale. Il s’agit donc d’une nouvelle

espèce que nous nommons Molineus mit/as n.sp.,

en référence au nom d’espèce de l’hôte.

Trois caractères primitifs sont présents chez de

nombreux Strongylina et disparaissent chez

presque tous les Trichostrongylina :

1. La bourse caudale de type 2-3, c’est-à-dire

avec les côtes 2 et 3 constituant un groupe séparé

du groupe formé par l’ensemble des côtes 4, 5 et

6. L’évolution se fait par une migration des

papilles 4 vers l’avant, ce qui conduit à une bour¬

se caudale de type 2-1-2, puis finalement à un

type 3-2 (voir Durette-Desset & Chabaud,

1981a);

2. Les côtes 4 très courtes se terminant loin du

bord de la bourse caudale ;

3. Le pore excréteur s’ouvrant au fond d’un pro¬

fond sillon transversal.

Il est intéressant de suivre l'évolution de ces trois

caractères dans le genre Molineus selon la réparti¬

tion géographique et selon l’hôte.

Neuf espèces sont connues en zone holarctique :

M. patens (Dujardin, 1845) ; M. europaeus

Zunker, 1928 ; Al. petrovi Durette-Desset et

Pesson, 1987 (= AI. patens sensu Petrov, 1928) ;

AI. americamts Sprehn, 1932 ; AI. barbatus

Chandler, 1942 ; AI. mustelae Schmidt, 1965 ;

AI. samueli Platt et Pence, 1981 ; M. legerae

Durette-Desser et Pesson, 1987 et M. springsmi-

thi yayeyamanus Hasegawa, 1989. À l’exception

de M. petrovi chez laquelle le sillon excréteur

n’est pas décrit et de M. americanus insuffisam-

448

ZOOSYSTEMA • 1998 • 20(3)

Un nouveau Molineus (Nematoda) chez un Primate de Guyane

ment connue, toutes les espèces possèdent les

trois caractères primitifs réunis.

Trois espèces sont connues en zone éthiopienne :

Al. genettae (Cameron, 1927) ; AI. cynictis cynictis

(Le Roux, 1933) ; A 1. cynictis thosi Tmncy, 1970.

Elles possèdent aussi les trois caractères primitifs

réunis.

Les trois espèces orientales, Al. planicipitis

(Cameron, 1928), Al. springsmitbi lnglis et

Ogden, 1965 et AI. inglisi Durette-Desset et

Pcsson, 1987 (= AI. païens sensu lnglis et Ogden,

1965) sont primitives par la longueur de la

côte 4 et par la présence du sillon excréteur, mais

toutes trois ont une bourse de « type 2-3 » ten¬

dant au « type 2-1-2 ». L’espèce décrite aux

Philippines, AI. asiaticus Tubangui et

Masilungati, 1937, est du « type 2-1-2 ».

En région néotropicale, sept espèces sont

connues chez les Carnivores et trois chez les

Primates. Les sept espèces parasites de

Carnivores, M. felineus Cameron, 1923, Al. bar¬

barie Cameron, 1936, AI. major Cameron, 1936,

Al. pardalis Cameron, 1936, Al. paraensis

Travassos, 1937, Al. nasuae Lent et Ereiras, 1938

et Al. brachiurus Costa et Freitas, 1967 possèdent

des côtes 4 courtes, six d'entre elles ont un sillon

excréteur et seule AI. paraensis en est dépourvue.

Trois espèces (du Brésil) ont une bourse caudale

de type 2-3 et quatre (une du Brésil et trois de la

Trinité) de type 2-1 -2,

Les trois espèces de Primates déjà connues.

Al. tomlasus (Molin, 1861), Al elegans Travassos

et Darriba, 1929 et Al. vexillarius Dunn, 1961

ont des côtes 4 courtes et un sillon excréteur pré¬

sent, mais toutes trois ont une bourse caudale de

type 2-1-2 tendanr au type 3-2. L’espèce décrite

ci-dessus partage avec les autres espèces de

Primates une bourse caudale de type 2-1-2 ten¬

dant au type 3-2, des côtes- 4 courtes, mais elle

est la seule à être dépourvue de sillon excréteur.

En résumé, le genre Alolineus conserve des carac¬

tères primitifs, de type Strongylina, chez les

Carnivores du monde entier. Une évolution mor¬

phologique vers un type plus spécialisé n'aurait

eu lieu que, soit dans une zone géographique très

limitée (nord du Brésil, Guyane, île de la

Trinité), soit chez les Primates. L’espèce décrite

ci-dessus, cumulant ces deux particularités, paraît

être la plus spécialisée actuellement connue.

RÉFÉRENCES

Cameron T. W. M. 1923. — Studies on two généra

and some little known species of the nematode

farnily Trichostrongylidae, Leiper. Journal of

Hehnintmlogy 1: 71-96.

— 1927. — On Microstrongylus genettae gcn. and

n.sp., a Trichosirongyle parasite of Genctla senega-

lensis. Journal of Helmintholngy 5: 81-88.

— 1928. — On some parasites of the ruscy tiger cat

(Felisplankrps). Journal of Helmintholngy 6: 87-98.

— 1936. — Studîes on the endoparasitic fauna of

Tnnidad. III. Some parasites of Trinidad

Carnivores. Canadian Journal of Research 14

25-38.

Chandler A. C. 1942. — The helminths of raccoons

in F.ast Texas. The Journal of Parasitology 28:

255-267.

Costa H. M. & Freitas M. G. 1967. — AJgunos hd-

minthos parasitos do Guara (Chrysocyon brachiurus

llliger), com a descricao de Alolineus brachiurus

n.sp. (Nematoda: Trichostrongylidae). Arquivos

EscoLi Veterinaria Alinas Gérais 19:25-29.

Dujardin F. 1845. — Histoire naturelle des Helminthes

ou oers intestinaux. Roi et, Paris, XVI + 650 ■>- 16 p.

Dunn F. L, 1961. —Alolineus vexitlarius n.sp.

(Nematoda Trichostrongylidae) tfoni a Petuvian

primate, Tamarintts ntgricollis (Spix, 18 23). Journal

of Parasitology 47: 953-956.

Durette-Desset M.-C. 1985. — Trichosrrongyloid

neijuiodes and their vertçbrate hosts: Recons¬

truction of the phylugeny of a parasitic group.

Advances in Parasitology 24; 239-306.

Durette-Desset M.-C. & Chabaud A. G. 1981a. —

Nouvel essai de classification des Nématodes

Trichostrongyloidea. Annales de Parasitologie

humaine et comparée 56 :297-312.

— 1981b — Sur les Molineinae parasites de

Mammifères. Annales de Parasitologie humaine et

comparée 56:489-502.

— 1995, — Note sur la nomenclature supra-familiale

des Srrongylida, Annales de Parasitologie Humaine

et Comparée 68 : 111-112,

Durette-Desset M.-C. & Pesson B. 1987. —Alolineus

patens (Dujardin, 1845) (Nematoda, Trichostron¬

gyloidea) et autres espèces décrires sous ce nom.

Annales de Parasitologie humaine et comparée 62 :

297-312.

Hasegawa H. 1989. — Two new nematodes from the

iriomote cat, Prionailurus iriomotensis. from

Okinawa: Uncinaria (Uncinaria) maya n.sp.

(Ancylostomatoidea) and Molineus springsmithi

yayeyamanus n.suhsp. (Trichostrongyloidea).

Journal of Parasitology 75: S63-S69.

lnglis W. &: Ogden C. G. 1965. — Descriptions of

some Strongyles (Nematoda) from mammals in

Easc Népal: with records of other parasitic nema¬

todes. Bulletin of the British Muséum (Natural

History) Zoology 13: 229-245.

ZOOSYSTEMA • 1998 • 20(3)

449

Durette-Desset M.-C. & Corvione M.

Lent H. & Freitas J. F. 1938. — Pesquisas helmintho-

logicas realisadas no Estado do Para. IV.

Trichosrrongylideos de mammifères. Memorias do

Instituto Oswaldo Cruz 33: 363-380.

Le Roux P. L. 1933. — On Tenuostrongylus cynictis

gen. and sp.n., a Trichostrongylid parasitizing the

yellow mongoose ( Cynictis penicullata). Armais and

Magazine ofNatural History, sériés 10, 10: 222-228.

Molin R. 1861. — Il Sottordine degli acrofalli ordina-

to scientificamente secondo i risultamenti délie

indagini anatomiche ed embriogeniche. Memorie

del Istituto Veneto di Science, lettere ed arti, Venezia

9: 427-633 (non consulté).

Petrov A. M. 1928. — Récognition of helminthofau-

na of fur-bearing animais of the USSR. II.

Nematodes of the digestive track. Proceedings of the

State Institute of Experimental Veterinary 5: 238-250

[in Russian],

Platt T. R. & Pence D. B. 1981. — Molineus samueli

n.sp. (Nematoda: Trichostrongyloidea: Molinei-

dae) from the badger, Taxidea taxas. Proceedings of

the Helmtnthological Society of Washington 48:

148-153.

Schmidt G. D. 1965 — Molineus mustelae sp.n.

(Nematoda: Trichostrongylidae) from the long-

tailed weasel in Montana and M. chabaudi nom.n.,

with a key to the species of Molineus. The Journal

ofParasitology 51: 164-168.

Sprehn C. 1932. — Lerhbuch der Helminthologie.

Gebrüder Borntraegen Berlin, 998 p.

Travassos L. 1937. — Revisao da familia

Trichostrongylidae Leiper, 1912. Monographias do

Instituto Oswaldo Cruz, 1,512 p.

Travassos L. & Darriba A. 1929. — Notas sobre

Heligmosominae. Medicina de los Paises calidas 2:

563-569.

Troncy B. M. 1970. — Contribution à l'étude des

helminthes d'Afrique, principalement du Tchad.

Bulletin du Muséum national d'Histoire naturelle

(Paris), série 3, 41 : 1487-1511.

Tubangui M. A. & Masilungan V. A. 1938. —

Nematods in the collection of the Philippine

Bureau of Science. III. Philippine Journal of

Sciences, Manilla 64: 257-267.

Zunker M. 1928. — Molineus europaeus spec. nov.,

ein neuer Nematode aus dem darm des Iltis

(Putorius putorius). Zeitsheiftung Parasitenkunde 2:

7-11.

Soumis le 2 octobre 1997;

accepté le 7 avril 1998.

450

ZOOSYSTEMA • 1998 • 20(3)

A study in medical history: introduction of

médicinal leeches into the West Indies

in the nineteenth century

Roy T. SAWYER

Biopharm (UK) Ltd, 2 Bryngwili Road,

Hendy, Carms. SA4 1XB (UK)

102364.1027 @ compuserve.com

Fred O. P. HECHTEL

School of Biological Sciences, University of Wales,

Swansea SA2 8PP (UK)

James W. HAGY

Department of History, University of Charleston,

Charleston, SC 29424 (USA)

Emanuela SCACHERI

Department of Biotechnology, Pharmacia & Upjohn,

Viale Pasteur 10,1-20014 Nerviano, Milano (Italy)

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E. 1998. — A study in medical histo¬

ry: introduction of médicinal leeches into the West Indies in the nineteenth century.

Zoosystema 20 (3) : 451-470.

KEYWORDS

Hirudinea,

Hirudinaria,

hirudin,

médicinal leech,

medical history,

West Indies.

ABSTRACT

Médicinal leeches were not found in the West Indies prior to 1822, but by

the turn of the century, a large, aggressive leech aboundcd on Puerto Rico,

St Lucia, Martinique and other islands. The authors conclude that this

“Caribbean leech", described as Hirudinaria (Poecilobdeltd) blanchardi

Moore, 1901, is a junior synonym of the “buffalo” leech Hirudinaria

manillensis (Lesson, 1842), the médicinal leech of India and neighbouring

countries of South-East Asia. The final proof of the true identity of this West

Indian leech came from comparison of the nucléotide séquences of the

cDNAs of the hirudin polypeptide from leeches from St Lucia and from

Bangladesh. The authors présent evidence chat this leech arrived from ships

carrying labourers from colonial India starting in the mid-l840’s. Each of

these ships were required to hâve leeches on board for médicinal purposes.

During this study, the existence of a second introduced leech species in the

West Indies was unexpectedly discovered, in Guadeloupe. The question

remains open whethet this second species is the médicinal leech intendonally

introduced into Guadeloupe from Sénégal by the French for breeding pur-

poses in the 1820’s.

ZOOSYSTEMA • 1998 • 20(3)

451

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

MOTS CLÉS

Hirudinea,

Hirudinaria,

hirudine,

sangsue médicinale,

histoire de la médecine.

Antilles.

RÉSUMÉ

Une étude d’histoire de la médecine : l'introduction des sangsues médicinales aux

Antilles au dix-neuvième siècle. Les sangsues médicinales n’avaient jamais été

rencontrées aux Antilles avant 1822, mais à la fin du siècle, une sangsue

agressive, de grande taille, était abondante à Porto Rico, Sainte-Lucie, en

Martinique ainsi que dans d autres (les. La conclusion des auteurs est que

cette « sangsue des Caraïbes » décrite comme Hirudinaria ( Poecilobdella )

blanchardi Muore, 1901 est un synonyme junior de Hirudinariu maniltensis

(Lesson, 1842), la sangsue médicinale d'Jnde et des régions voisines du Sud-

Est asiatique. La preuve définitive de la véritable identité de cette sangsue des

Antilles est apportée par la comparaison des séquences de nucléotides du

cDNA du polypeptide de I hirudinc des sangsues provenant de Sainte-Lucie

et du Bangladesh. Les auteurs démontrent que cette sangsue est arrivée par

les navires transportant la main-d’œuvre en provenance de Llnde coloniale à

partir du milieu des années 1840. Chaque navire devait avoir des sangsues à

bord, à des fins médicinales. Au cours de cette étude, l’existence d une

deuxième espèce de sangsue introduite aux Antilles a été découverte en

Guadeloupe. La question demeure de savoir si cette seconde espèce est bien

la sangsue médicinale du Sénégal introduite intentionnellement en

Guadeloupe par les français dans les années 1820.

INTRODUCTION

The introduction of a new animal or plant spe-

cies can hâve profou nd conséquences, especially

on islands and other isolated ecosystcms. Many

of such introductions h ave occurred. but usually

unrecorded, during the 300 years of active

European colonisation when there were mass

movements of people and materials to and from

the New World For example, yellow lever along

with the mosquito Aedes aegyipti (Linn.) was

introduced into the Caribbean and other parts of

the neotropical région aboard slave ships from

West Africa in the seventeenth century (Taylor

1971). Clearly, it is of general interest for future

environmental impact assessmems to identifv

spécifie examples of beretofore unrecognised

introductions of other bloodsucking animais. We

document in this paper a rare example of the

introduction of a médicinal leech into the West

Indies in the nineteemh century and give éviden¬

ce of yet another introduced leech.

On certain islands of the eastern Caribbean there

abounds today a large médicinal leech species

widely known as Canbeobdclla blanchardi. It was

described nearly a century ago and presumed to

be unique to the New World (Moore 1901;

Ringuelet 1976), having no near relatives what-

soever in the Western Hemisphere (Sawyer &

Kinard 1980; Sawyer 1986: 736). In this multi-

disciplinary paper, we présent molecular, mor-

phological and taxonomie evidence for the first

time that this remarkable West Indian species is

in fact identteal to Hirudinaria manillensis

(Lesson, 1842), the médicinal leech of India and

neighbouring countrics of South-East Asia. In

addition, we discovered in Guadeloupe that a

leech also known bv the spécifie naine blanchar-

di represents an undetermined species of

AsiaticohdeUa of African/lndian origin,

Current systematics recognises six species of

“buffàlo’’ leechcs in the Hirudinariinac, a subfa-

mtly ol the Hirudintdae cbaracterised by the pré¬

sence of a large vaginal caecum (“caecal pouch”)

in the femaie reproductive System (Sawyer 1986:

683-687) These six species ate divided into two

généra which are differentiated as follows: in

Poecilobdella , the femaie reproductive System has

a distinct “vagina" (termed “vaginal stalk" by

some workers) and the male System lacks ejacula-

cory bulbs (Fig. IB); whereas in Hirudinaria, the

femaie System lacks an elongate “vagina" and the

male System has ejaculatory bulbs (Fig. IA). Five

species occur naturally throughout tropical and

452

ZOOSYSTEMA - 1998 -20(3)

Médicinal leeches from the West Indies

subtropical Asia from the western limit of the We also address in this paper the question of

Indian subcontinent to the Pacific coast indu- when these leeches were introduced into the

ding numerous islands and archipelagcies. The West Indies and by what mechanism(s). We

sixth nominal species, “ Caribeobdella blanchar- document that no native médicinal leeches lived

dï", was First described as being from Puerto in the West Indies prior to 1822. Furthermore,

Rico as Hirudinaria ( Poecilobdella) blancbardi by there is no record of the existence of any leech

Moore (1901). resembling Hirudinaria on any of the West

6 - Xlbs/b 6 9 -Xllb 5 /b 6

Fig. 1 . — Comparison of taxonomically diagnostic features of the male (left) and female (right) reproductive Systems of the two géné¬

ra of “buffalo” leeches, Hirudinaria and Poecilobdella. A, Hirudinaria manillensis from St Lucia (the Martinique leech is the same,

unlllustrated); B, Poecilobdella granulosa (rom a dealer in "Madras area", India. See “Methods" for spécifie localities. co, common

oviduct; e. epididymis; eb, ejaculatory bulb; ed, ejaculatory duct; o, oviduct; os, ovisac; ps, pénis sheath; pt, prostate; v, “vagina"

sensu lato (portion between common oviduct and female gonopore); vc, vaginal caecum; vd, vas deferens. Scale bar: 1 mm.

Viewed from the left side, anterior to left. Note both species hâve a well-developed vaginal caecum.

ZOOSYSTEMA • 1998 • 20 (3)

453

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

Indian islands prior to 1827. However, starting

in the mid-1840’s ships taking émigrant labou-

rers from India to various British and French

islands of the West Indies were required to hâve

leeches on board for médicinal purposes. Over a

period of several décades these émigrant ships

took on thousands of the Indian médicinal leech

Hirudinaria , primarily at Calcutta and Madras.

We conclude that, about the late 1840 s, some ot

these leeches were releascd onto one or more

islands and that their descendants are the leeches

thriving today on St Lucia, Puerto Rico,

Martinique and other islands. The capability of

médicinal leeches to colonize very rapidly areas

where simdar species do not occur has been

documented in Guadeloupe (Pointier et al.

1988).

EARLY RECORDS OF MEDICINAL LEECHES IN THE

West Indies

The earliesr accounr of médicinal leeches in the

West Indies appears to be a report in 1817 by Dr

John Williamson who recorded: “Practitioners in

the West Indies labour under a great disadvanta-

ge, by not having leeches in that country. They

hâve been sent there at a great expense; but they

soon became sickly, and perished" (Williamson

1817: 361). The absence of leeches was later cor-

roborated in 1822 by M. J Achard, Government

Pharmacist at Port Royal, Martinique, who

recorded that there were no native species of

leeches in Martinique which could be used thera-

peutically (Achard 1825). In 1827, however,

Blainville tantalizingly described from

Martinique a 4 cm leech with blood in its crop

and other features of the Hirudinidae (Blainville

1827: 250; Moquin-Tandon 1846; 324). In

1893 Raphaël Blanchard of Paris, the foremost

leech taxonomïst at that time, reported without

further detail there was a species of

“ Hirudinaria" on Martinique, “whose presence

in the Antilles is a real cunosity”. In 1897

Blanchard went on to say, in rcference to a dis¬

cussion of "Hirudinaria ( Poecilobdella) gratiu-

losd\ an Asian species: “Also it is very interesting

to find it in the Antilles, where it has been,

without doubt, transported by man for médici¬

nal purposes; discarded in the streams where it

has acclimated. It is found in abundance in

Martinique, where it is very prosperous [,..]. We

hâve received numerous live animais, in two

batches [...]. One batch sent in 1891 by P.

Vanhaecke, Superior du Séminaire-Collège de

Fort de France [...]. In 1893 we received some

front another source, one of which was very

large, 245 mm in length. We know only by hear-

say that it occurs on islands other than

Martinique In 1901, a sintilar leech was

described as a new species Hirudinaria

(. Poecilobdella ) blanchardt by the éminent taxono-

mist J. Percy Moore of Philadelphia, from spéci¬

mens from Puerto Rico. In 1934, Oka obtained

specimens of the Martinique leech and compared

it externally with what he considered to be the

same species from Taiwan.

Early attempts to establish leeches IN THE

West Indies

During the colonial period the médicinal leech

Hirudo medicinalis Linnacus, 1758 was very

widely used for médicinal purposes throughout

Europe, including their colonies (Sawyer 1981).

From the beginning of the nineteenth century,

médicinal leeches had become increasingly rare

in Western European countries, most notably

France and England, and had to be imporred in

large numlters to meet an enormous demand. In

a single year, 1832, more than filry-seven mil¬

lions leeches were imported into France where

they were used mainly in hospitals in the vicinity

of Paris. Customs records document that during

the nineteenth cenrury more than one billion

leeches were imported into France alone from

easrern parts of Europe (Sawyer 1981). (loday

Hirudo medicinalis is listed as an endangered spe¬

cies and accordingly protected worldwide by the

CITES convention While overcollection

undoubtedly played a significant rôle, a full

understanding of the facrors underlying the

décliné of this and porenrially other médicinal

leech species worldwide is problemarical)

ln the meanwhile demand for médicinal leeches

in the New World was growing faster than sup-

ply. Although the United States had its own nati¬

ve leech species, Macrobdella décora (Say, 1824),

sometimes called the “American médicinal

leech”, it was generally recognized as inferior in

that it made a more shallow bite and bled much

454

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

less ( e.g. Wood & Bâche 1867: 442). We now

know the subfamily Macrobdellinae.

Macrobdella Verrill, 1872 and allies endemic to

North and South America hâve very reduced

bleeding rimes compared wirh the rrue médicinal

leeches Hirudo medicinalis and Hirudinaria

manillensis of the Eastern Hémisphère (Munro et

al. 1991), hence the need for che American colo¬

nises to import leeches. Accordingly, large num-

bers of H. medicinalis were imported into the

United States from Europe throughout the eigh-

teenth and nineteenth centuries and espccially in

the period before the American Civil War (Hagy

1991). Because of chronic supply and transporta¬

tion problcms, several serious attempts were

nrade to breed H. medicinalis in the United

States (Hessel 1881, 1884) but ail such efforts

failed, The same unsuccessful scénario also took

place in the Fnench West Indies,

Since the West Indies in the early nineteenth

century did not hâve any native médicinal

leeches, they were entirely dépendent upon

importation from abroad. At the rime the French

Antilles were, as was France irself, leech “manie"

and had been importtng thousands of H. medici¬

nalis from Europe since at least 1814 (Achard

1825). On at least one occasion in 1822 the

Antilles had even imported leeches, undoubtedly

Macrobdella décora, from “Newfoundland”

(Anonymous 1822).

In order to sacisfy increasing demand, the French

medical authoriries, as early as 1822, niade

serious attempts to breed Hirudo in the French

Antilles (Anonymous 1824; Achard 1825),

including French Guyana on mainland South

America (Conseil de Santé 1831). These

attempts at bteeding Hirudo are documented in

various reports in the Annales maritimes et colo¬

niales during the 1820’s and early 1830’s (see

Berget & Rey 1874 for full bibliography) but

were ail unsuccessful.

In 1829 leeches were imported from the French

colony of Senegambia in West Africa into the

Antilles {e.g. Dupuy 1830; Calve 1830). The

exact species involved is unclear (see Discussion).

In any evenr, the leech species the authors of this

paper collected in abundance in St Lucia,

Martinique and Pueno Rico are distinctly mem-

bers of the Hirudinariinae, a well-characterised

subfamily which does not live in Africa. The

bloodsucking (“hirudinid”) leeches of Africa are

very unlike Hirudinaria manillensis. No African

leech, for example, has a “vaginal caecum" so

characteristic of the Asian Hirudinariinae

(Sawyer 1986: 684); however, we leave open the

possibilitv that a leech species found today on

Guadeloupe is of African origin (sec Discussion).

By way ot summary, during the 1820’s the

French imported three species of leeches into the

French Antilles, including French Guyana. These

were Hirudo medicinalis from Europe,

Macrobdella décora from Newfoundland, and an

unidenrified hirudinid from Senegambia. None

of these species represent the large leech

Hirudinaria manillensis we collected in St Lucia,

Martinique and Puerto Rico.

MATERIALS AND METHODS

Materials

We sampled accessible streams and ponds by

slowly wading into the water, disturbing the mud

in the process. Leeches were collected by hand or

net while they swam near the water surface or

while they attached to the bare legs of the collec-

tors. Leeches were either taken alive to the labo-

ratory for breeding and turrher studies or were

preserved under field conditions with 5% forma-

lin ot 70% éthanol for later dissection and iden¬

tification.

We examined preserved .specimens from the

Caribbean and from Asia in the Smithsonian

Institution (Washington), Natural History

Muséum (London), Muséum national d’Histoire

naturelle (Paris) and Institute for Zoological

Taxonomy, Zoology Muséum. University of

Amsterdam. Flowever, owing to uncertainties of

labelling and constraints on dissecting old

muséum material, conclusions herein are based

on specimens recently collected alive from nature

by the authors or recently by colleagues.

Preserved specimens were pinned under alcohol

and dissected from dorsal midline to reveal dia¬

gnostic features of male and female reproductive

Systems. The drawings were made freehand with

the aid of an ocular micrometer. At least two

mature specimens were dissected from each loca-

ZOOSYSTEMA • 1998 • 20(3)

455

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

lity wherever possible. Dissected specimens arc

presently in the personal collection of the second

author but will eventually be lodged with the

Natural History Muséum (London) and the

Smithsonian Institution (Washington).

Specimens of leeches collected alive in St Lucia,

Martinique and Puerto Rico were examined

externally in detail and then carelully dissected

by F. O. P. Hechtel. Each was compared with

specimens collected in Bangladesh, India,

Philippines and other parts of Asia. The systema-

tics follows that of Sawyer (1986). Sawyer (1986:

683) inadvertently stated that the Hirudinariinae

hâve pharyngeal ridges terminating independent-

ly between the jaws. ThJs not the case for any

Hirudinariinae examined by us in the West

Indies nor in Asia.

Molecular genetics

To establish if the leech collected in St Lucia was

identical ro Hirudinaria manillensis, a genetic

comparison was niade of specimens from

St Lucia with specimens from an Asian popula¬

tion of H. manillensis. Although H. manillensis

occurs throughout South-East Asia we chose for

this study leeches of tliis species from Bengal

from which, according to historical évidence pre-

sented below, the West Indies leeches probably

originated. We chose a population from Sylhet,

Bangladesh, because it had been the basis of a

prior molecular study (Scacheri et al. 1993).

Toward this end arrangements were made to ship

live specimens of Hirudinaria manillensis collec¬

ted from Bangladesh to the laboratory of

Biopharm (UK) Ltd in Wales. Similarly, Sawyer

collected specimens from St Lucia in 1989 and

maintained them alive in Wales. Ir, 1990 Sawyer

took live individuals from each population to the

laboratory of the fourth author E. Scacheri, in

Milan, Italy. Individual heads were dissected

from the bodies, washed in 5 M NaCL and

quickly trozen in liquid nitrogen prior to storage

at- 80 °C.

Total cellular RNA was prepared from leech

heads essentially as described by Harvey et al.

(1986). The reverse transcriptase reaction was

carried out in a 40 pl volume as follows: 10 pg

total RNA from leech heads was mixed with 1 pg

oligo(dT) primer, 8 pi 3 mM dNTP mix and

8 pi reverse transcriptase buffer (230 mM

Tris/HCl pH 8.3, 300 mM KC1, 50 mM

MgCl2, 5 mM dithiothreitol), heated to 65 °C

for 2 min and quickly chilied on ice.

10 II RNasin (Pronicga) and 20 U avian myelo-

blastosis virus reverse transcriptase (Boehringer

Mannheim) were added, and the tube was incu-

bated ai 42 °C for 2 hours. The reaction mixture

was phenol/chloroform-extracted, isopropanol-

precipitated and resuspended in 60 pl stérile dis-

ulled water. Oligonucleotide primers were

synthesized on an Applied Biosystems model

380B DNA synthesizer. To obtain the complété

sequence of HM1 cDNA. three rounds of PCR

amplification were perforrned. Amplified pro-

ducts were analyzed on 1.5% agarose gel, phe-

nol-purified and ethanol-precipitated. For

further details see Scacheri et al. (1993).

Historical Research

During the course of this study, we idcntified

that the species from St Lucia, Martinique and

Puerto Rico ptobably originared from the Indian

subcontinent sometime in the last century.

Accotdingly, the question arose as to how and

when the leech could hâve been imported front

that far away région. The historical archives of

this period in the Oriental and India Office

Library, Blackfriars Road, London, and the

Colonial Office (CO) records of the Public

Record Office (PRO), Kew, London, were a rich

source of information. Much of this archivai

research was conducted with the invaluable assis¬

tance ofMrs Betty Thomson, Richmond, Surrey.

The third author J. W. Hagy documentcd the

importation of leeches into the French West

Indies in the 1820s, using archives of the British

Library, London, as well as the Intctlibrary

resources of the University of Charleston. With

funding from the University of Charleston, Hagy

focussed on Indian émigration archives of the

1840s at the Oriental and India Office Library,

London. Our research eventually focussed on the

mass movement of indentured labourers in the

1840's front India following the émancipation of

slaves on the West Indian islands. Archivai évi¬

dence is presented below which documents that

leeches regularly accompanied these labourers

456

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

HM1 St Lucia TCAAAAG

ATG

TTC

TCT

CTC

AAG

TTG

TTC

GTT

GTC

TTC

CTG

Amino acids

Met

Phe

Ser

Leu

Lys

Leu

Phe

Val

Phe

Leu

HM1 Bangladesh

ATG

TTC

TCT

CTC

AAG

TTG

TTC

GTT

GTC

TTC

CTG

HM1 St Lucia

GCT

GTT

TGC

ATC

TGC

GTG

TCT

CAA

▼

GCA

GTG

AGC

TAC

ACT

Ala

Val

Cys

Ile

Cys

Val

Ser

Gin

Ala

Val

Ser

Tyr

Thr

HM1 Bangladesh

GCT

GTT

TGC

ATC

TGC

GTG

TCT

CAA

GCA

GTG

AGC

TAC

ACT

HM1 St Lucia

GAT

TGT

ACG

GAA

TC A

GGC

CAG

AAT

TAT

TGT

CTA

TGC

GTG

Asp

Cys

Thr

Glu

Ser

Gly

Gin

Asn

Tyr

Cys

Leu

Cys

Val

HM1 Bangladesh

GAT

TGT

ACG

GAA

TCA

GGC

CAG

AAT

TAT

TGT

CTA

TGC

GTG

HM1 St Lucia

GGA

GGT

AAT

CTC

TGC

GGT

GGA

GGC

AAA

CAT

TGT

GAA

ATG

Gly

Asn

Leu

Cys

Gly

Lys

His

Cys

Glu

Met

HM1 Bangladesh

GGA

GGT

AAT

CTC

TGC

GGT

GGA

GGC

AAA

CAT

TGT

GAA

ATG

HM1 St Lucia

GAC

GGT

TCT

GGA

AAT

AAA

TGC

GTC

GAT

GGG

GAA

GGT

ACT

Asp

Gly

Ser

Gly

Asn

Lys

Cys

Val

Asp

Gly

Glu

Gly

Thr

HM1 Bangladesh

GAC

GGT

TCT

GGA

AAT

AAA

TGC

GTC

GAT

GGG

GAA

GGT

ACT

HM1 St Lucia

CCG

AAG

CCT

AAG

AGC

CAG

ACT

GAA

GGC

GAT

TTC

GAA

Pro

Lys

Pro

Lys

Ser

Gin

Thr

Glu

Gly

Asp

Phe

Glu

HM1 Bangladesh

CCG

AAG

CCT

AAG

AGC

CAG

ACT

GAA

GGC

GAT

TTC

GAA

HM1 St Lucia

ATC

CCA

GAT

GAA

GAT

ATA

TTG

AAT

TA A

CGAACGCATAT

Ile

Pro

Asp

Glu

Asp

Ile

Leu

Asn

End

HM1 Bangladesh

ATC

CCA

GAT

GAA

GAT

ATA

TTG

AAT

TA A

CGAACGCATAT

Fig. 2. — Comparison of the nucleotid6 and deduced amino acid sequences of the cDNAs o! the hirudln polypeptide variant HM1

from leeches Irom St Lucia and from Sylhet, Bangladesh. The arrow points to the signal peptidase cleavage site. The deduced

amino acid sequence corresponds to lhe complété aqiipo acid sequence determlned from leeches from Bangladesh by peptide map-

ping analysis published elsewhere (Scacheri et al. 1993).

aboard ship for medical purposes on the long

journey from India to the West Indies.

Since Mauritius in the sourhern Indian Océan

was a comrnon port of call for such ships, Savvyer

searched through selectcd baclc issues of the colo¬

nial newspaper Le Cernéen, Journal de L'Ilc

Maurice, from 1833 to 1872 looking for éviden¬

ce for the importation of leeches into this tsland

during this period. This research was conducted

in the reading room of the National Archives,

DBM Complex, Petite Rivière, Mauritius.

RESULTS

Taxonomy of leeches from Puerto Rico,

Martinique and St Lucia

Hirudinaria manillensis, the most comrnon and

widespread of the “buffalo” leeches of Asia, was

originally described as being from the Philippine

island of Luzon by Lesson (1842) (see also

Harding & Moore 1927). In 1986, Hechtel col-

lected specimens from this same island.

Figure 3A shows the large vaginal caecum charac-

teristic of the Hirudinariinae, as well as the pré¬

sence of ejaculatory bulbs and absence of an

elongate “vagina” characteristic of the genus

Hirudinaria, Following detailed morphological

examination, numerous specimens obtained

from Bangladesh proved unequivocally to be

Hirudinaria manillensis (Fig. 3B). [Hechtel noted

that Hirudinaria manillensis is polymorphie,

occurring in two main colour phases: a green

phase (darkish green dorsum and paler green

venter) and a reddish phase (dark reddish brown

dorsum and paler brick-red venter). Though

both phases occur together, at least in Asia, one

phase prédominâtes overwhelmingly in each

population.]

The type specimens of Hirudinaria ( Poecilob -

ZOOSYSTEMA • 1998 • 20(3)

457

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

délia) blanchardi could not be located and are

presumed lost. This being the case, dissections

(Fig. 3C) of specimens collected live from Puerto

Rico revealed no sigruficant différences between

them and rhe Luzon and Bangladesh leeches.

Specimens collected later from St Luda (Fig. IA)

and Martinique (not illusrrated) also proved to

be the same species. Specimens from Puerto

Rico, Martinique and St Lucia ail possess the

large vaginal caecum, ejaculatory bulbs and laclc

an elongatc “vagina”. On morphoiogical

grounds, we hâve no hésitation in assigning

médicinal leeches ol these three islands to the

spedes Hirudinaria manillensis.

Molecular studies focussed on an inter-popula¬

tion compatison of the genomic organisation of

the leech polypeptide hirudin, a well-characteri-

sed inhibitor of thrombin. As part of another

study the thrombin inhibitor secreted by

H. manillensis from the Bangladesh population

was partially purified (Electricwala et al. 1991).

Two variants were eventually found and sequen-

ced, called HM1 and HM2, diffeting in

ten amino acids in the central part of the molé¬

cule (Scacheri et al 1993), The protein structure

of the two hirudin variants include sixty-four

amino adds with six cystéine residues, plus ïwen-

ty residues which constitutc the signal peptide

required for extracellular sécrétion. This signal

peptide is identica! in both isotorms. Based on

this structural information Scacheri and her col-

leagues were able to isolate cDNAs for both

HM1 and HM2 by extraction of leech head

RNA, subséquent DNA synthèses and PCR

amplification. Furthermore, by cloning the geno¬

mic fragments of both variants they were able to

elucidate for the first time the gene organisation

of hirudin-like antithrombins from leeches

(Scacheri et al, 1993). Fully active recombinant

F1M2 was then produced in Escherichia coli cells

following transformation with a synthetic gene

Having characterised the hirudin gene from the

Bangladesh population of Hirudinaria manillen¬

sis in another study (Scacheri et al. 1993),

Scacheri compaied the same gene in leeches from

St Lucia with that from Bangladesh leeches.

Working at the DNA level it was unnecessary to

sequence the hirudin protein, thereby greatly

reducing the number of leeches required.

Furthermore, an important feature ol this techni-

cal approach is rhe ability of isolating PCR-

amplified clones from total RNA préparations

extracted from very few leeches, sometimes even

from one leech head. For the St Lucia population

cesearch focussed exclusively on the cDNA of the

best characterised isoform of hirudin (HM1), By

way of summary of these data, Scacheri found

that the nucléotide sequence for the HM1

cDNA from the St Lucia population was identi-

cal to that from the Bangladesh population

(Fig 2), This applied also to the nucléotide

sequence corresponding to the twenty amino acid

signal pepdde. In this context, it is very interesting

to note in terms of the rate of évolution that,

aithough the St Lucia and the Bengal populations

hâve been isolatcd for approximately 1 50 years,

die hirudin gene is ver)' highlv conserved,

In conclusion, based on comparative morpholo-

gy, as well as on comparison of the nucléotide

sequence of the hirudin gene, we hereby designa-

te the leech described originally as Hirudinaria

( Poecilnbdella) blanchardi Moore, 1901 from

Puerto Rico, and found also in St Lucia and

Martinique, as the junior synonym of

Hirudinaria manillensis (Lesson, 1842) of che

Philippines and other parts of South-East Asia.

Systematics

Family HlRUDINIDAE Whitman, 1886

Subfamily FllRUDINARIINAE Sawyer, 1986

Genus Hirudinaria Whitman, 1886

Hirudinaria manillensis ( Lesson, 1842)

(Figs IA, 3)

? Hirudo Martinicensis Blainville, 1827: 250

(Martinique). (Not Hirudo Martinicensis Moquin-

Tandon, 1826: 139).

Hirudo manillensis Lesson, 1842: 8 (Philippines, type

material could not be located).

? Hinulo Unicolor Moquin-Tandon, 1846: 324 ( new

name for Hirudo Martinicensis preoccupied).

Limnatis ( Pnmlobdelhi ) granulnsa - Blanchard 1893:

28 (undissected); 1897-' 345 (undissected).

Hirudinaria ( Poecilnbdella) blanchardi Moore. 1901:

214, pl. 12 (Puerto Rico, type material could nor be

located).

458

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

â - Xlbg/be $ - Xllbg/bg

9 — Xllbs/b 6

Fig. 3. — Comparison of taxonomically diagnostic teatures of the male (left) and temaie (rlght) reproductive Systems ot “buffalo"

leeches collected from Asia and the West Indies. See “Methods" for spécifie localities. A. Hirudinaria manillensis from Luzon,

Philippines (type locality); B, Hirudinaria manillensis from Bangladesh; C, Hirudinaria manillensis from Puerto Rico. Scale bar; 1 mm.

See Fig. 1 for key to labelling and orientation.

ZOOSYSTEMA • 1998 • 20 (3)

459

Sawyer R. T., Hechtel F. O. P-, Hagy J. W. & Scacheri E.

Fig. 4. — Comparison of taxonomically diagnostic features of the male (upper) and female (lower) reproductive Systems of two spe-

cies of the genus Asialicobdella ; A, Asiaticobdella fenestrata from Gambia, viewed from dorsal side, anterior to top; B, undetermined

species of Asialicobdella from Guadeloupe, viewed from the left side, anterior to left. See Discussion for spécifie localities. vs, vagi-

na “sensu strictef; vt, vaginal duct (or “stalk”). See Fig, 1 for key to other labelling. Scale bars: 1 mm. Note neither species has a

vaginal caecum.

Limnatis grantdosa - Oka 1934: 286, fig. (externals)

(undissected).

Caribeobdell/i blanchardi- Ringuelet 1976: 13.

“ Poecilobdella" blanchardi — Sawyer &c Kinard 1980:

84 (Puerto Rico, dissected; Antigua and Haiti, undis¬

sected).

Hirndinaria manillensis — Sawyer 1986: 687,

fig. 18.9E.

MATERIAL EXAMINED. — Philippines. Calumpang,

Laguna, I4"1Q’N - 121 ° 18'E, Noveniber 1986, col-

lected by F. O. P. Hechtel.

Bangladesh. Sylhet, 24“53’N - 9l°51'E.

Puerto Rico. SW Puerto Rico, Cartagena Lagoon,

27.V(I1.1973, eollected by j.W. Miller and I.

Pomales, Department of Marine Sciences, U.P.R.

Mâyaguez, P.R. 00708.

St Lucia. Cattle pond 21 3 miles south of Micoud,

13°4S.2'N - 60”55 8’W, September 1989, eollected

by R. T. Sawyer.

Martinique. Small stream, Tributary of Lazarde

River, Route du Vert-Pré, Lamentin, 29.V. 1998, col-

lected by J. Vaubon.

460

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

Genus Poecilobdella Blanchard, 1893

Poecilobdella granulosa (Savigny, 1820)

(Fig. IB)

Sanguisuga granulosa Savigny, 1820: ! 15 (type locali-

ty: Pondichéry, India).

Poecilobdella granulosa - Sawyer 1986: 687, fig. 17.

16B.

MATERIAL EXAMINED. — India. — Specimens sup-

plied by a dealer in the “Madras area", 1984. —

Specimens obtained in 1997 by Dr Rames h Yadav

from a Bombay dealer who reported they had been

collected by the “Adivasi" people front the lakes neat

the city of Baroda, Gujarath State, India.

Subfamily HlRUDlNINAE Richardson, 1969

Genus Asiaticobdella Richardson, 1969

Asiaticobdella sp.

(Fig. 4B)

Himdinaria blanchardi — Pointier, Théron & Imbert-

Establet 1988: 38 (Guadeloupe, undissected).

MATERIAL EXAMINED. — Guadeloupe. Specimens

purchased by R. T. Sawyer in the market at Pointe-à-

Pitre, Guadeloupe, in August 1995. — Specimens

collected alive on 6.II. 1997 in little ponds to the

north of the airport, west of Abmes, by N. Barré.

Asiaticobdella fenestrata (Moore, 1939)

(Fig. 4A)

Limnatis fenestrata Moore,1939: 343. pis 27, 28 (type

locality: Botswana).

Asiaticobdella fenestrata — Sawyer 1986: 776, 777,

fig. 18.14D.

MATERIAL EXAMINED.— Gambia. I.ive specimens

acquired in July 1993 from a sacrcd crocodile pool at

Katchikaeli (13°28’N - 16°40’W) in Coastal Gambia,

close to the Southern bank of the Gambia River at its

mouth, through the kindness of C. M. Moiser.

Researchers are cautioned that médicinal leeches

in the West Indies cannot be distinguished from

external characters alone and précisé identifica¬

tions must be based on dissection of the repro¬

ductive Systems. To avoid confusion, researchers

are advised to detail the nature of rhe reproducti¬

ve System when maleing identifications, accor-

ding to the following simpltfied key. It cannot be

ruled out that more than one spccies ol médici¬

nal leech lives on any of the islands.

SlMPLIFIED KEY TO THE MEDICINAL LEECHES IN THE WEST INDIES

1. Vagina with a large caecum (“caecal pouch”) (Fig. 1 : vc) ..2

— Vagina lacking a large caecum (Fig. 4B) . Asiaticobdella sp.

Known from Guadeloupe

2. Female reproductive System has a distinct “vagina” (elongate portion between the

female gonoporc and the common oviduct); male System lacking ejaculatory bulbs

(Fig. IB) . Poecilobdella granulosa { Savigny, 1820)

Not recorded from the West Indies

— Female reproductive System lacks an elongate “vagina”; male System with ejaculatory

bulbs (Fig. IA) . Himdinaria manillensts (Lesson, 1842)

Known from Puerto Rico, Martinique and St Lucia

DISCUSSION

MeCHANISM OF TRANSPORT OF LEECHES FROM

India to the Caribbean

The leech on St Lucia, Martinique and Puerto

Rico is actually the Asian médicinal leech

Himdinaria manillensis which appeared on the

islands in the middle of the nineteenth century.

To try to explain how the leech could hâve been

introduced, we started looking for historié

ZOOSYSTEMA • 1998 • 20(3)

461

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

connections, especially medical, betwecn che

West Indies and South-East Asia ac about this

time. We discovered â significant connection in

the émigration of a large number of labourers

from India into both the British and French

islands (see Thomas 1985 for further back-

ground).

The economy of the West Indian colonies was

largely built on sugar. This required heavy labour

and African slaves were brought in for this pur-

pose. In England the Emancipation Act ot 1833

provided for the graduai treedom of the slaves in

the colonies, this having been completed by

1838. To replace the slaves, the West Indian

planters furned to India for immigrant labo tir.

After some taise starts amidst controvcrsy with

abolitionists, the Colonial Office in 1844 appro-

ved a scheme for indentured Indian émigration,

wholly managed by the English governmenr in

order to protect the health and safety of the

labourers. The émigrants were promised return

passages to India after five years. In 1845 two

shiploads of Indians reached British Guyana, and

one ship cach went to Jamaica and Trinidad. The

voyage from Calcutta to Trinidad took between

eighty-five and ninety-two days, typically stop-

ping at Cape of Good Hope or St Helena. By

further example, Captain J H. Wilson, West

India Emigration Agent, dispatched twelve ships

in the 1845-1846 season and seventeen ships in

1846-1847, varying in passengers number from

203 to423 (Public Record Office 1847: 158).

Because of a disruption foilowing the Sugar

Duties Act of 1846, large scale émigration from

India was not again fully underway until 1851.

In the lS50’s the Windward Islands were allowed

to recruit small numbers of Indian labourers on

the usual ternis, permission being granted to

Grenada in 1856, St Lucia in 1858 and

St Vincent in 1861. These small islands reques-

ted indentured labour only irregularly and in

small numbers For example, durlng the émigra¬

tion period to St Lucia from 1859 to 1869, the

English landed 4354 Indians from Calcutta.

The French emancipated their slaves in 1848 and

an acute shorrage of labour resulted in the colo¬

nies. The planters were very conscious of the

example set by the English colonies (and the

French island of La Réunion in the Indian

Océan). In 1852, an immigration law was passed

providing officiais to supervise recruiting and to

look after the welfare oi émigrants. Consequently

the Compagnie Générale Trans-Atlantique artan-

ged to supply 2000 or 3000 Indians each year. In

1854, an Immigration Committec was set up to

control the whole operation. Subsequendy, a full

code of immigration régulations appeared as laws

in 1855 and 1859 vvhich made the rudimentary

protective organization much more elaborate and

efficient. The Compagnie Générale Maritime

contractcd to supply Martinique with 1500

Indians over tour years.

The English government sought to persuade

France to give up recruiting in Africa and in

1861 agreed to let the French colonies recruit

labour in British India on much the same terms

and under the same régulations as did the

English colonies (Parliamentary Papers 1861).

Some Indians had been brought in by the French

in the 1850’s, over 9500 into Martinique and

perhaps 1000 into Guadeloupe from the French

Indian territorîes of Pondichéry and Chander-

nagore. But these territories could supply only

limited numbers, so British India became the

recruiting ground. Altogether, between 1853 and

the termination of the agreement with England

in 1885, 25509 Indians were landed in Marti¬

nique, embarking mostly from Calcutta and

Pondichéry. In Guadeloupe, until 1861, most of

the migrants, mainly Tamil, originated from

South India (mainly Pondichéry). From 1873

this strearn became secondary compared with the

Calcutta région (Centre d'Etudes 1982). From

1856 to 1889 over 40000 Indians landed in

Guadeloupe.

In order to ensure the health of the émigrants,

Her Majesrys Colonial Land and Emigration

Commissioners enforced strict conditions and

medical requirements onto the contracror of each

shipload of émigrants (Public Record Office

1847). As shcnvn below, each ship from India

raking émigrants io the West Indies from the

mid-1840’s to che early 1870s was required to

bave leeches on board for médicinal proposes.

Based on the foilowing historical evidence, we

propose this was the most probable mechamsm

by which Hirudinaria manille mis came to be in

St Lucia and other islands of the West Indies.

462

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

From British India; Calcutta and Madras

The records of the Public Record Office (PRO)

and the Oriental and India Office in London

include the following observations, relevant ro

our study. For administrative reasons, the Brirish

ships transporting lndian émigrants to the West

Indies embarked almost exclusively from either

Calcutta or Madras (Public Record Office 1847).

In order to ensure the health of the émigrants,

Her Majescy's Colonial and Emigration

Commissioners enforced strict conditions and

medical requirements on the contractors (Public

Record Office 1847). Each ship was required to

hâve a list of medical supplies before embarka-

tion. In 1847, this "List of Medicines and

Medical Comforts" included one hundred

leeches for up to 100 émigrants, to be increased

by 50 for each 100 émigrants beyond 100

(Public Record Office 1847: 21). From the

example of Captain J. H Wilson given above,

his twelve ships would hâve transporred toward

the West Indies a total of approximately

2 000 lndian leeches in the 1845-1846 season,

and siinilarly his seventeen ships would hâve car-

ried approximately 3000 leeches in the

1846-1847 season. Interestingly, these particular

ships origiuated from Madras and were bound

for British Guyana, Trinidad and J a ma ica, none

of which appears to harbour the leech today.

The leeches had to otiginate “fresh” at the port of

embarkation of Calcutta or Madras and “not

England”. In a lettet datcd 10 March 1847 to

the West India Emigration Office, Madras, the

same Captain J. H. Wilson recommendcd “that a

clause be introduced in the Charter strictly enjoi-

ning that vessels shall purchasc every article of

provision required by the régulations lresh at the

port of embarkation" (Public Record Office

1847: 337). Similarly, in the Emigration

Commissioners’ official “Tender for the

Conveyance of lndian Emigrants to the West

Indies” dated Juttc 1847, item eight records “[...]

and also a supply of Medicine and Medical

Comforts according to the annex [...]. Provided

alvvays, that ail articles of Provisions for the use

of the Emigrants shall be provided and put on

Board in India and not in England" (Public

Record Office 1847: 21). In other words the

leech species in question originated in British

India, i.e. Hirudinaria and therefore certainly

would not hâve been the European médicinal

leech Hirudo médicinal! wh ich does not live in

the lndian subcontinent (Sawyer I486: 571).

Records show that médicinal leeches were requi¬

red ro be on board émigrant ships from 1847 to

1871. In the 1856 “Rules for Rcgulating ail

Matters Connected with Emigration from

Madras to the West Indies” 500 leeches were

required for 50-100 emigranrs (India Office

1856: 17). The 1859 “Revised Rules Re Coolie

Emigration Including a List of Medicines" requi¬

red 50 leeches per 100 persons; 75 per 200 per-

sons; 100 per 300 persons; and 100 per 350

persons (Public Record Office 1859: 36). The

1864 “Rules for the Guidance of the Protector of

Emigrants in Calcutta" required 50 leeches per

100 persons; 75 per 200 persons; 100 per

300 persons; and 100 per 350 persons (India

Office 1864. 4). The 1871 "Emigration from the

Port of Madras. Rules under Act of 1871" requi¬

red 50 leeches per 100 persons; 75 per 200 per¬

sons; 100 per 300 persons; and 125 per

400 persons (India Office 1874). In the same

year the 1871 schedule 3 “The Medicines, Rules

Under Act Vil (The India Emigration Act)”

required “leeches" but no numbers were specified

(India Office 1872: 394). Interestingly, the 1883

“Rules Relating to Emigration from Calcutta”

required “one sixteen oz blood porringer” but no

mention of leeches (India Office 1884). We hâve

not found any record of the actual medical use of

leeches on board these émigrant ships, but such

records arc to be expccted sincc each ships sur¬

geon was required to keep a medical diary.

A few records document that leeches were used

mcdicinally in the Caribbcan région about this

time. Leeches were used successfully following

arterial surgery in tire Hospital of St Felipe and

Santiago, Havana, in 1849 (Wills 1849:

148. 149). Dr Heccor Gavin MD FRCS,

Lecturer on Forensic Medicine at Clearing Cross

Hospital, London, in his 1851 report regarding a

recent outbreak of Yellow Fever in Surinam,

enclosed a translation of a report by the Dutch

Medical Officer H. Schomnberg of 5 September

1851 on the treatment of Yellow Fevet near

Paramaribo, Surinam. Leeches were part of the

treatment: “leeches [...] generally proved very

ZOOSYSTEMA • 1998 • 20(3)

463

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

bénéficiai [...]” and Leeches produced a very

satisfactory resuit” (Publie Record Office 1851).

In France and England, the use ol leeches rea-

ched a peak about 1820-1845 and gradually fell

out of favour by the 1870 s (Sawyer 1981).

Bloodletting, but not necessarily leeching, conti-

nued on British ships up to rhe 1880’s and

1890’s and probably later, but the practice was

becoming suspect For example, Acting Assistant

Surgeon M. Elphington Greany of HMS Vestml

in 1869-1870 recorded the following: Dr.

Bellot, a great Havana authoriry on Yellow Fever,

is accustomed, 1 understand, ro bleed in almost

ail cases, if seen in the early stage, and 1 myself

hâve been advised in Port-au-Prince to use the

lancet, but I saw no case in which it would be

allowable to do so. I should rather fancy that

such a proceeding would be fatal to any chance

of a patients recovery” (Public Record Office

1870).

Although we hâve documented that thousands of

Indian médicinal leéches Hirudinaria were trans-

ported to the West Indies front the nmd- 1840s

to the early 1870 s, the final fatc of these leeches

is so far undocumented. We are aware thar:

“Ships surgeon shall receive charge of medical

stores. He ntust ascertam quality and that the

supplies are not short. On arrivai at the port of

debarkadon the surgeon is ro deliver the balance

of medical stores to the Emigration Agent at that

port with a statement of issues during the voya¬

ge’’ (India Office 1864: 4). We propose that

some of the Hirudinaria manillensis ended up in

local water and established themselves.

From French India: Pondichéry

Apart from the account above of labourers going

to Martinique and Guadeloupe from Pondichéry,

and later from Calcutta in coopération with the

English émigration policy, we hâve not found

direct evidence that leeches were put on board in

Pondichéry for .ships desdned to the West Indies.

Moquin-Tandon (1846: 341) did record that a

médicinal lecch, which he called “Htrudo granu-

losa" was “employed by the doctors of

Pondichéry”. In an 1857 report from the India

Board to the British Colonial Office regarding

Indian émigration on board British ships from

Pondichéry to French colonies in the West

Indies. a list of required médicaments did not

spedfy leeches (Public Record Office 1857: 318).

Howcver, Leuckart and Brandes elaimed, unfor-

tunately without giving any further detail, that

Hirudinaria at one time "was sbipped out of

India (Pondichéry) in large quantifies to the

islands of Bourbon and Mauritius” in the Indian

Océan (Leuckart & Brandes 1901: 879).

Interestingly, under the Convention with the

French, the British in 1862 transported over

4500 Indians into La Réunion from Calcutta

and Pondichéry (Public Record Office 1862).

Leech Importation into Mauritius

The économie and social history of Mauritius in

rhe Southern Indian Océan is remarkably similar

to that of islands of the French West Indies. In

1997 Sawyer visited Mauritius to détermine

whether médicinal leeches had been imported

into the island in the pasr and whether the leech

may hâve escaped, as in rhe West Indies (Sawyer

in press). By searching through adverrisements in

the colonial newspaper Le Cernccn, Journal de

Plie Maurice, clear evidence was found that for at

least the forty year period from 1833 to 1872,

large numbers of leeches were intentionally

imported from Pondichéry into Mauritius by

local pharmacists for médicinal purposes (Fig. 5).

A sélection of such adverrisements follows:

31 mai 1833. “Ch ez M. Grosjeau neveu, rue

St George : belles sangsues de l’Inde arrivées par

Y Antoinette. ”

Note: on 11 ]une 1833 “Arrivages [...] La barque

Y Antoinette, capit. Colin, partie de Madras, et de

Pondichéry le 15 avril ; cargaison riz et diverses mar¬

chandises. Passagers vingt Indiens.”

2 mars 1848. Belles Sangsues de Pondichéry, à 1 p.

la douzaine. S’adresser à M. Guîot ou à M. E.

Fleurot."

Note: previously recorded ships from Pondichéry,

reported on 22 Fcbruary 1848: (a) Brig. “ Mauritius

Packet ” from Pondichéry. 12 Janu.iry, “with sundries

for this port": and (bj Bark “ East Anglian from

Pondichéry, 18 January, “with sundries for this port”.

25 avril 1872. "Belles Sangsues de Pondichéry

s'adresser à la Pharmacie B. Perrot, rue Desforges,

11° 67.”

Note: this Street now is Sir Seewoosagur Ramgoolan

St, Port Louis.

464

ZOOSYSTEMA • 1998 • 20(3)

Médicinal leeches from the West Indies

A VENDRE.

Véritable tin de Constance blanc et ronge en chopiueï,bon

porter en bouteilles à 2 p. 7o e. la douzaine.

S'adrt s-or à A). D. Bonuelin, jeuue, rue St Georges.

— Ch. Z M. BENOIT, marchand, rue Uesforgcs : bi< re

(i'Uugd'on, pi enucrc qualité, ù 2 p. 25 c. lu douzaine et 4L a.

tu bouteille ; vin du Bouleaux, premitne qualité, à 25 ». U

bouteille ; buttgir diaphane (T Angleterre ; le tout nu compta m.

— En gros ou en détail, BELLES SANGSUES ,

années de Pondichéry par le navire l’Etmnée.

S'adresser au magasin Heyncmans.

ni la

de la

gnio,

»r la

*o rie

1 une

j*j ie

Belles SANGSUES

ue eosmciiÉBY.

S’adresser à la Pharmacie B. PEIIUOT,

ruo Desforoos, No. G7.

Fig. 5. — Représentative advertisements by pharmacists in Port Louis, Mauritius, in the colonial newspaper Le Cernéen, Journal de

LÏIe Maurice. Top; 6 December 1833. Bottom: 25 April 1872. Leeches sold in pharmacies in Mauritius in the Southern Indian Océan

were imported from Pondichéry, india, for over forty years.

No evidence vvas found that some of the ships

carrying leeches from Pondichéry went on to the

French West Indies to supply medical dernand,

but that possibility must be left open.

Unfortunately, the specics of leech imported into

Mauritius from Pondichéry in the nineteenth

century remains as yet undetermined

Although many médicinal leeches were indeed

imported into Mauritius, there is no evidence

that the leech species in question escaped and

established itself in the wild. Quite the contrary

while several leech species were collected on the

island, no bloodsticking species at ail were

encountered (Sawyer 1997, personal observation).

Local people very familiar with the wildlife of

Mauritius were unanimous in confimting that no

bloodsucking leeches occur on the island today.

Medical ethnology

Leeches were enormously valued for medical

purposes in the last century and practitioners

were highly motivated to acquire them. Since the

medical need for leeches undoubtedlv motivated

their importation into the Cartbbean area in

some numbers, an ethnological assessment of the

current medical use of leeches in this région is

relevant to this study. Sawyer carried out nume-

rous interviews of local people on various West

Indian islands, as well as French Guyana,

Surinam and Mauritius. Ofmuch value were the

local markets svhere leeches were still being sold

in recent tintes. Whenever possible such market

leeches were purchascd for later identification. A

more formai medical ethnological study was

undertaken in rural St Lucia, where in-depth

interviews were conducted by Sawyer with six

local people recognized as "healers’ 1 by the

St Lucia National Trust. The latter study was car¬

ried out in French patois in September 1989

with the invaluable assistance of Mr. Laurent

ZOOSYSTEMA • 1998 • 20(3)

465

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

Jean Pierre of the Sc Lucia National Trust,

Castries. Histotical évidence presented above

suggested that the St Lucia Icech may hâve origi-

nated from Bengal, where it had been used medi-

cinally in the last century. To learn more Sawyer

went to Dhaka, Bangladesh in July 1992, where

he conducted extensive interviews wich local

Street vendors who were still selling Hirudinaria

manillensis for médicinal purposes in time immé¬

morial tradition.

Locally collected leeches were still being sold for

médicinal purposes on some ofthe islands ofthe

West Indies, including the main markets in

Castries, St Lucia (1989), Fort-de-France,

Martinique (1993, but nor by 1998) and Pointe-

à-Pitre, Guadeloupe (1995). As late as 1976

médicinal leeches were being sold in at least one

pharmacy in Cayenne, French Guyana, but by

1993 no pharmacist contacted in Cayenne was

aware of anyone using leeches anymore in French

Guyana (Sawyer personal observation).

In St Lucia (1989) much effort was made to

interview market sellers, local pracritioners and

récipient users of leeches in urban as well as in

remote régions. Today, these are almost exclusi-

vely people of African descent who speak French

patois as dreir flrst or only lauguage, even on this

nominally "British” island. tn the Castries mar¬

ket more than one vendor sold leeches (“sans!

corrupted from the french “sangsues”). Leeches,

reportedly from Dermery, were purchased from

one vendor and latex identified definitely as

Hirudinaria manillensis. The market leeches were

in any sort of bottle, which invariably had a

piece of charcoal at the bottom. One seller

explained (erroneously) the leeches “ate the char-

coal” which could be “dried and rc-used . Most

people seemed to be aware of leeches and their

médicinal use, and usually knew a relative who

had used them some time in the past. Scveral

people made the comment that leeches were less

commonly used now than a couple of généra¬

tions ago, One young rnan said his grandmother

“used to keep them around hcr house”. Leeches

were used sparingly, but for appropriate condi¬

tions, such as for “black eyes”, “swollen feet”,

“boils”, “blood poisoning” and “snake bire”.

Some were applied “to the back” for undiagnosed

conditions, and one young woman’s grandfather

reportedly had leeches applied several times a

week for a while for some undiagnosed condi¬

tions. Leeches apparently were not used for eye

complaints except black eyes. They could be re-

used by placing sait cm them and squeezing out

the blood. One intelligent old man recognised as

a local “healer” volunteered that his grandmother

told him that ethnie lndian people (i.c. from

India) were “the best users of leeches”. This same

observation wos made by a French scientist in

St Lucia in reference to Guadeloupe. Several

people observed that leeches were less abundant

now than when they were young. They attribu-

ted this décliné to the “pesticide being used to

treat nematodes”, and to “banana irrigation”. It

was reported several times that more than one

species of leech lives in St Lucia (“one does not

bitc" by one account, and “one very aggressive

but not so good; the slow sucking one better” by

another account). Sawyer could not confirm a

second hirudinid species in St Lucia front his

limited fteld studies there. One wcll-traveled

St Lucian observed that leeches were more abun¬

dant in St Lucia than in any of the other islands.

Sawyer can certainly confirm that leeches, identi¬

fied as Hirudinaria manillensis, were locally very'

conimon in St Lucia in September 1989.

(Sawyer was reliably intormed, but has not yet

confirmed, rhat leeches occur and are also used

medicinally on Dominica and Granada).

Most of the West Indies today is populated

mainly by people of African descent, and many

aspects of the culture reflect this African héritage.

In a separate on-going srudy, Sawyer has found

no evidence that live leeches were ever used in

traditional medicine in black Africa. At the same

time. it is well-docurnerited that bloodletdng and

cupping were, and continue to be, widely practi-

ced there (e.g. Livingstone 1857: 129, 130). In

contrast, the médicinal use of leeches was com-

monplace to French and British colonists, as well

as widely practiced by people ofthe lndian sub¬

continent. The so-called "buffalo” or “cattle”

leeches hâve been used medicinally for over two

thousand years in the lndian subcontinent, a

practice which continues today. On rhe srreets of

Dhaka, Bangladesh, the species used is

Hirudinaria manillensis (Sawyer 1992, personal

observation). Leeches are still commonly used in

466

ZOOSYSTEMA • 1998 • 20 (3)

Médicinal leeches from the West Indies

rhe traditional (Ayurvedic) hospital.s and ciinics

of Bombay on the west coast of India. Some spé¬

cimens of rhe leech used clinically in Bombay

were sent to Hechtel for identification by Dr

Ramesh Yadav and were found to be

Patcilobdella granulosa. Leeches of an undetermi-

ned species are reportedly still being used. in the

villages in rhe hinterland of Sri Lanka (Sawyer

1995, personal observation).

The use of these large, aggressive leeches is des-

cribed in considérable detail in Chapter Xlll of

the Susruta Samita which records the ancient

Ayurvedic form of Indian medicine

(Bhishagratna 1963: 98-105), dating back by

some accounts to 200 BC. One vvould expect

these peopie to bring this tradition with them to

the West Indies, bearing in mind that the

Europeans had already been using leeches there.

A case of- récent leech colonisation

The ability ol a médicinal leech spccies to coloni-

ze an isolared pond very tapidly has been rho-

roughly documenred during a fifteen year study

from 1972 to 198” in a small lake in Guade¬

loupe (Pointier et al. 1988). Lake Grand Étang is

located in the tain forest at an altitude of 450 m

and has “no permanent humati habitation within

a radius of 3 lent". During a biological survey in

1972, there were no leeches in the lake, but in

1973 a locally commun species of leech was

introduced by local peopie. The leech population

increased rapidly where they fed mainly on the

tilapid fish Oreocbromis mossambicus (Peters,

1844). The leeches becarne “an important new

factor contributing to fish mortality; manydying

fish were seen floating on the lake or stranded in

rhe aquatic végétation 1 '. This well-documented

case may give us a due to rhe success of this and

pos.sibly other leech species, />. their ability to

thrive on the fish O. mossambicus which was

introduced to the West Indies from Africa as a

source of protein. (Note: this leech species which

understandably was called Hirudinaria blanchar-

di is proliably the same as the Guadeloupe “mar¬

ket leech” discussed below).

Evidence for a second introduced leech in

the West Indies: Guadeloupe market leech

Until recently, we had presumed in our investiga¬

tion that ail médicinal leeches found on the

various islands of che West Indies represented a

single species, Hirudinaria manillensis , which we

had concluded came from India in the mid-

1800’s. Most of our morphological and molecu-

lar genetics data were based on specimens

collecred alive in St Lucia, and corroboratcd with

morphological data from specimens collected

alive in Puerto Rico and Martinique. The first

hint that a second médicinal leech species is pré¬

sent in the West Indies resulted from dissecting

specimens purchased by Sawyer in the market at

Pointe-à-Pitre, Guadeloupe, in August 1995. To

our surprise this “market leech", while iooking

similar exfernally to Hirudinaria manillensis , is

quite different internally. The jaw structure and

reproductive organs clearly demarcate it from ail

other hirudinid species (Macrobdellinae) ol rhe

New World. (To confirm that the “market leech”

actually lives in the wild in Guadeloupe rather

than purchased from ou rade the island, N. Barré

kindly collected live specimens of the same leech

species west of Abymes. Hechtel confirmed by

dissection that these were the same as the “mar¬

ket" species).

Though in appearance very similar to

H. manillensis , dissection of the reproductive Sys¬

tems of the Guadeloupe “market leech” (Fig. 4B)

revealed it to be an undetermined species of the

genus Asiaticobdella. This genus belongs to the

Hirudininae, a subfamily disttnguished from the

Hirudinariinae by the absence of a spacious vagi¬

nal caecum. As currentlv defined (Sawyer 1986:

688), the genus Asiaticobdella occurs in both

India and Africa (Harding & Moore 1927).

Although the authors are unable to rule out an

Asiaticobdella of Indian origin, historié and taxo¬

nomie evidence presented below leaves open the

possibility that the Guadeloupe leech may be of

African origin.

After tàiling in the last century to introduce the

European médicinal leech Hirttdo medicinalis

into the Frc-nch West Indies, M. Gerbidon,

intérim governor of Sénégal, proposed in 1827

the idea of sending leeches from Senegambia on

the norrh-west coast of Africa ( e.g. Dupuy 1830;

Calve 1830) to the Antilles, including to

Cayenne, French Guyana. After at Ieast one fai-

led attempt, in June 1829, he successfully ship-

ZOOSYSTEMA • 1998 • 20 (3)

467

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

ped 50 000 leeches from Sénégal, of which

34 000 survived. Of these the French authorities

purposely rcleased about 16 000 into the ponds

and streams of Guadeloupe. In December 1829

it is recorded that “the Negroes hâve found them

on their legs and on the legs of animais, and

fîshermen hâve seen several in their nets (nasses).

Unfortunatcly, Negroes hâve sold a great nuxnber

of them in Pointe-à-Pitre.”

In order to elucidate the possibility chat leeches

may hâve been introduced from Senegambia,

Hechtel acquired live specirnens of a candidate

hirudinid leech from Coastal Gambia. After care-

ful study of its external and internai characters

(Fig. 4A) Hechtel conftdently idenrified this

Gambian leech as Asiaticobdella fenestrata (see

Sawyer 1986: 774-778 lor key to African

Hirudiniformes). This study also confirmed that

the Gambian leech was nor the same as the

Guadeloupe leech. They dilïer specifically in that

A. fenestrata has a vaginal duct about 2.5 to

3 times the length of the vagina (sensu stricto)

whilst the Guadeloupe leech has a vaginal duct

about 1.5 times the lengfh of the vagina. The

pénis sheath is also commensurately longer in

A. fenestrata. To date we hâve not been able to

match the Guadeloupe leech with any descrip¬

tions or specirnens known to us. Flowever, some

very old specirnens labelled “Marc d'Issy, Sénégal\

Bocall AS 11" were obrained from the Muséum

national d'PIistoire naturelle. Unlortunately, they

were too brittle for a definitive identification but

the relative shortness of the vaginal duct distin-

guished them from A. fenestrata and leaves open

the spéculative possibility they are the same as

the Guadeloupe "market leech”

The exact spccics of leech introduced from

Senegambia in the 1820 s is so far unclear from

historié records. In fact, Vire)' (1829) records

that “Sénégal has several hirudinid species in its

ponds, lakes and streams”. One candidate, howe-

ver, which should be eliminated, is a virtually

unknown species described by Henry, SéruJlas

and Vire)' in 1829 as Sanguisuga mysomelas from

“Sénégal, particularly in lakes Mboroo and

Nghier” (Virey 1829; Moquin-Tandun 1846: 14,

340), but apparently it has not been recorded

since. Interestingly, it was noted about this spe¬

cies that doctors at the time “must always use

double those of Europe for the same amount of

blood drawn” (Virey 1829).

ERRATIC DISTRIBUTION OF MEDICINAL LEECHES

on West Indian Islands

One of the inexplicable findings concerning

médicinal leeches in the West lndies is their erra-

tic distribution, being prolïfic on some islands

but totally absent on others. For example, such

leeches are known to occur on Martinique,

Guadeloupe (Pointier et ai 1988), Dominica, St

Lucia, Haiti, Antigua (Sawyer & Kinard 1980),

and Puerto Rico (Moore 1901; Sawyer & Kinard

1980). At the same time, it appears to be absent

from Jamaica, Barbados, St Maarten/St Martin,

Surinam and French Guyana, as well as from

Guyana, Trinidad, Bahamas and probably Cuba.

It is still unclear whether the reason(s) for this

unusual distribution are historical or ecological,

or both.

It is almost certain that leeches hâve been ship-

ped freely between islands for médicinal pur-

poses' for the past 150 years, a practice that is still

going on. During periodic visirs to the West

lndies région from 1974 to 1998. Sawyer ubtai-

ned crédible oral évidente that in recent times

leeches are still being shipped for médicinal pur-

poses to certain leech-free islands, as well as to

the mainland ot South America. For example,

local residents daim that a leech is still imported

from time to time into St Martin/St Maarten

from "another island”, and into Trinidad repor-

tedly from “Granada”. As late as 1976, leeches

were being shipped regularly from Martinique,

reportedly “from the ponds near the airport", to

Cayenne, French Guyana, but by 1993 no

leeches were apparently being imported into

French Guyana (Sawyer 1993, personal observa¬

tion).

Acknowledgements

This study began over twenty-five years ago in

1974 during a trip by R. T. Sawyer to Puerto

Rico where be first encountered and was per-

plexed by a large aggressive leech unlike any

other in the New World. It took many years to

prove, to our satisfaction, the true identity of the

leech and to détermine when and by what

mechanism it had arrived in the West lndies.

468

ZOOSYSTEMA • 1998 ■ 20(3)

Médicinal leeches from the West Indies

Along the way many people contrihuted to

pièces of the puzzle. We especially thank

N. Barré of the Département d’élevage et de

médecine vétérinaire CIRAD-EMVT, Pointe-à-

Pitre, Guadeloupe, and C. M. Moiser, Plymouth

College of Further Education, England for col-

lecting live spécimens of leeches from

Guadeloupe and Gambia, respectively; and to

Prof. J.-L. Justine, Laboratoire de Biologie

Parasitaire, Protistologie, Helminthoiogie,

Muséum national d Histoire naturelle, Paris, for

furnishing préserved specimens from

Guadeloupe and Sénégal. In Martinique, R. T.

Sawyer is particularly grateful to M.-J.

Lamorandière, Sofitel Bakoua, Les Trois îlets,

and to M. Tanasi, Office National des Forêts,

Fort-de-France for exceptional assistance in

obtaining specimens of the Martinique leech in

the “dry” season. We are also very grateful to

W. F. Kinard, Universily of Charleston; R.

Munro, Swansca, Wales; J. P. Pointier, Centre de

Biologie et d Ecologie Tropicale et Méditer¬

ranéenne, Perpignan, France; M. Théry, Labora¬

toire d’Ecologie générale, CNRS; R, K. Yadav,

Central Council of Indian Medicine, Bombay;

and to Mrs E. F. Thomson, Richmond, England.

For invaluable assistance in Mauritius, R. T.

Sawyer would lîke to thank the following: P.

Sooprayen, Chief Archivât; R. Gopaul, Ministry

of Health; C. Michel and Y. Martial.

REFERENCES

Achard M. j. 1825. — Sur la sangsue officinale, sa

reproduction aux Antilles, etc. Bulletin des Travaux

de la Société de Pharmacie 11 ; 296-300.

Anonymous 1822. — Sur une espèce de sangsue indi¬

gène à Terre-Neuve, et qu’il peut être utile

d’embarquer sur les vaisseux du roi ultérieurement

destinés pour les Antilles. Annales maritimes et colo¬

niales de l'Année J822, 2 (2) ; 561.

— 1824. —Notice sur la sangsue officinale, sa repro¬

duction aux Antilles. Annales maritimes et coloniales

de l Année 1824- 1 (2): 331.

Berger C. & Rey If. 1874. — Répertoire bibliogra¬

phique des travaux des médecins et des pharma¬

ciens de la marine française, 1698-1873. Archives

de Médecine navale Baillière, Paris 1874 : 23-

Bhishagratnu K, K, 1963. — An English Translation

ofTHE SUSHRUTA SAM HIT A based on Original

Sanskrit Text, Vidya Vidas Press, Varanasi, India:

98-105.

Blainville M. H. D. de 1827. — Sangsue, 47 :

240-273 in I.amarck J. B. de, Histoire naturelle des

Animaux sans Vertèbres. Paris.

Blanchard R. 1893. — Révision des Hirudtnées du

Piémont. Bollettino dei Musei di Zoologia ed

Anatomia Cornparata délia R. Università di Torino

8:28

— 1897. — Hirtidinécs des Indes néerlandaises.

Weber's Zoologisrhc Ergebnissc einer Reise in

Nederldnduch Ost- Indien 4- 45

Calve M. 1830. — Rapport de M. Calve, chirurgien

de la marine de première classe, chargé du seivice

par intérim, sur les sangsues du Sénégal Annales

maritimes et coloniales de T Année 1830, 2 (2) :

154-159.

Centre d’Étude de Géographie d’Outre-Mer (ed.)

1982. — Atlas des Départements Français d'Outre-

Mer. CNRS-ORSTOM, Paris, 36 p.

Conseil de Santé 1831. — Extrait d’un rapport du

Conseil de la Guyane Française, sur des expériences

relatives à la reproduction des sangsues. Annales

maritimes et coloniales de l’Année 1831, 45 :

304-308.

Dupuy M. 1830. — Seconde partie du rapport sut la

sangsue médicinale envoyée du Sénégal pour être

naturalisée à la Guadeloupe. Annales maritimes et

coloniales de l'Année 1830, 2 (2) 195-154.

Electricwala A , .Sawyer R T„ Powell Jones C. &

Atkinson T. 1991. — Isolation of thrombin inhibi-

tor from the leech Hirudinaria manillensis, Blond

Coagulation andEtbrinolysis 2: 83-89.

Hagy J. W. 1991. — Mosquitoes, leeches and medici¬

ne in Charleston, South CaroJina (1670-1861).

Blood Coagulation and Etbrinolysis 2; 65-68.

Harding W. A. & Moore J. P. 1927. — Hirudinea.

The rauna of British India, inclue!ing Ceylon and

Burma. Taylor and Francis, London, 302 p.

Harvey R. P., Degryse E.. Stéfani L., Schamber F.,

Cazenave J. P., Courtney M., Tolstoshev P. &

Lecocq J. P. 1986. — Cloning and expression of

cDNA coding for the anticoagulant hirudin front

the hloodsucking ieech Hirudo medicinatis.

Proceedings of the National Academy of Sciences of

the United States of America 83: 1084-1088.

Hessel R. 1881. — Artificial culture of médicinal

leeches and of spectes of Hélix. Bulletin of the

United States Fish Commission 1: 264

— 1884. — Leech culture. Bulletin of the United

States Fish Commission 4: 175, 176.

India Office 1856. —L/PÔt)/1 /89.

— 1864. —V/27/821/9,

—1872. —Pros#50, P/691.

_ 1874. —V/27/821/13.

— 1884. —V/27/821/11.

Lesson ]. P. 1842. — Description d’une nouvelle

espèce de sangsue. Revue Zoologique 1842 : 8.

Leuckart R. & Brandes G. 1901. — Die Parasitesi des

Menschen und die von ihnen herrührenden

Krankheiten. Fin Hand- und Lehrbuch fur Natur-

ZOOSYSTEMA • 1998 • 20(3)

469

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

forscher undÀrzte. Hirudlneen: 535-897, Leipzig.

Livingstone D. 1857. — Missionary Travels and

Researches in South Africa. John Murray, London,

129, 130.

Moore J. P. 1901. — Descriptions of two new leeches

from Porto Rico. Bulletin of the United States Fish

Commission for 1900, 2: 211-222.

— 1939. — Additions to our knowledge of African

leeches (Hirudinea). Proceedings of the Academy of

Natural Sciences of Philadelphia 90: 297-360.

Moquin-Tandon Â. 1846. — Monographie de la

Famille des Himdinées. Baillière, Paris, 448 p.

Munro R., Siddall M., Desser S, S. & Sawyer R.

1991 — Blecding in human voluiueefs from the

bite of the American médicinal leech Macrobdella

décora compared with its Etrropean counterpart

Hirudo ntedictnalis. Comparative Hacmatology

International 1 214-216.

Oka A. 1934. — Comparison des Limnath granulosa

provenant de la Forraosa et de la Martinique.

Proceedings of the Impérial Academy of Japon, Tokyo

10: 2S6-288.

Parliamentary Papers 1861. — Anglo-French

Convention on Emigration of Labour from India to

the French Colonies 65: 251.

Pointier ). P., Théron A» & Imbert-Establet D.

1988. — Décline of a sylvatic focus of Schistosoma

mansoniin Guadeloupe (French Wesrlndies) follo-

wing the compétitive displacement of the snail host

Biomphalaria glabrata by Ampullaria glauca.

Oecologia 75: 38-43.

Public Record Otïîce (1847). — PRO: CO 318/172.

_ (1851). — PRO: CO 318/194.

— (1857). — PRO: CO 318/216.

_ (1859). — PRO: CO 318/223.

— (1862). — PRO: CO 386/188.n.

— (1870). — PRO: ADM 101/233.

Ringuelet R. A. 1976. — Clave para las familias y

generos de sanguijuelas (Hirudinea) de aguas dulces

y terrestres de Mesoamerica y Sudamerica.

Limnabios 1: 9-19.

Savigny J. C. 1820 (1822). —- Système des Annélides.

Paris. Leeches : 105-120.

Sawyer R. T. 1981. — Why we need to save the

médicinal leech. Otyx 16: 165-168.

— 1986. — Leech Biology and Behaviour. Oxford

University Press, Oxford, 1065 p.

— (in press). — The trade in médicinal leeches in the

Southern Indian Océan in the nineteenth century.

Medical Flistory.

Sawyer R. T. & Kinard W. F. 1980. — A checklist and

key to the marine and freshwater leeches (Annelida:

Hirudinea) of Puerto Rico and other Caribbean

islands. Caribbean Journal of Science 15: 83-85.

Scacheri F,., Nitti G., Valsasina B., Orsini G., Visco

C., Ferrera M., Sawyer R. T. &i Sarmientos P.

1993. — Novel hirudin variants from the leech

Hirudinaria manillensis. Amino acid sequence,

cDNA and genomic organization. European

Journal ofBwchemistry 214: 295-304.

Taylor R. M. 1971. — Epidemiology: 442, in Strode

G. K. (ed.), Yellow Fever. McGraw-Hill. New York.

Thomas T. N. 1985. — lndtaru Oversear. A Guide to

Source Materials in the India Office Records for the

Study of Indian Emigration 1830-1950. British

Library, London.

Virey J. ). 1829. — Sur des sangsues du Sénégal.

Journal de Pharmacie 15: 640-646.

Williamson J. 1817. — Medical and Miscellaneous

Observations Relative to the West India Islands.

Oliphant, Waugh and Innés, Edinburgh (two

volumes).

Wills S. 1849. — Hospital of St Felipe and Santiago,

Havana, West lndies. The Lancet 1849 (1): 1^8,

149.

Wood G. B. & Bâche F. 1867. — The Dispensatory of

the United States of America. Lippincott,

Philadelphia.

Submitted on 28 November 1997,

accepted on 3 June 1998.

470

ZOOSYSTEMA • 1998 • 20 (3)

New Pisaniinae (Mollusca, Gastropoda,

Buccinidae) from New Caledonia,

with remarks on Cantharus and related généra

Geerat J. VERMEIJ

Department of Geology, University of California,

Davis, CA 95616 (USA)

vermeij @ geology.ucdavis.edu

Philippe BOUCHET

Muséum national d’Histoire naturelle,

55 rue de Buffon, F-75231 Paris cedex 05 (France)

bouchet@mnhn.fr

Vermeij G. J. & Bouchet P. 1998. — New Pisaniinae (Mollusca, Gastropoda, Buccinidae)

from New Caledonia, with remarks on Cantharus and related généra. Zoosystema 20 (3) :

471-785.

KEYWORDS

New Caledonia,

bathyal,

seamount,

Buccinidae,

new taxa.

ABSTRACT

The généra Cantharus Rôding, 1798, Pollia Gray in Sowerby, 1834, and

Cancellopollia n.g. (type species: C. gracilis n.sp.), are pisaniine buccinids

having a small tooth (labral spine) at the edge of the crenulated outer lip. As

defmed and restricted here, these généra hâve a mainly I ndo-West Pacific

distribution. Cantharus septemcostatus n.sp., Pollia pcllita n.sp., Cancellopollia

gracilis n.sp., and C. ustulata n.sp., are reported from deep water in the New

Caledonia région, and Cantharus leucotaeniatus Kosuge, 1985 and Pollia vic-

dani (Kosuge, 1984) n.comb. are reported from Vanuatu. Despite a narrow

bathymétrie (415-560 m) and horizontal (northernmost Norfolk Ridge) dis¬

tribution, Cancellopollia gracilis exhibits remarkable variation, with highly

localised morphs.

ZOOSYSTEMA • 1998 • 20(3)

471

Vermeij G. J. & Bouchet P.

MOTS CLÉS

Nouvelle-Calédonie,

bathyal,

mont sous-marin,

Buccinidae,

nouveau taxon.

RÉSUMÉ

Pisaniinae (Mollusca, Gastropoda, Buccinidae) nouveaux de Nouvelle-Calédonie

et remarques r«r Cantharus et genres apparentés. Les genres Canthams Roding,

1798, Pallia Gray in Sowerby, 1834, et Cancellopollia n.g. (espèce-type :

C. gracilis n.sp.) sont des Buccinidae Pisaniinae caractérisés par la présence

d’un petit denticule (épine labiale) faisane saillie au milieu des annulations

de la lèvre externe. Tels qu’ils sont redéfinis et restreints ici. sur la base de

leurs espèces-types, ces genres ont essentiellement une distribution Indo-

ouest Pacifique, Les espèces Canthams septemcostatus n.sp., Pollia pellita

n.sp., Cancellopollia gracilis n.sp. et C. ustulata n.sp. sont signalées des pentes

bathyales de la région néo-calédonienne, et Canthams leucotaeniatus Kosuge,

1985 et Pollia viedani (Kosuge, 1984) n.comb, sont cités du Vanuatu. On ne

connaît aucun signalement du genre Canthams 'a l'est des îles Fidji, où il est

présent dans le Pléistocène, mais n'est pas connu dans P Actuel, Malgré sa

répartition limitée à une étroite bande bathymetrique (415-560 m) dans

l'extrême nord de la Ride de Norfolk, Cancellopollia gracilis présente une

variation remarquable, avec des formes morphologiquement reconnaissables

très étroitement localisées sur des monts sous-marins isolés.

INTRODUCTION

The Pisaniinae comprises a large subfamily of

mainly tropical carnivorous buccinid neogastro-

pods. Although Cemohorsky (1971, 1975) pro-

vided an extensive review of Indo-West Pacific

species, the generic classification of the

Pisaniinae requires révision, and several deep-

water taxa remain to be describcd. In the very

extensive material collectcd by the MUSORS-

TOM expéditions to New Calcdonia and nearby

régions, several hitherto undescrihed species rela-

ced to the généra Canthams and Pollia hâve corne

to light. Our purpose here is to describe these

species, to propose the new genits Cancellopollia

for two of them, and briefly to comment on the

limits and distinguishing features of Cantharus

and Pollia.

The names Cantharus Roding, 1798 (type spe¬

cies: Buccinum tranquebaricum Gmelin, 1791),

Pollia Gray in Sowerby 1834 (type species;

Buccinum undosum Linnaeus, 1758), and

Tritomdea Swainson, 1840 (type species:

Buccinum undosum Linnaeus, 1758; thus an

objective synonym of Pollia) hâve been applied

to a diverse array of Paleogene to Recent gastro-

pods with ovate to fusiform shells, 3 convex cre-

nulated outer lip, and sculpture consisting of

strong spiral cords Crossing strong to obsolète

axial ribs or folds (sec, e.g., Bellardi 1872;

Cossmann 1901; Pcyrot 1927; Wenz 1941;

Glibcrt 1963; Cemohorsky 1971, 1975). This

broad and inconsistent usage stems from the fai-

lure ol most earlier autbors to take into account

important shell characters rhat can serve to dis-

tinguisb a number of disrina: clades. Although

we do not wisb to attempt a complété generic

révision here, we take the oppommity to rede-

finc Cantharus and Pollia in order to place the

new genus Cancellopollia in the proper contcxt.

We restrict the genus Cantharus to pisaniine buc-

cinids with the following characters; shell ovate;

releoconch consisting of six or more shouldered

whorls separated by a dceply impresscd suture;

base of last whorl constricred above short sipho-

nal canal; sculpture consisting of strong. widely

separated axial folds, strongest on adapica! part

of whorls and fading helow, crossed by strong

spiral cords and threads o( varions sires; siphonal

fasciole prominent, sculptured svitli fine spiral

threads, demarcatinga narrow umhilical «lit rhat

in many species is closed; outer lip weakly

convex in profile, crenulated ai edge; fourrh or

fifth crenulation from abapical end of outer lip

enlarged to form blunt labraJ tooth or spine, for-

med in adult shell at termination of external

groove; external terminal varix absent; inner side

472

ZOOSYSTEMA * 1998 • 20(3)

New Pisaniinae from New Caledonia

of outer lip Iirate, the number of lirae correspon-

ding with the number of external spiral cords;

columella smooth; inner lip adhèrent or wealdy

erect on abapical two-thirds, marked by a few

ridges on adapical sector and by a weak pariétal

rib at its adapical end; pariétal rib flanked on

outer lip by one or cwo slighdy enlarged tooth-

like lirae; adapical end of aperture broad.

As restricted bere, the genus Cantharus is exclusi-

vely Indo-West Pacific in distribution. lt occurs

along continental nvargins and adjacent to high

islands. Living species are known from the

Indian Océan east to the Philippines, north-

eastern AuStralia, New Caledonia and Vanuatu

(see below). During the Pliocène and Pleis-

tocene, the genus was also represenred in

Okinawaand Fiji (see below).

The genus Pollia differs front the closely related

Cantharus by having a more fusiform shell, ada-

pically more tapered and narrower aperture, and

a stronger pariétal rib; by the presence of an

external terminal adule varix; and by having a

very small labral tooth located midway along the

edge of the outer lip instead of on its abapical

one-third. The labral tooth is absent on many

individuals of the type species, P. undosa , which

also differs from other species of Pollia by having

obsolète axial folds. Most species ol Pollia occur

in the Indo-West Pacific. The very widely distri-

buted P fumosa (Dillwyn, 1817). however,

extends east to the Galapagos Islands; and P ver-

meuleni (Knudsen, 1980) is a West African spe¬

cies.

Most other species that have been assigned to

Cantharus and Pollia (or its synonym Tritonidea)

lack a labral tooch or, in tire case ol some tropical

eastern Pacific species of Muricantbarus Olsson,

1971 (type species; Pseudoneptunea pana mica

Hertlein et Strong, 1951), have a small labral

rooth formed as an extension ol a spiral cord.

Although Cernohorsky (1975) synonymized

Muricantbarus and other taxa front tropical

America and the eastern Atlantic with Cantharus ,

species from these régions belong to généra other

than Cantharus and Pollia. Similarly, none of the

many species assigned to Cantharus and Pollia

(or Tritonidea) Irom Cenozoic strata in Europe,

West Africa, or the Americas belongs to

Cantharus or Pollia.

Arrreviations

AMS Australian Muséum, Sydney

MNHN Muséum nauonal d’Histoire naturelle, Paris

NM Natal Muséum, Pietermaritzburg

NMNZ Muséum of New Zealand Te Papa Tonga-

rewa, Wellington

dd dead specimen

Iv live specimen

SYSTEMATICS

Class GASTROPODA

Superfamily BUCCINOIDEA Rafinesque, 1815

Family BUCCINIDAE Rafine.sque, 1815

Subfamily PISANIINAE Gray, 1857

Genus Cantharus Roding, 1798

Cantharus septemcostatus n.sp.

(Figs IA. 2A, 3A, 7A)

Type MATEIUAL. — Holotvpe in MNHN; paratypes: 2

(MNHN), 1 (AMS), 1 (NMNZ), 1 (NM).

TYPE EOCALITY. — North of New Caledonia,

19°05'S, 163°22'E, 220-225 m (R.V. A lis, SM1B 6,

stn DW129).

EtYMOLOGY. — Latin septem, seven, and costatus , rib-

bed, wilh reference to the axial sculpture.

MA l’ERLAl EXAMINED. — North of New Caledonia.

MUSORSTOM 4, stn CPI 52, 19°06’S, 163°22’E,

223 m, I dd. — Stn CP189, 19°07’S, 163“29'E,

210 m, f juv. dd.

LAGON, stn 1147, 19°08'S. 163°30'E, 210 m,

2 dd. — Stn 1148, 19°07'S, 163”30'E. 220 m, 1 Iv

(paratype MNHN).

SM1B 6, stn DW 108, ^O/’S, 163°30'E,

210-220 m, 1 dd. — Stn DW110, 19°05’S,

163“30'E, 225-230 m, 1 Iv, 1 dd. — Stn DWI13.

19°03’S, 163°30’E, 250 m, 1 lv (paratype

MNHN). — Stn DWI27, 19°07’S, 163°23’E,

190-205 m, 1 d.f. - Stn DW128, 19°06’S,

163°22'E, 205-215 m, 2 dd. — Stn DW129,

19°05'S, 163 U 22’L, 220-225 m, 3 lv (paratypes AMS,

NMNZ), 4 dd (holotype, paratype NM). —

Stn DW 130, 19°05’S, 163 D 21’E, 225-230 m, 1 lv,

3 dd.

BATHUS 4, stn DW934, 19°05’S, 163°29’E,

231-240 m, 2 lv, 1 dd.

DISTRIBUTION. — North of New Caledonia, alive in

220-250 m.

ZOOSYSTEMA • 1998 • 20(3)

473

Vermeij G. J. & Bouchet P.

Fig. 1. — Cantharus-, A, C. septemcostatus n.sp., north of New Caledonia, holotype, 36.2 mm; B, C. sp., Nakasi Beds, Early

Pleistocene (1.8 Ma); Saunilambu. 18'’02'S. 178°29'E, Viti Levu, Fiji, Kohn coll. 1982, 28 mm; C, C. leucotaeniatus Kosuge. 1985,

Pta Engano, Cebu, Philippines, leg. Kosuge (MNHN), 47.5 mm.

474 ZOOSYSTEMA ■ 1998 • 20(3)

New Pisaniinae from New Caledonia

Fig. 2. — Opercula; A, Canlharus septemcostatus n.sp., north of

New Caledonia, SMIB 6, stn DW128, 12.3 mm;

B. Cancellopollia gracilis n.sp.. Norfolk Ridge, Stylaster Bank,

CHALCAL 2, stn DW76, 8.1 mm.

Description (Holotype)

Shell large, ovate, consisting of 1.2 prococonch

and 6.0 teleoconch whorls, constricted slightly

just above siphonal canal, spire moderately high.

Protoconch (Fig. 4A) smooth, with five or six

crowded, hroad axial ribs betore terminal varix,

protoconch-telcoconch discontinuity sharply

demarcated. Spiral cords appcar along with less

crowded axial ribs on First teleoconch whorl;

sculpture of last whorl consisting ol seven widely

separated axial Poids, strongest adapically, crossed

by four primary and numerous secondary spiral

cords; whorl broadest at level of adapical cord,

which forms shoulder; lourth (abapical) cord

separated Irom the other three primary cords.

Outer lip edge erect beyond last-lormcd axial

fold, which does not form a distinct terminal

varix; lip edge with eleven crenularions, the four-

th from the abapical end being larger than the

others and forming a short, blunt, labral spine;

this spine located at termination of external

groove immediately above fourth primary spiral

cord. Siphonal fasciole rounded, sculptured with

about twelve fine spiral threads; umbilical slit

absent; inner side of outer lip lirate, the two ada-

pical lirae enlarged as denticles opposite very

weak pariétal rib at adapical end of inner lip;

columella smooth, more or less straight, with

fold at base of siphonal canal; inner lip slightly

erect along abapical two-thirds of its length,

adhèrent on adapical one-third.

Background colour creamy beige, axial ribs light-

ly tinted with rust brown especially at shoulder

angulation, two rniddle spiral cords white at their

intersection with axials. Aperrure and columella

white. Rather thick periostracum with hairy pro¬

jections aligned along incrémental Unes, hairs

longest on main spiral cords.

Dimensions: holotype height 36.2 mm; diameter

23.1 mm; aperture heighr 23.6 mm; aperture

width 12.0 mm. Other material up to 40.7 mm

in height.

Remarks

Canthdrus septemcostatus is closelv related to two

other species, C. leucotaeniatus Kosuge, 1985,

from ihe Philippines, and C. okinawa MacNeil,

1961, from the Pliocène of Okinawa. Together,

chese three species form a group characterized by

having spiral sculpture consisting of four primary

cords, ol which the niost abapical (fourth) cord is

remote from the other three. A groove ending at

the outer lip in a blunt labral spine lies immedia-

rely above the fourth (counted from adapical

end) primary spiral cord.

Cantharus leucotaeniatus (Fig. IC) differs from

C. septemcostatus by having a larger number of

axial folds (ten to twelve as compared to seven on

the last whorl), a larger number of crenularions

on the outer lip (eighteen as compared to ele¬

ven), and by having the second instead of the

first primary spiral cord (counting from the su¬

ture) marking the broadest point of whorl.

C. leucotaeniatus also reaches a slightly larger size

(6U mm vs 40.7 mm) and is more vividly colou-

red, with spiral cords dark brown ovet a white

background. C. okinawa is cxtrcmely similar to

C. leucotaeniatus , but has ninc or ten axial folds

(compared to ten to twelve), twelve instead offif-

teen to eighteen crenularions and lirae on the

outer lip, and an adhèrent instead of abapically

minutely erect inner lip. The fossil species

reaches a maximum length of only 29.4 mm âs

compared to 60 mm for C. leucotaeniatus. It may

be that, as more material becomes available,

C. leucotaeniatus will prove to be a junior syno-

nym of C. okinawa. MacNeil (1961) described

ZOOSYSTEMA • 1998 • 20(3)

475

Vermeij G. J. & Bouchet P.

C. okinawa from the Shinzato Tuff (Pliocène) in

the Shimajiri Group of Okinawa A. J. Kohn

(pers. comm.) has collected addicional specimens

from the underlying Yonabaru Formation (Early

Pliocène) in Okinawa. Examination of these spe¬

cimens reveals that they agréé in every detail wirh

MacNeils shells. C okinawa difFers from C. sep-

temcostatus by having a larger number of axial

folds (nine or ten instead of seven), by having the

inner lip adhèrent instead of abapically mtnutely

erect, and by being somewhat smaller (height

29.4 mm instead of 40.7 mm).

A probable fourth species in this group is repre-

sented by a single well-preserved fossil specimen

(Fig. IB) from the Pleistocene of Fiji, collected

by A. J. Kohn. Although the spire is broken, the

last whorl is perfectly preserved. It bears eight

sharply rounded axial folds, crossed by six prim-

ary spiral cords. The groove terminating at the

outer lip in a blunt labral tooth is located just

above the lowest primary spiral cord. There are

eighteen annulations and lirae on the outer lip,

of which four are situated below the annulation

that is enlarged to form the labral tooth. This

specimen represents the easternmost occurrence

of the genus Cantharus. Widi only a single spéci¬

men at hand, we prefer to postpone formai des¬

cription until more material becomes available.

Cantbanis leucotaeniatus Kosuge, 1985

(Fig. IC)

C. albozonatus Kosuge, 1983: 137 (non Cantharus

[ Tritonidea ] albozonatus Smith, 1890).

Cantharus leucotaeniatus Kosuge, 1985: 20 (nom.nov.

pro C. albozonatus Kosuge, 1983, non Smith, 1890),

NEW RECORD. — Vanuatu. Off Espiritu Sanro,

MUSORSTOM 8, stn C.P 1102, 15' J 04'S, 167°09’E,

208-210 m, 1 Iv juv.

This new record represents a considérable extension of

the known range, otherwise restricted to the

Philippines (Springsteen & Leobrera 1986: 166).

Genus Pollia Gray in Sowerby 1834

Pollia pellita n.sp.

(Fig. 4A-C)

Type material. — Holotvpe (ly) and 2 paratypes

(dd) in MNHN.

TYPE I.OCAUTY. — New Caledonia, Loyalty Ridge,

20°42’S, 167°00’E, 270 m (R.V. /Km/MUSORS-

TOM 6, stn CP400).

ÊTYMOLOGY. — Latin pellitus (adj.), dressed with furs;

with reference to the thick hairy periostracum of

livîng specimens.

Fig. 3. — Protoconchs; A, Cantharus septemcostatus n.sp., north of New Caledonia, SMIB 6, stn DW129: B, Cancellopollia gracilis

n.sp., Norfolk Ridge, Stylaster Bank, CHALCAL 2, stn DW76. Scale bars: A, B, 0.5 mm.

476

ZOOSYSTEMA • 1998 • 20 (3)

New Pisaniinae from New Caledonia

Fig. 4. — Pollia; A-C. P. pellita n.sp.; A, B, Loyalty Ridge, holotype, 26.5 mm (A) and paratype, 29 mm (B); C, Coral Sea,

MUSORSTOM 5, stn 260, 33.5 mm; D, P. vicdani (Kosuge, 1984), Vanuatu, off Espiritu Santo, MUSORSTOM 8, stn DW1097,

19.8 mm.

ZOOSYSTEMA • 1998 • 20 (3)

477

Vermeij G. J. & Bouchet P.

MATERIAL EXAMINEP. — Loyalty Ridge. MUSORS-

TOM 6, stn CP400, 20°42’S, 167°00'E, 270 m, 1 Iv

(holotype), 2 dd (paratypes).

Coral Sea. MUSORSTOM 5, stn 260, Capel Bank,

25°59’S, 159°44’E, 285 m, 1 dd. — Stn 275, Argo

Bank, 24°47’S, 159 o 40’E. 285 m, I iv.

Distribution. — Loyalty Ridge and Lord Howe

Rise, alive at 270-285 m. Nor round around New

Caledonia proper or on the Norfolk Ridge.

Description

(Type material front Loyalty Ridge, Fig. 4A, B)

Shell up to 29 mm in height, slender, fusiform,

consisting of 2.0 protoconch and up to 6.1 teleo-

conch whorls. Protoconch diameter 2000 pm,

exposed height 2080 pm, high, bulbous, with

small nucléus, diameter 250 pm, whorls convex

with deep suture, smooth, protoconch-teleo-

conch discontinuity distinct. Teleoconch spire

whorls evenly convex on most exposed part, aba-

pical part wirh narrow concave gutter, suture

impressed, last whorl evenly convex to tapering

base. Sculpture consisting of strong, broad axial

ribs, crossed over by well-defined spiral cords.

Eight axial ribs on ali teleoconch whorls, inter-

spaces narrower than ribs, with strong incrémen¬

tal lines. Spiral cords uneven, narrower than

interspaces, main cords increasing Iront four on

first whorl to nine on penultimate whorl, inter¬

spaces with one or two thinner secondary cords;

one cord stronger, siruated low on spire whorls,

overhanging suture, equal to other cords and

situated on periphery on last whorl; last whorl

with ca. nventy primary cords extending onto

siphonal canal, interspaces with one to four

(most frequently one to three) secondary cords,

some of these almost as strong as primary cords.

Outer lip erect beyond external terminal varix,

with thirteen internai lirae, adapical and abapical

crenulations slightly enlarged, tenth lira (counted

from adapical side) prolonged as a small labral

tooth; inner lip erect, with ai. filteen lirae corres-

ponding to position of main cords below, parierai

wall marked by two thickened folds, coluntella

with bîfid terminal swelling; aperture broad*

tapering adapically; siphonal canal short but dis-

tinctly set off. No umbilicus. Background colour

orange brown, spiral cords darker, white spiral

band just below periphery. Periostracum very

thick, tightly adhering, with long hairy expan¬

sions aligned aloiig spiral cords.

Holotype height 26.6 mm, diameter 14.5 mm;

aperture height 15 mm, aperture width 7 mm.

Remarks

The two spécimens from the Coral Sea (Fig. 4C)

dilfer from the three type .specimens by reaching

a slightly larger size (up to 34 mm), details ol the

sculpture (nine, rather than eight, axial ribs per

whorl), and a sharper colour pattern of dark

brown ribs contrasting against light background.

These différences could be indicative of genetical

isolation of the two groups of populations, but

the material is ton scanty to speculate further on

these différences, We prefer to provisionally treat

them as geographical variants of a single biologi-

caJ species.

Pollia pellita differs from P. viedani (Fig. 4D;

Bouchet & Warén 1986: ftg. 63) by being pro-

portionally distinctly broader ( pellita

h/D = 1.77-1.90, mean 1.82, n = 5; viedani

h/D = 1.95-2.15, mean 2.03, n = 9) and with a

smaller rib number (eight or nine vs fifteen per

adult whorl in specimens of 26-34 mm).

Pollia viedani (Kosuge, 1984) n.comb.

(Fig. 4D)

Cantbarus viedani Kosuge, 1984: 146, pl. 49, figs 6-9.

New records. —Vanuatu, Off Espiritu Santo,

MUSORSTOM 8, stn DW1097, 15°05’S, 167°11’E,

281-288 m, 1 dd. — Stn DW1106, 15°05’S,

167°12'E, 305-314 m, 2 dd.

Remarks

We have not exantined the type material (holoty¬

pe from Bohol, Philippines, in 220 m), but we

have seen several lots from the I’hilippines

[MNHN (Bouchet & Warén 1986. 471. fig. 63,

as Cantharus délicat us), NM] thaï conform the

original description and illustration. The status

of Cantharus viedani in relation to Tritunidea

delieata Smith, 1899 is unclear. Bouchet &

Warén (1986: Ftg. 62) illustrated the holotype of

T. delieata from the Gulf of Bengal in 165 m,

and recorded the species from the Mozambique

Channel and the Philippines. We have reexami-

ned the Philippines material, which differs from

478

ZOOSYSTEMA • 1998 • 20(3)

New Pisaniinae from New Caledonia

the type materia! ol T. delkata by its sttong axial

sculpture extetiding over the whole whorl heighr.

The general shell proportions are however similar

and these spécimens ntigiit represent local

variants of a single biologica! spccies. Without

further évidente, we prefer to treat the

Philippines specimens as distinct from those

from the Gulf of Bengal, and identify them as

C. viedanr. The Vanuatu shells beat a general

resemblancc- to the Philippines shells, but appear

to be adult at a smaller size (19.8-20.5 inm vs up

to 39.6 mm) and with a correlatedly lower num-

ber (ten) of axial ribs per adult whorl. They also

hâve a more bulbous protoconch Obviously,

more material is needed from intermediate loca-

lities to evaluate the status of the various popula¬

tions.

We transfer Qmtharus viedani to Pollia , based on

the position of che obsolète tooth in the middle

of the outer lip of Philippines specimens.

Genus Cancellopollia n.g.

Type SPECIES. — Cancellopollia gracilis n.sp.

ETYMOLOGY. — Latin cancelli, lattice; and suffix

Pollia, suggesting close relationship to Pollia ; gender

féminine.

DlAGNOSIS

Pisaniine buccinids of medium size having fusi-

form shell; teleoconch whorls evenJv convex,

separated by shallow suture; sculpture on last

whorl consisting ot numerous (eighteen or more)

spiral cords, which cross fifteen or more axial

ribs, giving the surface a cancellatc appearance.

Umbilical slit absent. Outer lip convex in profile,

its edge crenulated, one to three crenulations

near midpoint of outer lip slightly enlargcd to

form very blunt labral tooth; edge ot outer lip

erect, extending slightly beyond external terminal

adult varix; inner side of outer lip bearing short

lirae; inner lip adhèrent; columella smooth, with

fold at base of siphonal canal; aperture narrow,

tapering adapically, about three times as high as

wide; pariétal rib at adapical end ot inner lip pro¬

minent.

Radula with central tooth moderately arched,

with three strong cusps medially and eventually

smaller cusps marginally; lacerais with large,

strong cusp between the inner and outer cusps.

Remarks

The nevv genus Cancellopollia consists of deep-

watet spccies that appear to be elosely related to

Pollia. Characters in common with Pollia include

the terminal adult varix, the enlarged centrally

placed crenulations on the edge of the outer lip,

the fusiform shape, and the tapeted adapical end

of the narrow aperture. Boeh généra hâve a pro¬

minent pariétal tooth. Cancellopollia differs front

Pollia by the much finer and mote numerous spi¬

ral cords and axial riblets, by having a higher

spire and more slender torm, by having a relari-

vely narrower aperture, and by having tewer lirae

on the inner side ot the outer lip than there are

spiral cords. In Pallia , the number of lirae rough-

ly corresponds to the number of cords, The spe-

cies currently assigned to Cancellopollia bave a

bulbous and pauçispiral protoconch, indicating

non-planktotrophic development, whereas those

of species assigned to Pollia are generally multi-

spiral, indicating planktotrophic larval develop¬

ment. However, we do not consider the mode of

larval development to be a generic character, and

we do not exclude that further deep-water explo¬

ration may lead to the discovery of species of

Cancellopollia with multispiral protoconch.

Buccinum cinis RêeVe, 1846, from the Galapagos,

which has alrèadv been placed in Alonostiolum

Dali, 1904 (Keen 1971), Caduaftr Dali, 1904

(Skoglund 1992). and a new unnamed genus

(Cernoliorsky 1975), is also remotely similar to

species of Cancellopollia. h rescmblcs them in

having a slender fusiform shell, relatively nume¬

rous (eighteen) spiral cords on the last whorl, an

external terminal adult varix, a hrate outer lip,

and a prominent pariétal rib. lt differs, however,

by the structure of the spiral sculpture, in which

main spiral cords are flanked by secondary cords,

plus spiral chreadlets in between; two twin cords

form a thick subsutural fold. Axial riblets are

absent, instead the spiral cords are ornamented

by about rwenty-five granules on the last whorl.

The inner lip is raised, with irregular thickened

wrinkles, and there is a strong anal tooth on the

outer lip. For these reasons, we believe that the

ZOOSYSTEMA • 1998 • 20(3)

479

Vermeij G. J. & Bouchet P.

resemblance of Buccinum cinis to species of

Cancellopollia is superficial, but wc cannot sug-

gest a proper generic allocation.

Cancellopollia gracilis n.sp.

(Figs 2B, 3B, 5A, 6, 7C, D)

Cantharus sp. - Cernohorsky 1981: 998, pl. 3, fig. 21.

Type MATERIAL. — Holorvpe in MNHN; paratypes: 2

(MNHN), 1 (NMNZ), 1 (AMS), 1 (NM).

Type LOCALITY. — South of New Caledonia, S ry las ter

Bank, 23°38’S, 167°43’E, 435 m (R.V. Coriolis,

CHALCAI. 2, stn DW77).

Etymoloc.y. — Latin gracilis (adj.), graceftil.

MATERIAL EXAM1NED, — New Caledonia slope.

Vauban 1978-1979, stn 15, 22°49’S, 167°12 E,

390-395 m, 1 Iv (Cernohorsky 1981).

BIOCAL: stn CP52, 23°06'S, 167°47’E, 540-600 m.

1 Iv.

MUSORSTOM 4, stn DW212, 22°47’S, 167 o 10T,

375-380 m, 2 Iv. — Stn DW220. 22°58 S, 167°38'E,

505-550 m. 3 Iv, 1 juv. dd. — Stn DW221, 22°59’S,

167 Ü 37’E, 535-560 m, 8 lv, 1 dd. — Stn DW222,

22°58'S, 167°33’E. 410-440 m, 3 lv, 1 dd. —

Stn DW230, 22°52’S, 167°12'E, 390-420 m, 1 juv.

lv, 1 dd.

SMIB 1, stn DW2, 22°52’S, 167“13’E. 415 m, 2 lv,

1 dd.

SMIB 2, stn DW1, 22 0 53’S, 167°13’E. 438-444 m,

1 dd. — Stn DW3, 22°56’S. 167 D 15’E. 412-428 m,

2 lv. 2 dd, — Stn DW6, 22°56’S, 167°16’E.

442-460 m, 1 lv. — Stn DW8, 22“54'S, 167°13’E.

435-447 m, 1 lv. — Stn DW9. 22 D 54 ! S, 167°I5'E,

475-500 m, 1 lv, 1 dd. — Stn DW17, 22°55'S,

167° 15’E, 428-448 m, 2 lv.

SMIB 3, stn DW21, 22’ 5 59'S, 167°19’E, 525 m,

2 dd. — Stn DW22, 23°03'S, 167°19’E, 503 m,

3 dd. — Stn DW26, 22°55’S, 167°16'E, 450 m, 2 lv.

SMIB 8, stn DW193-I96, 22 o 59’-23°00’S,

167 0 21’-167 0 23'E, 491-558 m, 6 lv, 6 dd. —

Stn DW197-199, 22"5 1 ’-22''52’S, 167"12’E,

408-436 m, 5 lv, 22 dd. — Stn DW200, 23°00’S,

167 n 21’E, 514-525 m. 2 dd.

BATHUS 2, stn DW729, 22°52’S, 167°12’E, 400 m,

5 lv, 2 dd.

Aztèque Bank. SMIB 8, stn DW189, 23' , 18’S,

168°06'F., 400-402 m, 1 dd.

BATHUS 3, stn DW829, 23 0 21’S, 168°02’E,

386-390 m, 3 dd. — Stn DW830, 23°20’S,

168°0TE, 361-365 m, 1 dd.

Stylaster Bank. CHALCAL 2, stn CP25, 23°39'S,

167°43’E, 418 m. 1 lv. — Stn DW76, 23“4l’S,

167°45'E, 470 m. 6 lv (paratypes AMS, NMNZ,

NM), 7 dd. — Stn DW77, 23°38’S, 167°43'E,

435 m, 6 lv (holotype). 1 dd.

SMIB 3, stn DW 12, 23°38'S, 167°42'E, 470 m,

5 dd. — Stn DWT3, 23°38'S, 167°42’E, 448 m, 2 Iv

(1 patatvpe MNHN), 1 dd.

SMIB 8, stn DW 166, 23°38S, 167°43’E, 433-

450 m, 2 lv, 2 dd, 3 juv. dd. — Stn DW4 67,

23°38 S, I67”43’E. 430-452 m, 4 lv. 2 juv. Iv, 3 dd,

1 juv. dd- — Stn DW168, 23°48’S, 167 C 43'E,

433-450 m, 2 lv. 3 dd. — Stn DW169, 23 D 37‘S,

l67 a 42'E, 447-450 m, 1 lv, 3 juv. lv.

BERYX 11, sut DW'27, 23“37’S, l67 0 4rE,

460-47Ü tn, 2 lv ( 1 patatype MNHN), 7 juv.

Iv/dd. — Stn CP32, 23"38’S, I67°43'E, 420-460 m,

I lv. — Stn DW38, 23"38'S, l67 a 39’E, 550-690 m,

4 lv, 14 dd. - Stn DW r 39, 23 Ü 37'S, 167°40’E,

490-500 m, 2 dd.

Norfolk Ridge. BERYX 11. stn DW.34, 23°33’S,

167°17'E, 560-570 m, I lv. ! dd.

DISTRIBUTION. — South of New Caledonia, Norfolk

Ridge. alive at 415-560 m.

Description

Shell up to 31.8 mm in height, slender, fusiform,

cousisting oi 1.2 protoconch and up to 5 teleo-

conch whorls. Protoconch (Fig. 3B) bulbous,

cousisting of 1.2 whorls, paucispiral, smooth,

terminating with a varix followcd by a fraction of

a whorl witli teleoconeh spiral cords sculpture

already présent, and a second discontinuity mar-

king transition to teleoconeh. Teleoconeh whorls

evenly convex separated by indistinct, appressed

suture; sculpture consisting of about forty fine

spiral cords Crossing thirty-five axial riblccs on

die last whorl, and eleven cords Crossing thirty-

one axial ribs on the penultimatc whorl; outer Iip

crenulate, erect beyond external terminal varix;

three centrally placed crenulations on outer Iip

slightly ventraJly and adaxially ertlarged; inner

side of outer lip with thirteen to eiglueen iirae;

inner lip adhèrent, its adapical end marked by a

prominent parierai rib; aperture narrow, tapering

adapically; siphonal canal short; umbilieal slit

absent.

Colour off-white with üghtly adhering, light yel-

lowish beige periostracum, formitig short lamel-

lar expansions along incrémental lines.

Holotype height 30.0 mm, diameter 13.2 mm;

480

ZOOSYSTEMA • 1998 • 20(3)

New Pisaniinae from New Caledonia

aperture height 19.2 mm, aperture width

6.1 mm.

Discussion

Despite a geographical distribution restricted to

the northern part of the Norfolk Ridge,

Cancellopallia gracilis exhibits major variation,

and morphological extrêmes might easily be mis-

taken for separate species (Fig. 6). Numerous

dredge hauts hâve been made between 300 and

800 in in the area considered, and we consider

that the known narrow bathymétrie distribution

Fig. 5. — Cancellopollia-, A, C. gracilis n.sp., Norfolk Ridge, Stylaster Bank, holotype, 30.0 mm; B, C. ustulata n.sp., Coral Sea,

Chesterfield Plateau, holotype. 15.2 mm.

ZOOSYSTEMA • 1998 • 20(3)

481

Fig. 6. — Geogtaphical variants ol Cancellopollia gracilis ri.sp. positloned over three-dimensiûnal bathymetry ot southemmost New

Caledonia and northem Norfolk Ridge. The mode! is positioned with rfie NW al top angle and tde main axis of Norfolk Ridge omning

from the top hght to lower lett. Llght areas are above 800 m deep, dark areas deeper than 800 m. Shell variants are prînted al the

same magnilicalion. A, Stylasler Bank, CHALCAL 2, stn DW77, 435 m, 28.5 mm; B, Norfolk Ridge, BERYX 11, stn DW34,

560-570 m, 19.1 mm; C. New Caledonia slope, SMI6 8, stn DW197-199, 408-436 m, 18.3 mm; D. New Caledonia slope. SMIB 2,

stn DW6, 442-460 m, 17.5 mm; E, Aztèque Bank, BATHUS 3, stn DW&29,386-390 m, 15.6 mm.

(415-560 m) is not a sampling artifact. Within

the horizontal range are four discrète forms sepa-

rated by deeper water. This, in combination with

non-planktotrophic development, explains why

the observed variation appears to hâve a srrong

geographical component.

1. A large number of samples originare ffom rhe

slope south of New Caledonia and the Isle of

Pines where bottoms at adéquate depths form a

continuons belt. AJI specimens hâve in common

a small adulr size (h = 14.9-21.1 mm) and a

broad shell (h/D ratio = 1.78-2.01, mean 1.87,

n = 34), Colour and sculpture are stable within a

sample but show a depth-related variation. In the

shallower part of the range (415-450 m), speci¬

mens hâve a pattern of interrupted brownish spi¬

ral cords (Fig. 6C); the axial ribs are low, almost

obsolète on the last adult whorl. In the deeper

part of the range (475-540 m), specimens hâve a

uniform olT-white shell (Fig. 6D), axial sculpture

is much stronger and extends onto the last adult

whorl.

2. Three samples originare from Aztèque Bank

(Fig. 6E), a seamount to the southcast of the pre-

ceding. This population is characterized by a

small adult size (h = 15.4-16.4 mm), a slc-nder

shell (h/D ratio = 1.97-2.15, mean 2.03, n = 4)

with high conical spire, and a colour pattern

with at least a few brownish spiral cords. Empty

shells only hâve been dredged in 360-400 m.

482

ZOOSYSTEMA • 1998 • 20(3)

New Pisaniinae from New Caledonia

Fig. 7. — Radulae; A, Cantharus septemcostatus n.sp., nortti of New Caledonia, SMIB 6, stn DW128; B, Cancellopollia ustulata

n.sp., Coral Sea, Chesterfield Plateau, MUSORSTOM 5, stn 379; C, C. gracilis n.sp., Stylaster Bank, CHALCAL 2, stn DW76;

D, C. gracilis n.sp., New Caledonia slope, SMIB 3, stn DW26. Scale bars: 100 pm.

suggesting that C. gracilis lives shallower on

Aztèque Seamount than elsewhere.

3. Samples from Stylaster Bank (Fig. 6A) form

an isolated and homogeneous group to the

south. This population is characterized by a large

adult size (h = 28.0-33.7 mm), a slender shell

(h/D ratio = 1.94-2.16, mean 2.09, n = 12) and

a uniform off-white colour with yellowish beige

periostracum.

4. Finally a single sample, with two specimens

(Fig. 6B), originales from an isolated projection

along the Norfolk Ridge, to the Southwest. The

shells look like a miniature of the Stylaster Bank

form, with a much smaller adult size (h = 19.0

and 19.1 mm), broader outline (h/D ratio =

1.93), but similar sculpture and colour.

Cancellopollia ustulata n.sp.

(Figs 5B, 7B)

Type MATERIAL. — Holotype and 3 paratypes in

MNHN.

Type LOCALITY. — Coral Sea, Chesterfield Plateau,

19°53’S, 158"38’E, 400 m (MUSORSTOM 5,

stn 361).

ETYMûIOGY. — From the Latin ustulatus (adj.), a lit-

rle brown as a resuit of burning, with référencé to the

brownish colour patches.

MATERlAl EXAMINED. — MUSORSTOM 5, stn 361,

I9 0 53 S, I58°38’E, 400 m, 1 lv (holotype). —

Stn 379, 19°33’S, 158°40’E, 370-400 m, 2 lv, 1 dd

(paratypes).

ZOOSYSTEMA • 1998 • 20(3)

483

Vermeij G. J. & Bouchet P.

Description

Shell up to 16.0 mm in height, slender, fusiform,

consisting of 1.2 protoconch and five teleoconch

whorls. Protoconch bulbous, paucispiral, smooth,

protoconch-celeoconch discontinuiry distinct.

Teleoconch whorls evenly convex separated by

shallow, impressed suture. Sculpture consisting

of strong, broad axial ribs, crossed over by well-

defined spiral cords. Axial ribs eleven on first

teleoconch whorl, then nine on spire whorls,

twelve on penultimate, and eighteen on last

whorl, interspaces narrower than ribs. Spiral

sculpture consisting of six cords on exposed part

of spire whorls, interspaces broader than cords;

cord 5 (cord 1 = adapical) stronger, situated low

on spire whorls, equal to other cords and situated

on periphery on last whorl; last whorl with nine

evenly spaced ptimary cords, separated by a

slightly broader interspace from another group of

ca. ten cords on shell base, crowded on siphonal

canal; secondary cords présent between pritnary

cords on last whorl. Outer lip crenulate, erect

beyond exrenial terminal varix; one crenulation

slightly enlarged, situated at level of fasciole bet¬

ween two groups of spiral cords; inner side of

outer lip with nine spirallv elongated swellings;

inner lip adhèrent, pariétal wall marked by a

weak but distinct lira, columella with hiftd ter¬

minal swelling; aperture narrow, tapering adapi-

cally; siphonal canal short but disrinctly set off.

No umhilicus.

Colour ofF-white with irregular, axially elongated

brown patch es. Periostracum cighrly adhering,

light yellowish beige, lorming short lamellar

expansions along incrcmental Unes.

Holotype height 15.2 mm, diameter 7.3 mm;

aperture height 6.5 mm, aperture width 3.1 mm.

Discussion

Canceüopollia ustulata differs from C. gracilis by

its smaller adult site, more slender spire with

more convex spire whorls, coarser axial and spiral

sculpture, with fewer ribs and cords, and correla-

tedly fewer apertural lirae It also diffère from the

coloured individuals of C, gracilis by the brown

colour forming distinctly axial blotches, rather

than spirally discontinuous cords.

C. ustulata is known only from the slope of the

Chesterfield Plateau, a mid-Tertiary isolated

structure in the middle of the Coral Sea, separa¬

ted from New Caledonia and Australia by abyssal

depths.

Aclcnowledgements

We thank Emmanuel Bouniot (Service des

Méthodes Administratives et de l’Informatique,

Nouméa) who prepared the 3D bathymetrical

map to our spécifications, based on data provi-

ded by the ZoNeCo programme. Serge Gofas

(MNHN) drew the protoconchs in figure 3,

Anders Warén (Naturhistoriska Riksmuseet,

Stockholm) prepared and scanned the radulae,

and Alan Kohn (Universiry of Washington) pla-

ced hts fosstl material at our disposai.

REFERENCES

Bellardi L. 1872. — I Molluschi dei terreni Terziari

de/ Piernonti e délia Lignria. Pane 1. Gasteropoda

(,Mtirkidae et Tritonidae), Stamperia Reale, Torino,

264 p.

Bouchet P. & Warén A. 1986. — Mollusca

Gastropoda: Taxonomical notes on tropical deep

water Buccinïdac with descriptions of new taxa.

Mémoires du Muséum national d‘Histoire naturelle ,

série A, 133:457-517.

Cernohorskv W. O. 1971. — Indo-Pacific Pisaniinae

(Mollusca: Gastropoda) and related buccinid géné¬

ra. Records of the Auckland Instituer and Muséum 8:

137-167.

— 1975. — Supplementary notes on the taxonomy of

buccinid species of the subfamily Pisaniinae

(Mollusca: Gastropoda). Records of the Auckland

Instituée and Muséum 12: 175-211.

— 1981. — On a collection of buccinacean and

mitracean gasttopods (Mollusca, Gastropoda) from

the Mozambique Channel and New Caledonia.

Bulletin du Muséum national d Histoire naturelle ,

série 4, 3 (A) : 985-1009.

Cossmann M. 1901. — Essais de Paléoconchologie

Comparée, Volume 4. Paris, 293 p.. 10 pis.

Glibert M. 1963. — l.es Muricacea et Buccinacea fos¬

siles du Cénozoïquc étranger des collections de

l'Institut Royal des Sciences Naturelles de Belgique.

Mémoires de l'Institut Royal des Sciences Naturelles de

Belgique, série 2, 74 : 1-179.

Kecn À. M. 1971. — SeashelkafTropical West America.

Stanford University Press, Pulo Alto, 1064 p.

Kosuge S. 1983. — Description of a new species of

the geuus Cantharus (Gastropoda, Buccinacea).

Bulletin of the Institute of Maûcology , Tokyo 1 (9):

137-138.

484

ZOOSYSTEMA • 1998 • 20(3)

New Pisaniinae from New Caledonia

— 1984. — Studies on the collection of Mr. Victor

Dan (6). Descriptions of new species of the généra

Pterochelus , Takia , Muricopsis and Cantharus.

Bulletin of the Institute of Malacology, Tokyo 1 (10):

143-146.

— 1985. — New name for the genus Cantharus.

Bulletin of the Institute of Malacology, Tokyo 2 (2):

20 .

MacNeil F. S. 1961 [“I960”]. — Tertiary and

Quaternary Gastropoda of Okinawa. U.S. Geolo-

gical Survey Professional Paper 339: 1-148.

Peyrot A. 1927. — Conchologie Néogénique de l’Aqui¬

taine, Tome V (1) : gastropodes. Bordeaux, 206 p.

Springsteen F. J. & Leobrera F. M. 1986. — Shells of

the Philippines. Carfel Seashell Muséum, Manda,

377 p.

Wenz W. 1941 [in 1938-1944], — Gastropoda. Teil

I, Allgemeiner und Prosobranchia: 961-1200, in

Schindewolf O. H. (ed.), Handbuch der

Paldozoologie. Borntraeger, Berlin, 1639 p.

Submitted on 12 May 1997;

accepted on 20 March 1998.

ZOOSYSTEMA • 1998 • 20(3)

485

A new species of Tityus C. L. Koch, 1836

(Scorpiones, Buthidae) in Colombia, with

a check list and key to the Colombian species

of the genus

Wilson R. LOURENÇO

Laboratoire de Zoologie (Arthropodes), Muséum national d’Histoire naturelle,

61 rue de Buffon, F-75231 Paris cedex 05 (France)

arachne@mnhn.fr

Lourenço W. R. 1998. — A new species of Tityus C. L. Koch, 1836 (Scorpiones, Buthidae)

in Colombia, with a check list and key to the Colombian species of the genus. Zoosystema

20 (3) : 487-497.

KEYWORDS

Scorpiones,

Buthidae,

Tityus oteroi n.sp.,

Darien,

Colombia,

new species,

check list,

taxonomie key.

ABSTRACT

Tityus oteroi n.sp. (Scorpiones, Buthidae) is described on the basis of five spé¬

cimens (one male and fout females), collectcd in the région of Darién in the

north of Colombia. With the description of Tityus oteroi, the number of spe¬

cies of the genus Tityus known to be distributed in Colombia is raised to

twenty. Some comments concerning the taxonomie position of the new spe¬

cies are added. A check list of Columbian species of Tityus as well as keys for

their détermination are also included.

MOTS CLÉS

Scorpiones,

Buthidae,

Tityus oteroi n.sp,,

Darien,

Colombie,

nouvelle espèce,

« check list »,

clé taxonomique.

RÉSUMÉ

Une nouvelle espèce du genre Tityus C. L. Koch, 1836 (Scorpiones, Buthidae) de

Colombie, avec une liste complète et des clés pour les espèces colombiennes de ce

genre. Tityus oteroi n.sp. (Scorpiones, Buthidae) est décrit à partir de cinq

exemplaires (un mâle et quatre femelles), collectés dans la région de Darién

au nord de la Colombie. Avec la description de Tityus oteroi, le nombre

d’espèces appartenant au genre Tityus, connues pour la Colombie s’élève à

vingt. Quelques commentaires sur la position taxonomique de la nouvelle

espèce sont ajoutés. Sont également incluses une liste des espèces colom¬

biennes de Tityus, ainsi que des clés pour leur détermination.

ZOOSYSTEMA • 1998 • 20(3)

487

Lourenço W. R.

INTRODUCTION

The Colombian scorpion fauna has attracted the

attention of arachnologists since the middle of

the 19th century (e.g. Gervais 1844; Thorell

1876; Pocock 1893; Kraepelin 1912, 1914;

Mello-Leirâo 1945)- Only since the 1980s,

however, and especially during the last ten years

has this fauna been studied intensively. Several

new species hâve been added (Lourenço 1984a,

b, 1991, 1993, 1994, 1995a, b; Lourenço &

Florez 1989, 1990a, b, 1995). A more recent

contribution by Lourenço (1997) represented the

first attempt to produce a summary of this fauna

associated with biogeographical considérations. It

was admitted, however, that the results of tins

summary are tentative, and that a number of

additional species should exist in Colombia. A

much larger inventory will reveal a more realistic

number of taxa présent in the country. At pré¬

sent, a total of four families (Buthidae,

Chactidae, Diplocetridae and Ischnuridae), nine

généra, and forty-one species has been confirmed

for Colombia.

Recent study of old specimens collected in the

région of Darién by F. Geay back in 1899 and

deposited in the collections of the Muséum

national d’FIistoire naturelle, Paris, revealed one

more new species of Tityus in Colombia.

Moreover, because many pcoplc hâve been stung

by scorpions in this country during the last fcw

years, probably by species belonging to the genus

Tityus, it seemed uscful to présent a check list

and keys to the species known to be présent in

Colombia.

SYSTEMATICS

Genus Tityus C. L. Koch, 1836

Tityus oteroi n.sp.

(Figs 1-3)

Holotype. — Colombia. Darién, 1899, F. Geay

leg.: S , deposited in the MNHN, Paris (RS-0853).

Etymology, — Patronym in honour of Dr. Rafael

Otero Patino of the Universidad de Antioquia,

Colombia.

Table 1. — Measurements (in mm) of Tityus oteroi n.sp.

Holotype S

Allotype 2

Carapace;

length

8.7

9.2

anterior width

6.8

7.2

posterlor width

9.4

10.6

Metasoma, segment 1:

length

6.4

5.8

width

4.3

5.0

Metasoma, segment V:

length

10.5

9.6

width

4.5

5.0

depth

4.4

4.3

Vesicle:

width

3.6

depth

3.4

3.3

Fémur:

length

16.2

10.2

width

2.7

2.6

Tibia:

length

16.3

10.8

width

3.2

3.7

Chelae:

length

25.7

19.2

width

2.5

3.5

depth

2.5

3.6

Movable finger:

length

15.2

12.6

MEASUREMENTS. — See Table 1.

Description of male holotype

Coloration

Base colour blackish-brown. Prosoma: carapace

blackish-brown; coxostctnal région ydlowish-

brown. Mesosoma: dorsum blackish-brown with

two confluent yellowish latéral stripes. Venter

yellowish-brown, Metasoma: segments I to IV

blackish-brown; V darker, more blackish. V'esicle;

reddish-black. Chelicerae yellowish-brown with a

very darlc thread; ftngers datk. Pedipalps: palm

reddish-yellow; ftngers reddish-brown. Legs red-

dish-brown with diffuse fuscous spots.

Morphology

Carapace moderately granular; anterior margin

488

ZOOSYSTEMA • 1998 • 20(3)

A new Tityus from Colombia

with a médian concavity. Anterior médian,

superciliary, and posterior médian keels moderate

to srrong. Ail turrows moderately deep. Médian

ocular tubercle distinctly anterior to centre of

carapace. Eyes separated by more than onc ocular

diameter. Three pairs of latéral eyes. Sternum

subtriangular. Mesosoma: tergires moderately

granular. Médian keel strong on ail tergites: tergi-

te VII pentacarinate. Venter: génital operculum

divïded longitudinally. Pectines: pectinal tooth

count 20-21; basal middle lamellae of thc pec¬

tines not dilated. Sternites smooth with elongate

srigmata; sternite Vil with four very feeble keels.

Metasoma: segments I to IV with dorsolateral

keels serrate and latéral supramedian keels crcnu-

late. Dorsolateral keels on segments II to IV with

numerous strongly spinoid granules (this charac-

ter is diagnostic for thc new species) (Fig. IA).

Latéral infratnedian keels on segment 1 complété,

strongly crenulate; on segments 11 and 111 repre-

sented by only two to three distal granules;

absent on segment IV. Ventrolateral and ventral

submedian keels strong, crenulate. Intercarinal

spaces moderately granular. Segment V with

dorsolateral and lateromedian keels vestigial; ven¬

trolateral and ventromedian keels strong, crenu¬

late. Latéral intercarinal spaces moderately

granular. Telson, feebly granular, almost smooth

with a long and strongly curved aculeus. Dorsal

surface smooth; ventral surface feebly granular;

subaculear tooth moderate and spinoid.

Chcliceral dentition charactcnstic of the farnily

Buthidae (Vachon 1963); ventral aspect of both

fingers and manus with long dense setae.

Pedipalps: fémur pentacarinate; tibia with six to

seven keels; chelae with vestigial keels; ail laces

leebly granular, almost smooth. Movable fingers

with 16-16 oblique rows of granules.

Trichobothriotaxy: orthoborhriotaxy A-a

(Vachon 1973, 1975; Figs 2, 3). Legs: tarsus ven-

trally with numerous short fine setae.

Description of allotype and paratypes

One allotype (female) and three paratypes

Fig. 1. — A, Tityus oteroi (6 holotype); metasoma, latéral aspect, showing the carinal structure and the spinoid dorsal granules;

B, Tityus oteroi (9 allotype); pecten. Scale bars : A, 4 mm; B, 2 mm.

ZOOSYSTEMA • 1998 • 20(3)

489

Lourenço W. R.

(females) with same data as for the holotype.

Coloration similar to that of the holotype, only

slightly darker, General morphology different to

that of the holotype hy mophometric values (see

Table 1). Female pectines smaller, with 19-19

(allotype), 18-19, 19-20, 20-20 teeth; basal

middle lamellae strongly dilated (Fig. IB).

Taxonomic POSITION

The new species belongs to the “ Tityus asthenef

group. It is closely associated with Tityus nemato-

chirus Mello-Leitâo, 1940; however, it can readly

be distinguished from this last species by the pré¬

sence of the strongly spinoid granules on the

dorsal keels of metasomal segments II to IV. In

T. nematochirus these keels are poorly carinated

and the granules are round and smooth. It can be

distinguished trom 77 asthmes Pocock, 1893 by

having distinctly more elongated pedipalps (see

taxonomic key).

CHECK LIST

Family BUTHIDAE Simon, 1880

Genus Tityus Koch, 1836

Species belonging to the “ Tityus clathratus ”

group:

Tityus bastasi Lourenço, 1984

77 betschi Lourenço, 1992

77 colurnbianus (Thoreil, 1876)

77 tayropta l.ourenço, 1991

Species belonging to the “ Tityus bahiensis ” group:

Tityus blanci Lourenço, 1994

77 charalaensis Mello-Leitâo, 1940

77 engelkei Pocock, 1902

77 lourençui Flore/., 1995

77 rebierei Lourenço, 1997

77 sabinede Lourenço, 1994

77 sastrei Lourenço et Florez, 1990

Species belonging to the “ Tityus asthenef group:

Tityus asthenes Pocock, 1893

77 cuellan Lourenço, 1994

77 ftttae Borelli, 1899

T forcipula (Gervais, 1844)

77 fuhrmanni Kraepelin, 1914

77 macroehtrus Pocock, 1897

77 nematochirus Mello-Leitâo, 1940

77 oteroi n.sp.

77 pachyurus Pocock, 1897

Total: 20 species (see also map Fig. 4).

D1AGNOSIS OF THE GENUS Tityus Koch,

1836 WITH KEYS TO THE SPECIES

KNOWN FROM COLOMBIA

Scorpions of small, medium or large size ranging

from 25 to 110 mm in total length. The general

coloration can présent almost ail the colour pat¬

terns that are observed among scorpions in gene¬

ral, ranging from pale-yellowîsh or reddish-

yellow to reddish-brown, dark-brown and black-

ish. and with dark spots which may be dtstribu-

ted in many different configurations. Ail

segments are in general strongly granulated.

Dentate margins of pedipalp-chela fmgers com-

posed of twelve to seventeen oblique rows of gra¬

nules, but with supernumerary granules not

présent. Pectine teeth ranging in number from

ten to thirty. Metasomal segments paralle) in

both males and females or elongated or enlarged

posteriorly in males. Sexual dimorphism présent

in almost ail species, and with several different

patterns or configurations.

KEY TO THE THREE CROUPS OF Tityus PROPOSED IN THE CHECK LIST

1. Small species ranging from 25 to 40 mm in total length with variegated pigmenta¬

tion and a rhomboidal subaculear tooth .. Tityus clathratus group

— Species of medium or large size, ranging from 50 to 100 mm in total length; pig¬

mentation varying from yellowish to brown and black; subaculear tooth spinoid .. 2

492

ZOOSYSTEMA • 1998 • 20 (3)

A new Tityus from Colombia

^ Tityus asthenes Pocock, 1893

T. bastosi Lourenço. 1984

T. betschi Lourenço, 1992

] T. blanci Lourenço, 1994

g T. charalaensis Mella-Leitâo, 1940

0 T. cotumbianus (Thorell, 1876)

fa T. cuetlari Lourenço, 1994

<£> T. engelkei Pocock, 1902

A T. festae Borelli, 1899

(•) T. forciputa (Gervais 1844)

^ T. fuhrmanni Kraepelin. 1914

@ T. lourençoi Florez 1995

fôl T. macrochlrus Pocock. 1897

(g T, nematochirus Mello-Leitâp, 1940

Q T. oteroi n.sp.

y T. pachyurus Pocock, 1897

Q T. rebierei Lourenço, 1997

fj) T. sabineae Lourenço. 1994

| T. sastrei Lourenço ef Florez, 1990

A T. tayrona Lourenço, 1991

4 * '

' L "-v f

7 /

Fig. 4. — Known distribution of the Tityus species in Colombia.

ZOOSYSTEMA • 1998 • 20(3)

Lourenço W. R.

2. Species of medium size, ranging from 50 to 70 mm in total Iength; coloration

rather pale varying from yellowish to reddish-brown or brownish, never black; often

with conspicuous dark spots; basal rniddle lamellae of female pectines not dilated in

almost ail species. Tityus bahiensis group

— Species of large size, ranging from 65 to 100 mm in total Iength; pigmentation blac-

kish in the adult and yellowish/variegated in immature individuals; subaculear tooth

always spinoid; basal rniddle lamellae of female pectines dilated in almost ail species

..... Tityus asthenes group

Key TO THE SPECIES OF THE Tityus clathratus GROUP

1. Dorsolateral keels of metasomal segments I to IV without a spinoid posterior gra¬

nule or with only a very feeble granule .2

— Dorsolateral keels of metasomal segments I to IV with a strong spinoid posterior

granule; Amazonia ... T. bastosi

2. Subaculear tooth strong and very rhomboidal .3

— Subaculear tooth moderate and feebly rhomboidal . T. betschi

3. Pectines with ten to fourteen teeth ... T. columbianus

— Pectines with fourteen to seventeen teeth . T. tayrona

Key TO THE SPECIES OF THE Tityus bahiensis GROUP

1. Ventral submedian keels of metasomal segments III and IV converging proximally

to form a Y shape . T, rebierei

— Ventral submedian keels of metasomal segments III and IV parallel through entire

Iength .2

2. General coloration from yellowish to reddish-yellow .3

— General coloration from dark-reddish to blackish.5

3. Dorsolateral keels of metasomal segments II to IV with several granules modified as

spines ... T. blanci

— Dorsolateral keels of metasomal segments II to IV without granules modified as

spines .4

4. Basal rniddle lamellae of female pectines strongly dilated; dentate margins of pedi-

palp-chela fingers composed of thirteen to fourteen oblique rows of denticles .

. T. engelkei

494

ZOOSYSTEMA • 1998 • 20(3)

A new Tityus from Colombia

— Basal middle lamellae of female pecdnes not dilated; dentate margins of pedipalp-

chela fingers composed of sixteen oblique rows of dendcles . T. sastrei

5. Male pedipalps slighdy longer and more slender than those of females.6

— Male pedipalps larger and more bulky than those of females . T. sabineae

6. Pectines with sixteen to seventeen teeth; dentate margins of pedipalp-chela fingers

composed ofseventeen oblique rows of denticles . T. lourençoi

— Pectines with fourteen teeth; dentate margins of pedipalp-chela fingers composed of

twelve oblique rows of denticles . T. charalaensis

Key TO THE SPECIES OF THE Tityus asthenes GROUP

1. Male pedipalps longer and more slender than those of females .2

— Male pedipalps larger and more bulky than those of females .4

2. Values of (male) fémur, tibia and chelae length respectively 6/8, 8/10 and

10/13 dmes longer than width .....3

— Values of (male) fémur, tibia and chelae length respectively 5, 3 and 5 times longer

than width ...... T. asthenes

3. Dorsolateral keels of metasomal segments I to IV with several strong spinoid poste-

rior granules ..... T. oteroi

— Dorsolateral keels of metasomal segments I to IV without spinoid posterior granules

....... T. nematochirus

4. Dorsolateral keels of metasomal segments I to IV with several strong spinoid poste¬

rior granules, or with one very strong spinoid posterior granule .5

— Dorsolateral keels of metasomal segments I to IV with moderate or feeble spinoid

posterior granules.......7

5. One very strong spinoid posterior granule on dorsolateral keels of metasomal seg¬

ments II to IV .. T. fuhrmanni

— Several strong spinoid posterior granules on dorsolateral keels of metasomal seg¬

ments I to IV ...........6

6. Adult size around 60 mm; legs and pedipalps blackish .. T, forcipula

— Adult size around 45 mm; legs and pedipalps yellowish to reddish-yellow.

. T. cuellari

ZOOSYSTEMA • 1998 • 20(3)

495

Lourenço W. R.

7. Moderate spinoid posterior granules on dorsolateral keels ot metasomal segments I

to IV; male pedipalps larger and more bulky than those of temales . T. pachyurus

— Feeble spinoid posterior granules on dorsolateral keels of metasomal segments I to

IV; male pedipalps only slightly larger and more bulky than those of females .8

8. Pectines with fifteen to sixteen teerh; dentate margins of pedipalp-chela fingers

composed of fifteen oblique rows of denticles . T. macrochtrus

— Pectines with twenty-one to twenty-two teeth; dentate margins of pedipalp-chela

fingers composed of sixteen oblique rows of denticles . T. festae

Acknowledgements

I am very grateful to Mr, Jacques Rebière,

Laboratoire de Zoologie Arthropodes (MNHN),

for preparing several illustrations, and especially

to Prof. John L. CIoudsley-Thompson of

University College, London, for reviewing the

manuscript.

REFERENCES

Gervais P. 1844. — Remarques sur la famille des

Scorpions et description de plusieurs espèces nou¬

velles de la collection du Muséum. Archives du

Muséum national d'Histaire naturelle 4 : 203-240.

Kraepelin K. 1912. — Ncuc Beitrage zur Systematik

der Gliederspinnen. Mitteilungen ans dem

Naturhistorischen Muséum. Hamburg 29: 45-88.

— 1914. — Bcitrag zur Kcnntnis der Skorpione und

Pedipalpec Columbicns. Mémoires de la Société des

Sciences naturelles, Neuchâtel 5 : 15-28.

Lourenço W. R. 1984a. — Analyse taxonomique des

Scorpions du groupe Tityus clathratus Koch, 1845

(Scorpiones, Buthidae). Bulletin du Muséum natio¬

nal a Histoire naturelle , Paris, série 4, A 6 (2) :

349-360,

— 1984b. — Etude systématique de quelques espèces

appartenant au complexe Tityus forcipula (Gervais,

1844) (Scorpiones, Buthidae). Bulletin du Muséum

national dHistoire naturelle, Paris, série 4, A 6 (3) :

729-739.

— 1991. — Les Scorpions (Chelicerata) de

Colombie. II. Les faunes des régions de Santa

Marta et de la cordillère orientale. Approche bio¬

géographique. Senckenbergiana biologica 7 (4/6):

275-288.

— 1992. — Biogéographie des espèces du groupe

naturel « Tityus clathratus » (Chelicerata,

Scorpiones, Buthidae). Bulletin du Muséum natio¬

nal d’Histoire naturelle , Paris, série 4, A 14 (2) :

473-481.

— 1993. — Opistbacanthus lepturus (Palisot de

Beau vois), Scorpion à modèle complexe de distri¬

bution, Btogeographica 6 9 (2); 87-88.

— 1994. — Scorpions Chelicerata de Colombie.

VI. Quatre nuuvelles espèces de Buthidae des

régions amazonienne. Sud-pacifique et de la cor¬

dillère orientale. Revista de la Academia Colornbiana

de Çiencuis 19 (73): 387-392.

— 1995a. — Nouvelles considérations sut la classifi¬

cation et la biogéographie des Opistbacanthus néo¬

tropicaux (Scorpiones, lschnuridae). Biogeographica

71 (2): 75-82.

— 1995b. — Considération s sur la biogéographie des

espèces appartenant au. genre Teutbraustes Simon,

1878 (Chelicerata, Scorpiones, Chactidae), Revue

Ârachnologique 10 (II) : 201-206.

— 1992. — Synopsis de la faune de scorpions de

Colombie, avec des considérations sur la systéma¬

tique et la biogéographie des espèces. Revue suisse de

Zoologie 104 (1) : 61-94.

Lourenço W. R. & Flore?. F.. 1989. — Los escor-

piones (Chelicerata) de Colombia. I. La fauna de la

isla de Gorgona. Approximation hiogeografica.

Caldasia 16 (76): 66-70.

— 1990a. — Scorpion (Chelicerata) front Colombia.

III. The scorp'io-fauna of pacifie région (Choco),

with somc biogeograpbic considérations.

Amazoniana 11 (2): 119-133.

— 1990b, — Scorpions (Chelicerata) de Colombie.

IV. Biogéogiaphie et diversité biologique des

Scorpions de Colombie, avec des commentaires sur

les refuges quaternaires. Comptes Rendus des Séances

de la Société de Biogéographie 66 (2) : 65-74.

— 1995. — Caractérisation géographique d’une nou¬

velle espèce de Scorpion appartenant au genre

Tarsopvrosus Francke (Diplocentridae). Biogeogra-

phicaV 1 (3): )43, 144.

Mello-Leitâo C. 1945. — Escorpiôes Sul Americanos.

Arquivos do Museu Nacional, Rio de Janeiro 40:

1-468.

496

ZOOSYSTEMA • 1998 • 20(3)

A new Tityus from Colombia

Pocock R. I. 1893. — A contribution to the study of

Neotropical Scorpions. Annals Magazine Natural

History, sériés 6, 12: 77-103.

Thorell T. 1876. — On the classification of scor¬

pions. Annals Magazine Natural History. 1-15.

Vachon M. 1963. — De l’utilité, en systématique,

d’une nomenclature des dents des chélicères chez

les Scorpions. Bulletin du Muséum national d’His-

toire naturelle, Paris, série 2, 35 (2) : 161-166.

-— 1973. — Étude des caractères utilisés pour classer

les familles et les genres de Scorpions (Arachnides).

1. La trichobothriotaxie en arachnologie. Sigles tri-

chobothriaux et types de trichobothriotaxie chez les

Scorpions. Bulletin du Muséum national d'Histoire

naturelle, Paris, série 3, n° 140, Zoologie 104 :

857-958.

— 1975. — Sur l'utilisation de la trichobothriotaxie

du bras des pédipalpes des Scorpions (Arachnides)

dans le classement des genres de la famille des

Buthidae Simon. Comptes Rendus sommaires de

l’Académie des Sciences, Paris, série D 281:

1597-1599.

Submitted on 8 December 1997;

accepted on 3 April 1998.

ZOOSYSTEMA • 1998 • 20(3)

497

Réhabilitation du genre Harmonicon (Pickard-

Cambridge, 1896) et description d’une nouvelle

espèce de Guyane française (Araneae,

Mygalomorphae, Dipluridae)

Patrick MARÉCHAL

Laboratoire de Zoologie (Arthropodes), Muséum national d’Histoire naturelle,

61 rue de Buffon, F-75231 Paris cedex 05 (France)

Christian MARTY

Villa Goarand, 9 rue du Dr Moges, F-97354 Montjoly, Guyane (France)

MOTS CLÉS

Araneae,

Mygalomorphae,

Dipluridae,

Guyane française,

nouvelle espèce,

Harmonicon.

Maréchal P. & Marty C. 1998. — Réhabilitation du genre Harmonicon (Pickard-Cambridge,

1896) et description d'une nouvelle espèce de Guyane française (Araneae,

Mygalomorphae, Dipluridae). Zoosystema 20 (3) : 499-504.

RÉSUMÉ

Une nouvelle espèce de la famille des Dipluridae est décrite de Guyane fran¬

çaise. Ses affinités avec une espèce brésilienne conduisent à la réhabilitation

du genre Harmonicon (Pickard-Cambridge, 1896). Quelques éléments de la

biologie de cette nouvelle espèce sont exposés.

KEYWORDS

Araneae,

Mygalomorphae,

Dipluridae,

French Guiana,

new species,

Harmonicon.

ABSTRACT

Reinstatement of the genus Harmonicon (Pickard-Cambridge, 1896) with des¬

cription of a new species from French Guiana (Araneae , Mygalomorphae,

Dipluridae). A new species of the Dipluridae family is described from French

Guiana. lts affinities with a Brasilian species lead to the reinstatement of the

genus Harmonicon (Pickard-Cambridge, 1896). Some biological aspects of

this new species are described.

ZOOSYSTEMA • 1998 • 20(3)

499

Maréchal P. & Marty C.

INTRODUCTION

Dans son dernier travail consacré aux araignées

de la Guyane française, Caporiacco (1954) recense

trois espèces de la famille des Dipluridae.

Poursuivant l’étude systématique de la faune

mygalomorphe de ce département d’outte-mer

récemment entreprise (Maréchal 1996), nous

décrivons ici une nouvelle espèce de cette famille.

Cette donnée inédite nous conduit à revenir sur

certaines synonymies établies par Raven (1985)

au sein de ta sous-famiJIe des Diplurinae et plus

particulièrement dans te genre Diplura.

Les informations écologiques et comportemen¬

tales collectées à plusieurs reprises sur le terrain

permettent également de présenter quelques

aspects de la biologie de cette Dipluridae guya-

naise.

Les Dipluridae américaines

La famille des Dipluridae comprend actuelle¬

ment quatre sous-familles (Raven 1985). Parmi

celles-ci, celle des Diplurinae est spécifique du

continent américain et, plus précisément, de la

région néotropicale, Il parait donc tout à lait

naturel de trouver de nouvelles espèces de cette

sous-famille en Guyane française, département

dont l’étude de la faune arachnologique est à

peine entamée.

Le genre Diplura

Le genre Diplura a été créé par Simon (1892) sur

la base de scopulae peu denses sur les tarses.

Après la découverte par Pickard-Cambridge

(1896) d'une lyre sur la coxa des pédipalpes chez

Melodeus, Simon (1903) la suppose absente chez

Diplura, Par la suite, tous les auteurs ont classé

les espèces munies d’une lyre dans d'autres

genres. Ce n’est que lors du travail de révision de

Raven (1985) que la présence d’une lyre est

constatée chez Diplura macrura (Koch, 1842) et

que toutes les Diplurinae munies d’un tel organe,

exceptées celles du genre Trechona (Koch, 1851),

y sont rassemblées. Pas moins de onze genres,

souvent monospécifiques, sont ainsi mis en syno¬

nymie par Raven (1985).

La découverte d'une mygale proche de Diplura

rufescens (Pickard-Cambridge, 1896) et l’examen

du type de cette dernière révèlent un certain

nombre de différences morphologiques d’impor¬

tance générique au niveau de la lyre et des pattes.

Nous proposons donc la réhabilitation du genre

Harmonicon (Pickard-Cambridge, 1896)

jusqu’ici monospécifique et en décrivons une

nouvelle espèce.

Genre Harmonicon Pickard-Cambridge, 1896

Espèce-type. — L’espèce-type du genre est

Harmonicon rufescens Pickard-Cambridge, 1896.

Diagnose différentielle. — Diffère de toutes les

autres Diplurinae par la formule ambulatoire 1423.

Diffère de Diplura par des pattes proportionnellement

plus longues et plus fines, la lyre dont l’extrémité des

soies est aplatie et recourbée en crochet, et l’absence

de protubérance sur les métatarses I des mâles adultes.

Description

Dipluridae de grande taille (corps supérieur à

25 mm), gracile, aux pattes longues et fines dont

le rapport ambulatoire est très nettement infé¬

rieur à dix (supérieur chez les espèces du genre

Diplura). La lyre est constituée de cinq soies, ce

qui pourrait être un caractère générique. Les

filières postérieures latérales sont longues et effi¬

lées.

Remarque

Le spécimen-type de H. rufescens est mutilé d’ori¬

gine et la quatrième paire de pattes manque. La

formule ambulatoire ne peut donc être connue.

De plus, Raven (1985) considère la formule 4123

comme l un des caractères distinctifs des

Diplurinae. Les deux espèces du genre Harrno-

nicon possédant par ailleurs tous les caractères de

la sous-famille, ce critère ne peur être considéré

comme pertinent pour caractériser les Diplu¬

rinae.

Harmonicon audeae n.sp.

MATÉRIEL EXAMINÉ. — Guyane française. Stn de

Saint-Eugène, 4 n 50’N - 53 o 05 W, capturée le

22.X. 1996, cnil, de Massary : holotypc ?. — Piste de

Saint-Élit, pk 10. 5°15’N 53“*05’W, capturé le

21.X. 1995, coll. P. Gaucher : allotype â. — Stn de

Saint-Eugène, capturé le 22.X.1996 : 1 cl ; capturé le

18.V. 1996, coll, de Massary) : I immature.

L’ensemble du matériel est déposé au Muséum natio¬

nal d Histoire naturelle, Paris (MNHN).

500

ZOOSYSTEMA • 1998 • 20(3)

Le genre Harmonicon (Aranaea, Dipluridae)

ÉTYMOLOGIE. — Cette espèce est baptisée en l’hon¬

neur de la naissance de la fille de l’un des auteurs.

Diagnose différentielle. — Diffère de H. rufescens

par la présence sur les tarses des pédipalpes d’une sco-

pula et d'une griffe munie d’une double rangée de

dents (femelle). Diffère également par la présence

d’épines sur les fémur des pattes I et II.

Description

Holotype femelle

Longueur totale sans les chélicères, 33 mm.

Couleur. Céphalothorax brun orangé. Pattes,

sternum et pièce labiale fauve clair, légèrement

jaunâtre ou orangé. Abdomen uniformément gris

violacé recouvert d une fine pubescence grisâtre

et parsemé de long poils noirs.

Carapace. Longueur 13,5 mm ; largeur 12 mm.

Recouverte d une pubescence noire clairsemée et

bordée de soies noires courbées vers l’avant.

Fovéa. Petite, profonde et légèrement récurvée.

Yeux, Tubercule distinct mais peu élevé, yeux

antérieurs en ligne légèrement procurvée, les pos¬

térieurs en ligne faiblement récurvée. Les yeux

médians antérieurs sont les plus gros (Fig. IA).

Chélicères. Munies d’une plage dorsale longitu¬

dinale de longs poils qui s’évase vers la base du

crochet. Présence d’une bande pubescente en

position dorso-latérale et d’une autre plus étroite

Fig. 1. — Harmonicon audeae n.sp., holotype S. A, aire ocu¬

laire, vue dorsale ; B, cbéllcère droite, vue ventrale ; C, lames

maxillaires, labium et sternum, vue ventrale. Échelles : A,

1 mm ; B, C, 3 mm.

er plus discrète en position latéro-exteme. Marge

interne portant neuf dents coniques en ligne ter¬

minée, dans la partie basale, par un groupe de

Fig. 2. — Harmonicon audeae n.sp., <3, MEB ; A, face interne de la coxa du pédipalpe droit montrant l'emplacement de la lyre ;

B, détail de la lyre constituée de cinq soies dont l’extrémité, aplatie, est recourbée en crochet. Échelles : A, 1 mm ; B, 100 pm.

ZOOSYSTEMA • 1998 • 20(3)

501

Maréchal P. & Marty C.

cinq dents plus petites. Gouttière munie d’une

série longitudinale de denticules qui se termine

en plage dans la partie basale. De longs poils

orangés et denses garnissent la marge externe qui

porte quatre soies isolées dans sa partie basale

(Fig. IB).

Lames maxillaires. Longueur 4,5 mm ; largeur

3,15 mm. Munies, dans leur angle interne, de

quarante à quarante-cinq denticules. Serrula pré¬

sente dans la partie interne près de la marge api¬

cale. Présence, sur la face interne, d’une lyre

constituée de cinq soies dont l’extrémité, aplatie,

est recourbée en crochet (Fig. IC, 2).

Labium. Longueur 2,25 mm ; largeur 2,25 mm.

De forme quadrangulaire, le labium ne porte que

quelques poils épars. Pas de denticules (Fig. IC).

Sternum. Longueur 6,6 mm : largeur 5,7 mm.

Suture Jabio-sternale constiruée par deux larges

fossettes semi-circulaires épousant le contour de

la base du labium. Sigillés bien distincts, ronds et

profonds. Les postérieurs sont les plus gros.

Pédipalpe. Muni d’une griffe portant, à sa base,

une double rangée de six à sept dents. Présence

d’une scopula peu dense mais nettement visible

sur la moitié apicale du tarse. Présence de courtes

trichobotbries en une rangée médiane sur le tarse

et en une double rangée latérale sur le tibia.

Pattes. Formule ambulatoire (classement des

pattes par ordre de longueur décroissant) 1423.

Pattes longues et Fines, le rapport ambulatoire

(diamètre du fémur/longueur totale X 100) com¬

pris encre 6.4 et 4,5- Griffés paires munies d’une

double rangée de six à huit dents. Griffe impaire

mutique. Tarses pseudosegmentés. Scopula peu

dense séparée par une double rangée médiane de

setae sur les tarses I et II. Scopula quasi inexis¬

tante et setae plus fortes sur les tarse III et IV.

Mensurations (mm)

III

Fémur

12,5

11,5

8,5

Patelle

5,5

3,5

Tibia

11,5

10,5

9,5

Métatarse

20,5

Tarse

6.5

6,5

Total

49,5

45,5

Diamètre fémur

3,1

2,9

2,8

2,7

rapport ambulatoire

6,26

6,37

6,22

4,5

—

Fig. 3. — Harmonicon audeae n.sp„ allotype 6 . A, aire oculaire,

vue dorsale ; B, chélicère droite, vue ventrale ; C et D, éperon

du tibia I en vue latéro-externe (C) et ventrale (D) ; E, abdomen

et filières en vue latérale. Échelles A, 1 mm ; B-E, 3 mm.

Présence d’une scopula clairsemée sur le métatar¬

se I et dans le tiers apical du métatarse II.

Trichobothries longues et fines, présentes en une

rangée médiane sur les tarses et les métatarses et

en une double rangée latérale sur les tibias.

Épines nombreuses et très irrégulières par leur

baille et leur force, présentes sur les métatarses, les

tibias et les fémurs. Leur nombre et leur disposi¬

tion est très variable d’un individu à l’autre et d’un

côté à l’autre sur un même individu. Tarses tou¬

jours mutiques a l’exception des pédipalpes qui

portent une ou deux épines sur la lace ventrale.

Filières. Longueur des postérieures médianes

3.5 mm. Longueur des postérieures latérales

22.5 mm ; article basal 6,5 mm ; article médian

6,25 mm ; article apical 9,75 mm. Filières posté-

502

ZOOSYSTEMA ■ 1998 • 20(3)

Le genre Harmonicon (Aranaea, Dipluridae)

Fig. 4. — Harmonicon audeae n.sp., allotype â, pédipalpe droit. A, profil externe ; B, vue ventrale ; C, profil interne. Échelle : 3 mm.

rieures latérales longues et fines avec un article

apical effilé.

Allotype mâle

Longueur totale sans les chélicères, 28 mm.

D’aspect plus grêle, le mâle diffère de la femelle

par les points suivants :

Carapace. Longueur 13 mm ; largeur 10.5 mm.

Yeux. Yeux médians antérieurs proportionnelle¬

ment plus gros que chez la femelle (Fig. 3A).

Chélicères. Marge interne avec onze dents

coniques en ligne. Gouttière munie d’une simple

série longitudinale de denticules ; plage de la par¬

tie basale présente, mais peu visible car non chiti-

nisée comme chez la femelle (Fig. 3B).

Lames maxillaires. Serrula absente. Lyre iden¬

tique.

Pédipalpe. Présence d'une épine prolatérale dans

le tiers apical du tibia. Embolus piriforme termi¬

né en courte pointe effilée (Fig. 4).

Pattes. Proportionnellement plus longues et fines

avec des rapports ambulatoires qui varient de 3 à

3,5. Scopulas plus denses que chez la femelle.

Présence, dans la partie apicale des tibias I, d’un

éperon constitué d’un processus surmonté d’une

forte épine (Fig. 3C, D).

Mensurations en mm

•

III

Fémur

15,5

13,75

12,75 15

7,5

Patelle

3,25

Tibia

10,5

13,5

6,75

Métatarse

13,5

14,25 17

Tarse

9,25

8,75

7,25

8,5

2,5

Longueur totale

68,25

60,75

Diamètre fémur

2,15

2,1

1,85

Rapport ambulatoire

3,15

3,46

3,3

3,03

Filières. Longueur des postérieures médianes

2,75 mm. Longueur des postérieures latérales

21.5 mm ; article basal 5,5 mm ; article médian

6.5 mm ; article apical 9,5 mm. Pas de différence

significative avec les filières de l’holotype femelle

(Fig. 3E).

ZOOSYSTEMA • 1998 • 20(3)

503

Maréchal P. & Marty C.

Habitat

Harmonicon audeae, à l’instar de H. rufescens, est

une Dipluridae forestière. Elle affectionne parti¬

culièrement la base des arbres, les souches ou les

cavités naturelles des talus où elle tisse une toile

en nappe horizontale munie d’un tube retraite

qui s’enfonce profondément dans les anfractuosi¬

tés du substrat. La partie plane de la construction

soyeuse, soutenue par des fils verticaux, est large¬

ment étendue à l'extérieur et peut atteindre une

superficie de près d'un mètre carré.

Obscuricole, H, audeae n’a jamais été vue de

jour. Dès la tombée de la nuit, elle se poste à

l’affût dans sa toile, tout près de l’entrée de son

tube. D’une grande agilité, cette araignée est

prompte à regagner sa retraite à la moindre alerte.

Déjà noté par Pickard Cambridge (1896) pour

H. rufescem, ce comportement rend sa capture

difficile. Les mâles, errants, sont plus aisés à cap¬

turer, notamment à l’aide de pièges dits de

Barber ou « pit fait traps ».

Selon les témoignages d’Amérindiens locaux,

plus particulièrement des Oyampis, cette mygale

serait très venimeuse. Toutefois, aucun cas

d’envenimation n’est actuellement connu des ser¬

vices médicaux guyanais.

Remerciements

Tous mes remerciements vont au Dr P. Hillyard

du Natural History Muséum de Londres qui a

bien voulu me confier le matériel-rype nécessaire

à l’élaboration de ce travail. Mes remerciements

également à M. J. Rebière pour la réalisation des

dessins de la figure 4.

RÉFÉRENCES

Caporiacco L. di 1954. — Araignées de la Guyane

française du Muséum national d'Histoire naturelle

de Paris. Commentationes pontificia Academia scien-

tiarum 16: 45-193.

Koch C. L. 1842. — Die Arachniden. Neunter Band,

Nürnberg: 1-108.

Maréchal P. 1996. — Psalistops gasci n.sp., première

Barychclidac de Guyane française (Araneae,

Mygalomorphae). Bulletin du Muséum national

d'Histoire naturelle, Paris, série 4, 18 (3-4) : 589-594.

Pickard-Cambridge F. O. 1896. — On the

Theraphosidae of the Lovver Amazons: being an

account of the new généra and species of tliis group

of spiders discovered during the expédition of the

steamshîp “Faraday” up the river Amazons.

Proceedings of the Zoological society of London:

716-766.

Raven R. J. 1985. — The spider infraorder

Mygalomorphae (Araneae): Ciadistics and systema-

tics. Bulletin of the American Muséum of natural

History 182, 180 p.

Simon E, 1892. — Histoire naturelle des araignées.

Tome 1, fascicule 1. Paris, 256 p.

— 1903. — Histoire naturelle des araignées. Tome 2,

fascicule 4. Paris : 669-1080.

Soumis le 4 décembre 1997;

accepté le 3 avril 1998.

504

ZOOSYSTEMA • 1998 • 20(3)

A new pedunculate cirripede (Thoracica,

Heteralepas ) from the Northeast Atlantic Océan

René-Pierre CARRIOL

Laboratoire de Paléontologie, Muséum national d’Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

rene.pierre.carriol@wanadoo.fr

KEYWORDS

Crustacea,

Cirripedia,

Thoracica,

taxonomy,

Meteor Seamount,

Northeast Atlantic.

Carriol R.-P 1998. — A new pedunculate cirripede (Thoracica, Heteralepas) from the

Northeast Atlantic Océan. Zoosystema 20 (3): 505-509.

ABSTRACT

A new species of Heteralepas is described from waters of 300 m at Meteor

Seamount, south of the Azores, in the Northeast Atlantic Océan. This spe¬

cies possesses a peduncle longer than the capitulum, a carinal crest without

protubérances and continuing onto the peduncle, and it lacks scuta.

MOTS CLÉS

Crustacea,

Cirripedia,

Thoracica,

taxonomie,

banc Météor,

Atlantique (Nord-Est).

RÉSUMÉ

Un nouveau cirripede pédoncule (Thoracica, Heteralepas,) du nord-est de

l’océan Atlantique. Description d’une nouvelle espèce à’Heteralepas trouvée

par 300 m de fond, sur le banc Météor, au sud des Açores, dans l’est de

l’Atlantique nord. Cette espèce possède un pédoncule plus long que le capi¬

tulum et une crête carinale, sans protubérances, se prolongeant le long du

pédoncule, mais n’a pas de scuta.

ZOOSYSTEMA • 1998 • 20(3)

505

Carriol R.-P.

INTRODUCTION

The Meteor Seamount was sampled by the

Lesivy during a rrawling survey in August 1990.

The expédition was sponsored by the FROM-

Bretagne. M. H. Du Buit identified the Fish

fauna (Du Buit 1991) and provided the spéci¬

mens of Heteralepas for study. This new taxon is

the sixth species of Heteralepas from the Adantic

Océan.

SYSTEMATICS

Subclass CIRRIPEDIA Burmeister, 1834

Superorder THORACICA Darwin, 1854

Order PEDUNCUt.ATA Lamarck, 1818

Suborder HETERALEPADOMORPHA

Newman, 1987

Family HFTKRAUÎPADIDAK Nilsson-Cantell, 1921

Genus Heteralepas Pilsbry, 1907

Species included. — The 21 taxa presendy attribu-

ted to this genus inelude : H. corauta (Darwin, 1851);

//. japomca (Aurivillius, 1892); H, lankesteri (Gruvel,

1900); H. belli (Gruvel, 1902); H. microstoma

(Gruvel, 1902); H. gigiis (Annandale, 1905); H. ovalis

(Hoelc, 1907); H. tennis (Hoek, 1907); H. rex

(Pilsbry, 1907a); H. èygnus Pilsbry, 1907b; H. nicobd-

rica Annandale, 1909; H. verula Pilsbry, 1909;

H. dubia Broch, 1922; H. batstii Hiro, 1937; H. Uti-

nomii Newman, 1960; H. mystacophora Newman,

1964; H. lundi/s Zevina, 1975; H. adiposa Zevina,

1982; H. fulva Zevina, 1982; H. smilius Ren, 1983;

plus the new species described here: Hetcralepiis meteo-

rensis n.sp.

DlAGNOSlS. — “Inner rami of cirri V and VI atro-

phied; articles of cirri (cxccpr cirrus I and inner rami

of V and VI) rectangular; cirri long: lesser curvature of

cirri supporting large pair of setae and one or two

smaller pairs from tne same origin; greater curvature

supporting a few unpaired setae." (Newman 1960).

Heteralepas meteorensis n.sp.

(Fig. F)

Type MATERIAU. — The types and additional spéci¬

mens are deposited in two places, the Laboratoire de

Zoologie-Arthropode:,, Muséum national d’Histoire

naturelle, Paris [nolotype MNHN-Ci2689; paratypes

MNHN-Ci2690 (specimen opened) and MNHN-

Ci2691 (dissected, appendages preserved in alcohol);

additional specimens MNHN-Q2692 (1 specimen)

and MNHN-Ct‘2693 (a small colony of about 15

adults) and the Laboratoire de Biologie marine du

Collège de France, Concarneau [additional tnatcrial (a

colony of scveral tens of adults)).

EtymoI-OGY. — Narned for the Meteor Seamount,

the type locality.

TYPE LOCALITY — Meteor Seamount, south of Azores

Islands (30°N - 28°30’W), at 300 ni depth, living on

Callogorgia sp. (Alcyonaria).

DlAGNOSlS

Peduncle obviously longer than capitulum; capi-

tulum lacking scuta; distinct carinal crest,

without protubérances, continuing onto the

peduncle.

Description

Specimens yellow pink in alcohol. Capitulum

globular, laterally compressed, carinal margin

broadly convex with carinal crest extending from

orifice over the capitulum to the base of pedun-

cule; scutal crest absent; lacking valves. Orifice

between one fourth and one third height of capi¬

tulum; lips slightly crenulate, Peduncle long,

annulated, two to four tintes length of capitulum

(Table 1), cylindrical, with diameter equal to

capitulum width

Tabi e 1. — Measurernents (in cm) of capitulum and peduncle.

Holotype Paratype Paratype

MNHN-Ci2689 MNHN-Ci2690 MNHN-CI2691

capitulum height

1.6

1.8

1.2

depth

1.4

1.7

1.2

width

0.7

0.8

peduncle length

3.4

7.2

3.1

diameter

0.7

1-1.3

0.8

Labrum slightly bullate, anterior portion with

tufts of short fine setae, crest with twenty-five

small, sharp teeth; palps nearly tciangular, superior

margin with numerous long spines.

Mandible with four teeth including inferior

angle, covered with short spinules; lower margin

of first tooth with few spinules and some minute

setae; lower margin of second and third teeth

with several tiny spinules; upper margin of

506

ZOOSYSTEMA • 1998 • 20(3)

A new Heteralepas from Atlantic Océan

second, third and fourth teeth with tinv spinules.

Superior margin of mandible with tuft of long

thin spinules and tuft of shorter spinules, inferior

margin supporting row of numerous short thin

spinules.

First maxilla notched, portion above notch with

a single strong spine; notch supporting a long

spine below and two long spines above with a

short spine and two long thin spines between;

lower cutring edge with long and some shorter

spines; short spines in two groups along superior

margin, tuft of few spinules near base of first

major spine, brush of spinules along inferior

margin; surface clothed with numerous spinules

arranged in rows.

Second maxilla broadly rounded; distal portion

Table 2. — IMumber of articles of right cirri (l-VI) and caudal

appendage (ca) from the paratype MNHN-Ci2691.

lit

VI ca

Inner ramus

26 14

Outer ramus

with tufts and rows of long thin pinnate spi¬

nules; proximal portion with a tuft of very long

pinnate spinules.

Inncr rami of cirri V and VI atrophied (Table 2).

Lesser curvature of cirrus V supports one pair of

long, one pair of short, and one pair of minute

setae, from nearly common base on each seg¬

ment. At base of long pair of setae, a very short

Fio. 1. — Heteralepas meteorensis n.sp.; A, holotype MNHN-Ci2690; B-G, paratype MNHN-CÎ2691; B, labium and right palp;

C, mandible; D, first maxilla; E, second maxilla; F, Intermediate article of outer ramus of cirrus V; G, pénis and caudal appendage;

H. distal part of pénis. Scale bars; A, 4 mm; B-F, H, 0.5 mm; G, 1 mm.

ZOOSYSTEMA • 1998 • 20 (3)

507

Carriol R.-P.

Table 3. — Species of the genus Heteralepas of the world océan.

Heteralepas

Western Atlantic

Eastern Atlantic

Eastern Pacific

Indo-Pacific

H. eornuta

West Indies,

North Caroline

Western Africa

Chile

Adaman Sea, Philippines

H. japonica

Japan, China,

Indo-Malaysian waters,

Philippines, Australia,

New Zealand

H. lankesteri

West Indies, Brazil

H. belli

Cuba

—

H. microstoma

Madera

H. gigas

Malaysia

H. ovalis

Malaysia

H. tenuis

New Guinea

H. rex

Hawaii

H. cygnus

California

H. nicobarica

Nicobar Islands

H. vetula

Japan

H. dubia

New Zealand

H, hataii

Japan

H. utinomii

Tasmania

H. mystacophora

Southeast Pacific

H. luridas

Carribbean

H. adiposa

Japan

H. fulva

Southeast Pacific

H. smilius

China

H. meteorensis n.sp.

—

South of Azores

—

spinule arises. Greater curvature supports graded

sériés of three or tour short .setae at each articula¬

tion.

Caudal appendagc of fourteen articles with thin

setules at distal margins ( fable 2).

Pénis weakly annulated, slender, tapering gra-

dually throughout its length; covered with

numerous long, soft setae.

Discussion

Zevina (1982) divides the Heteralepas in two

groups: those with thepeduncle obviously longer

than the capitulum and those with the peduncle

shorter. Heteralepas meteorensis n.sp. belongs to

the first group. It differs frotn H. gigas by lacking

scuta. The carinal crest without protubérances

distinguishes H. meteorensis n.sp. from H. fulva

and H. nicobarica. H. meteorensis n.sp. cannot be

confused with H. cygnus , in which the carinal

crest does not continue on the peduncle.

Distribution

Heteralepas attains greatest diversity in the Indo-

Pacific (Zevina 1982; Ren 1983, Rosell 1991):

among the twentv-one species of which this

genus consists (Table 3), sixteen are exclusively

from Indian and Pacific Océans and four are

exclusively from Atlantic Océan. Only one spe¬

cies, H, cornura , is known from both the Atlantic

and Pacific Océans. H. meteorensis n.sp. is along

with H. microstoma and H. eornuta (the latter

two from Madera and Western Africa respectiv-

ely), H comuta also occurs in Western Atlantic

Océan (West Indies, North Caroline) as does

H. luridas (Caribbean), H. lankesteri (West

Indies, Brazil) and H. belli (Cuba).

Acknowledgements

1 wish to thank J. S. Buckeridge (UNITEC

Institute of Technology, Auckland, New

Zealand) and an ânonymous reviewer for several

useful comments on the manuscript, D. Defaye

(Laboratoire de Zoologie-Arthropodes, Muséum

national d'Histoire naturelle, Paris, France) who

kindly lent me her microscope, B. Battail

(Laboratoire de Paléontologie, Muséum national

508

ZOOSYSTEMA • 1998 • 20(3)

A new Heteralepas from Atlantic Océan

d’Histoire naturelle, Paris, France) who transla¬

tée! me some russian texts and C. Norrthon who

duplicated my drawings.

REFERENCES

Annandale N. 1905. — Malaysian Barnacles in the

Indian Muséum. Memoirs of the Asiatic Society of

Bengal 1 (5): 73-84.

— 1909. — An Account of the Indian Cirripedia

Pedunculata. Memoirs of the Indian Muséum 2 (1):

61-137.

Aurivillius C. W. S 1892. — Neue Cirripeden aus

dem Atlantischen, Indischen und Stillen Océan.

Ofversigt af Konglica Vetenskaps-Akademiens

FSrhandlingar 49 (3): 123-134.

Broch H. 1922. — Papers from Dr Th. Mortensen’s

Pacific Expédition 1914-1916, 10. Studies on

Pacific Cirripeds. Videnskabelige Meddeleiser fra

Dansk naturhistorisk Forening i Kobenhavn 73:

215-358.

Darwin C. 1851. — A monograph on the subclass

Cirripedia, with figures of ail the species. The

Lepaaidae or Pedunculated Cirripedes, Ray Society,

London, 400 p.

Du Buit M. H. 1991. — Faune ichtyologique des

bancs situés autour des Açores. Mésogèe 51 : 25-28.

Gruvel A. 1900. — On new species of the genus

Alepas {A. lankesteri), from tne collection of the

Bridsh Muséum. Armais and Magazine of Natural

History, sériés 7, 6 (32): 195-198.

— 1902. — Sur quelques Lepadides nouveaux de la

collection du British Muséum. Transactions of the

Linnean Society of London 8 (8): 277-295.

Hiro F. 1937. — Cirripeds of the Palao lslands. Palao

Tropical Biological Station Studies 1: 37-72.

Hoek P. P. C. 1907. — The cirripedia of the Siboga

Expédition, Pedunculata. Siboga-Ex p edi lie 31a:

1-127.

Newman W. A. 1960. — Five pedunculate cirripeds

from the Western Pacific, including two new

forms. Crustaceana 1 (2): 100-116.

— 1987. — Evolution of cirripedes and their major

groups: 3-42, in Southward A. J. (ed.), Crustacean

Issues 5, Bamacle biology.

Nilsson-Cantell C. A. 1921. — Cirripeden - Studien.

Zur Kennrnis der Riologie, Anatomie und

Systematik dieser Gruppe. Zoolagiska Bidrag fran

Uppsalal-. 75-390,

Pilsbry H. A. 1907a. — Hawaiian cirripedia. Bulletin

of the Bureau of Fisheries 26: 179-190.

— 1907b. — The barnacles (Cirripedia) contained in

the collections of the U. S. National Muséum.

United States National Muséum Bulletin 60: 1-122.

— 1909. — Report on the Barnacles of Peru. Praceed-

ings ofthe United States National Muséum 37: 63-74.

Ren X. 1983. — Five new species of suborder

Lepadomorpha (Cirripedia Thoracica) from

Chinese waters. Haiyangyu huzhao 14 (1): 74-87

[in Chinese].

Rosell N- C, 1991. — Crustacea Cirripedia

Thoracica : MUSORSTOM 3 Philippines collec¬

tion, in Crosnier A. (éd.), Résultats des Campagnes

MUSORSTOM, volume 9, Mémoires du Muséum

national d’Ffistoire naturelle (A) 152 : 9-61.

Zevina G. B. 1975. — Cirripedia thoracica of the

American Mediterranean. Trudy Instituta

Okeanologii 100: 233-258 [in Russian).

— 1982. — Barnacles of the suborder Lepadomorpha

of the world océan. II, in Fauna U.S.S.R.,

Zoological Instirute, Russian Academy of Sciences,

Leningrad 133: 1-222 [in Russian].

Zullo V. A. Ce Newman W. A. 1964. — Thoracic

Cirripedia from a Southeast Pacific Guyot. Pacific

Science 18 (4): 355-372.

Submitted on 29 October 1997;

accepted on 3 April 1998.

ZOOSYSTEMA • 1998 • 20(3)

509

A new species of the genus Pasiphaea from

the South Indian Océan (Crustacea, Decapoda,

Pasiphaeidae)

Ken-lchi HAYASHI

Department of Applied Aquabiology, National Fisheries University,

Shimonoseki 759-6595 (Japan)

hayashik@fish-u.ac.jp

John C. YALDWYN

Honorary Research Associate, Muséum of New Zealand Te Papa Tongarewa,

P.O. Box 467, Wellington (New Zealand)

Hayashi K.-l. & Yaldwin J. C. 1998. — A new species of the genus Pasiphaea from the

South Indian Océan (Crustacea, Decapoda, Pasiphaeidae). Zoosystema 20 (3) : 511-519.

KEYWORDS

Pasiphaea gelasinus ,

new species,

Pasiphaeidae,

Caridea,

Decapoda,

Crustacea,

South Indian Océan.

ABSTRACT

Pasiphaea gelasinus n.sp. belongs to a group of Pasiphaea species characterized

by a distaüv notched telson and a mid-dorsa] earina on carapace and some or

ail abdominal segments, Within this group, it is distinguished by the rostral

profile wirh anterior margin nearly vertical with a distinctive convcxicy, and

by the presence of a distinct hollow laterally un the carapace on each side of

the rostral base. The merus/ischium/basis spine formula for the first pereo-

pod is I-3/0/0 and for the second pereopod 7-12/0/0.

MOTS CLÉS

Pasiphaea gelasinus,

espèces nouvelles,

Pasiphaeidae,

Caridea,

Decapoda,

Crustacea,

océan Indien Sud.

RÉSUMÉ

Une nouvelle espece du genre Pasiphaea du sud de l’océan Indien. Pasiphaea

gelasinus n.sp. appartient à un groupe de Pasiphaea caractérisé par un telson

terminé par une encoche et la présence d’une carène dorsale sur la carapace et

sur quelques-uns ou tous les segments abdominaux. Dans ce groupe, cette

espèce se distingue par le profd de son rostre dont le bord antérieur est

presque vertical avec une convexité distincte et par la présence d’une dépres¬

sion latérale nette sur la carapace, de chaque côté de la base du rostre. La for¬

mule donnant le nombre des épines sur le mérus, l’ischion et le basis des

premiers péréiopodes est 1-3/0/0 et, pour les seconds, 7-12/0/0.

ZOOSYSTEMA • 1998 • 20(3)

511

Hayashi H.-I. & Yaldwyn J. C.

INTRODUCTION

Recently, che Muséum national d’Histoire natu¬

relle, Paris, carried out, under the leadership of

Guy Duhamel, an exploratory fishing cruise on

the seamounts off Saint Paul and Amsterdam

Islands. The area is interesting from the zoogeo-

graphical view point, but little systematic research

has been donc in the area so far. During the exa¬

mination of pclagic shrimps collccted from that

area, we foünd an undcscribed large species of

the genus Pasiphaea Savigny, 1816. Il is nearly

150 mm in total length and is characterized by

the carinated carapace and abdomen, the armed

meri of the first and second pereopods, and the

deeply forked distal margin of the telson. An

unusual dimple-likc hollow is uniquely présent

on either side of the rostral base. Ail type mate-

rial is preserved at the Muséum national

d’Histoire naturelle, Paris.

Abbreviations

CL carapace length, measured in decapod

shrimps from the orbit to the mid-dorsal

point of the posterior margin of the carapace;

CP pclagic rrawl;

MNHN Muséum national d’Histoire naturelle, Paris.

SYSTEMATICS

Genus Pasiphaea Savigny, 1816

Pasiphaea gelasintis n.sp.

(Figs 1-3)

Material examiner. — South Indian Océan.

Seamounts off Saint Paul and Amsterdam Islands,

research eompaign 1996, CP 10, 37°37-8'S -

77°51.8’E, depth 730-905 m, 3.VII. 1996: holotype

â , CL 35.2 mm (MNHN-Na 13438); paratypes 1 d,

CL 32.0 mm, 1 9 , CL 34.0 mm (MNHN-

Na 13439). — CP 6, 37°37.2’S - 77°55.5’E, depth

310-520 tn, 3i VIL 1996: paratype 9, CL 22.0 mm

(MNHN-Na 13460). — CP 12, 37°37.9’S -

77°51.7’E, depth 685-830 m, 4.V1I.1996: paratypes

1 <? , CL 36.0 mm, 2 9 9, CL 33.1, 49.5 mm

(MNHN-Na 13459).

ETTMOLOGY. — The spécifie name gelasinus is Latin

for “dimple” used as a nominative in apposition. The

English word ‘‘dimple” (French 11 fossette ") is a small

natural dent or crease in the flesh, especially on the

cheek or chin, or else a slight dépréssion in any surface,

and refers in this species to the dimple-like hollow on

each side of the rostral base.

Distribution. — Known only from the South

Indian Océan, ncar Saint Paul and Amsterdam

Islands, at depihs of310-905 m.

SlZE. — The complété male selected as the holotype is

CL. 35.2 mm. The females are CL 22.0-49.5 mm. No

ovigerous females are available, but the female of CL

33.1 mm shows many ovarian eggs through the cara¬

pace.

Diagnoses

Shell firm. Rosrrum continuous from dorsal

margin of carapace, not reaching anterior margin

of carapace. Carapace dorsaliy carinated; a dis¬

tinct hollow at each side of rostral base; bran-

chiostegal sinus small. Ail abdominal segments

wirhour Terminal spine; first segment dorsaliy

rounded; second to sixth segments dorsaliy cari¬

nated. Telson slightly shorter than sixth segment;

posterior margin deeply notched, with more

than six pairs of spines. First pereopod with one

to three spines on merus; ischium unàrmed; basis

unarmed, posterodistal end rectangular with

setae but sometimes acutely pointed; finger

slightly shorter than palm. Second pereopod

with ten to twelve spines on posterior margin of

merus; unarmed on ischium; unarmed on basis

excluding posterodistal spine; finger longer dian

palm. Ischium of third pereopod without spi-

nules on posterior margin. Pleurobranch on

eighth thoracic segment well-dcveloped, as large

as arthrobranch on sixth segment. Male holotype

35.2 mm in CL and largesr female 49.5 mm in

CL.

Description

Large pasiphaeid (Fig. 1). Rostrum triangular in

latéral view, but posterior (or upper) margin

smoothly continuous with dorsal margin of cara¬

pace, anterior (or lower) margin nearly or obli-

quely vertical with slight convexity along entire

margin, apex acute, pointed forward nearly rea¬

ching anterior margin of carapace (Fig. 2A).

Carapace dîstinctly carinated on dorsal margin

from posterior margin of rostrum to posterior

512

ZOOSYSTEMA ■ 1998 • 20(3)

New Pasiphaea from the South Indian Océan

Fig. 1. — Pasiphaea gelasinus n.sp. Holotype 6, CL 35.2 mm. Scale bar: 10 mm.

1/4-1/5 of carapace; distinct Jtepatic ridge conti¬

nuons backward with branchiocardiac ridge, but

ill-defined posteriorly; shallowly depressed

around rostral base and with a distinct liollow

laterally along each side of rostral base (Fig. 2A,

B). Branchiostegal spine arising just inside antéro¬

latéral margin ol carapace, not supported by cari-

na; apex overreaching anrerior margin (Fig. 2A),

Branchiostegal sinus small but distinct (Fig. 2A).

Only three specimens including holotype with

distal part of abdomen. First abdominal segment

rounded do rsa II y. Second ro fifth segments with

distinct dorsal carina (Fig. 2C). Sixth segment

compressed, 1.3-1.4 rimes as long as fifth seg¬

ment and about 1.3-1.5 times as long as deep

(Fig. 2D); dorsal margin sharply carinated on

anterior rwo thirds, and flattened on posterior

third; from latéral view, posterior end somewhat

produced posteriorly and dorsal margin before

posterior end slightly depressed (Fig, 2D). Telson

0.8-0.9 times as long as sixth segment, dorsally

grooved for enrire length (Fig. 2E); distal margin

deeply forked, distal end not complété in ail spe¬

cimens, but at least ten spmes recognizable on

one side in one specimen (Fig. 2F).

Eye well-developed; cornea larger than eyestalk;

from dorsal view, cornea and eyestalk obliquely

divided into two equal parts (Fig. 2G). Stylo-

cerite nearly reaching end of First antennular seg¬

ment; upper distal end small and spiniform,

directed obliquely upward (Fig, 2A). Antennal

scale 3.9-4.0 times as long as wide (Fig. 2H), and

as long as or slightly longer (1.0-1.1) than First

chela and a little shorter (0,85-0.95) than second

chela (Fig, 1); basicerite with medium-si/.ed

spine on lower distal corner (Fig, 2A), Mourh-

parts of typical shape for genus (Fig, 3A-G).

Endopod of First maxilliped small process with

single seta (Fig. 3E).

Posterodistal corner of basis of first pereopod rec-

tangular with seta, not forming typical spine but

sometimes ending in acute point; ischium unar-

med; merus with one to three spines on distal

half of posterior margin; chela of typical shape,

finger shorter (46-47% of chela) than palm;

sériés of more than ten spinules on lower mesial

margin of palm to base of immovable finger

(Fig, 3H). Basis of second pereopod ending in

distinct spine, unarmed on posterior margin.

ischium unarmed on posterior margin; merus

with seven to twelve spines on posterior margin;

palm shorter (39-43% of chela) than Angers and

as long as or slightly shorter than that of first

(Fig, 31). No spinules on ischium of thtrd pereo¬

pod (Fig. 3J). Fourth and fifth pereopods of typi¬

cal shape for genus (Fig. 3K, L).

ZOOSYSTEMA • 1998 • 20(3)

513

New Pasiphaea from the South Indian Océan

Endopod of male First plcopod composed of rwo

lobes, mesial lobe small, with many retinaculae

in central part; outer lobe large, stirrounded by

many long plumose setae (Fig. 3M). Endopod of

male second pleopod of typical shape for genus;.

appendix masculina with several long setae, not

reaching end of appendix interna (Fig. 3N).

Branchial formula as follows ;

Maxillipeds

First

Second

Third

Pleurobranchs

—

Arthrobranchs

Podobranchs

—

Epipods

Exopods

—

“

Pereopods

First Second Third Fourth Fifth

Pleurobranchs

Arthrobranchs

Podobranchs

Epipods

Exopods

Remarks

The more than sixty species in the genus

Pasiphaea Savigny, 1816 (Burukovsky 1996) can

be divided into a number of groups of species

based on characters of the telson, carapace, abdo¬

men and first nvo pereopods The présent new

species belongs ro a species group vvhich is cha-

racterized by a deeply forked telson end, by a

dorsally carinated carapace and abdomen, and by

the armed rneri o! the first and second pereo¬

pods. The distal margin of the telson must be

clearly notched, not just weakly concave. The

carapace carina may not necessarily extend ail the

way from the rostral blade to the posterior dorsal

margin of the carapace but it must extend poste-

riorly from the rostrum along a considérable pro¬

portion of the dorsal surface. The carapace dorsal

carination may not necessarily be sharp and

blade-like, it may be somewhat rounded, but it

must appear as a distinct dorsal carination. Ail

abdominal segments need not be carinated but at

least most of the third, the fourth, the fifth and

at least the anterior part of the sixth abdominal

segments must be carinated. Once again the

abdominal dorsal segmentai carination need not

ail be sharp and blade-like, but the segments

must he clearly carinated. The sharpness of the

carapace and abdominal dorsal carination, if pré¬

sent, in the genus Pasiphaea appears, at least in

some species, to be lelated to sue, âge, and posi¬

tion in molt cycle of the individual specinten.

The number of meral spines on the First and

second pereopods varies considerably with growth

or size, but the presence or absence, many or a

few in number, and their position, if présent, are

good spécifie characters.

The présent group in the genus Pasiphaea

includes the following twelve nominate species

listed in alphabetical order:

P acidifions Bâte, 1888;

P. alcocki (Wood Mason, 1891) in Wood Mason

& Alcock (1891);

P. balssi Burukovsky et Romensky, 1987;

P. harnardi Yaklwvn, 1971 (synonyms P meirin-

gnaudei Kensley, 1977; P herentsae Kensley et al.,

1987);

P faxûni Rathbun, 1902;

/? grandtcuhi Burukovsky, 1976;

P. korzuni Burukovsky, 1995;

P. multidentata Esmark, 1866;

P pacifica Rathbun, 1902;

P rathbunae (Stebbing, 1914);

P sinensis Hayashi et Miyaké, 1971;

P. tarda Kroyer, 1845 (synonym P. principalis

Sund, 1913).

Within this group, rwo species can be immedia-

tely separated from P gelasinus as follows; in

P alcocki, the first to fifth segments of abdomen

are dorsally smooth and only the sixth segment is

dorsally carinated (Wood Mason 1893); in

P. pacifica , the branchiostegal spine is placed on

the side of the carapace, more or less above the

branchiostegal sinus, well inside the antérolatéral

margin of the carapace and not extending any-

where near the antérolatéral margin, and the

merus of the first pereopod is usually unarmed

but up to four spines are présent on some spéci¬

mens (Rathbun 1902; 1904; Butler 1980).

Seven further species of this group can be

ZOOSYSTEMA • 1998 • 20(3)

515

Hayashi H.-I. & Yaldwyn J. C.

separated from P gelasinus as their second pereo-

pods are spined on the basis and often on the

ischium, as well as on the merus: in P balssi the

merus/ischium/basis spine formula for the

second pereopod is 16-19/1-2/10-16 (rostrum

extending wcll beyond Iront of carapace)

(Burukovsky & Romensky 1987); P- barnardi

second pereopod spine formula, 7-21/0-2/1-7,

though some small specimens may hâve one or

both second pereopod bases unarmed (rostrum

extends to just beyond front ot carapace)

(Yaldwyn 1971; Burukovsky 1978; Kensley 1977

as P, meiringnauder, Kensley et al. 1987 as

P. bcrcniuiê) ; P. grandi, eu la second pereopod spine

formula, 11-44/2-5/6-41, number of spincs

increasing with sizc (rostrum extending beyond

front of carapace) (Burukovsky 1976); P multi-

dentata second pereopod spine formula,

9-37/0-3/4-16 (rostrum narrow and acute, nea-

ching beyond tront of carapace) (Esmark 1866;

Sivertsen & Holthuis 1956; Zariquiey 1957,

1968); P mihbunae second pereopod spine for¬

mula, 7-17/2/3-8 (branchiostegal spine arising

from margin of carapace posterior to branchios¬

tegal sinus) (Stebbing 1914; Haie 1941); P tarda

second pereopod spine formula, 12-22/0-1/1-9

(rostrum extending well beyond front of carapace

and in mature specimens with a characteristic

convex bulge on lower border of rostrum more

or less above front of carapace) (Kroyer 1845;

Sund 1913 as P. principale, Sivertsen & Holthuis

1956; Butler 1980; Hayashi 1990).

P. gelasinus is similar to the remaining four spe-

cies of this group in having rhe ischium and basis

of the first and second pereopods unarmed,

P gelasinus has an acute rostrum, not extending

beyond the front of the carapace, with a distinc¬

tive convexicy on the nearly vertical lower margin

of the rostrum in latéral view. P acutifrons,

P faxoni and P. korzuni hâve rather similar rosirai

profiles, acute, usually not extending beyond

front of carapace but without a nearly vertical

lower margin with or without a distinctive

convexity. P. sinensis has a rostral profile rather

different from ihat of P. gelasinus, in having the

lower margin with a convexity on midlengrh,

extending beyond rostral apex and usually with a

small concavity at base ot lower margin.

P. gelasinus has a distinct hollow laterally on the

carapace at either side of the rostral base (the

"dimple” of its spécifie namc). Neither P acuti¬

frons, P faxoni, P. korzuni nor P sinensis show this

feature and, as far as we are aware, this feature is

unique to P. gelasinus among described species of

the genus Pasiphaea, though a similar hollow

may be shown in figures ot P. diaphana

Burukovsky et Romensky, 1980 (Burukovsky

1993, fig. 2.1, 2.3). Such a hollow on the carapace

is not memioned in the original 1980 descrip¬

tion of P duiphana, nor is it shown in the carlier

figures of this species published by Burukovsky

& Romensky (1979, figs 1, 2) under rhe name

Pasiphaea fidgellata non Rathbun. P. diaphana

belongs to the Pasiphaea species group characteri-

zed by a distally truncate telson, a non-carinated

carapace and non-carinated abdominal segments,

but with meri of first and second pereopods

armed.

P gelasinus has the merus of the first pereopod

with one to three spines, and the merus of the

second pereopod with seven to twelve spines.

The four species of this group discussed in the

last two paragraphs hâve first and second pereo¬

pod meral spines (based on published and unpu-

biished information available) compared to those

of P gelasinus as follows:

First pereopod

Meral spines

Second pereopod

Meral spines

P. gelasinus

1-3

7-12

P. acutifrons

1-8

10-32

P. faxoni

3-6

9-18

P. korzuni

1-5

11-20

P. sinensis

6-12

19-25

The dosest species to P gelasinus in this group

would appear to be P. sinensis at nearly the same

size, and with the abdomen also carinated on

second to fifth, and anterior two thirds of sixth,

segments. The lack of carapace hollows on either

side of the rostrum, the dissimilar rostral profile,

and the higher meral spine counts on both first

and second pereopods in P. sinensis would appear

to disringuish these two species on careful exami¬

nation.

Although missing the posterior part of the ab¬

domen, the smallest specimen of P. gelasinus

516

ZOOSYSTEMA • 1998 • 20(3)

Hayashi H.-I. & Yaldwyn J. C.

examinée! differs front ihe other larger specimens

in having a slightly different shape and propor¬

tion; the dorsal margins of the remaining first

and second segments are smooth with no trace of

a carina, the eyes are comparatively longer as are

the fingers of the second pereopod. These diffé¬

rences are probably due to the immaturity of the

specimen.

The authors are aware of sortie giant specimens

of Pasiphaea with CL up to 85 mm from the

Tasman Sea, from ofl eastern Australia and from

Southern waters of Western Australia, under

study in Japan and New Zcaland, that are ver)'

similar in many features to P gelas mus, and may

or may not prove to be conspecific. The large

specimens at least hâve a deep and anteriorly

open furrow on the carapace each .side of the ros-

trum, a carinated carapace and abdomen, and a

distally notched telson The spécifie status of

these specimens (more than one species may be

involved) is as yet undecided.

REFERENCES

Bâte C. S. 1888. — Report on the Crustacea Macrura

collected by H. M. S. “Challenger” during the years

1873-1876. Report on the Scienttfic Results oj the

Voyage of H. M. S. “Challenger " during the years

1873-1876. Zoology 24, 2 vols, 942 p.

Burukovsky R. N. 1976. — A new species of shrimp

Pasiphaed grandiculd sp.n. (Decapoda, Crustacea)

and a short outline of the genus species. Biologiya

Morya4\ 17-28 [in Russian],

— 1978. — About two species of shrimps (Decapoda,

Caridea) from the South-West Atlantic.

ZoologicheskyZhumal 57: 1729-1732 [in Russian].

— 1993. — Shrimps of genus Pasiphaea (Crustacea,

Decapoda, Pasiphaeidae) from the western part of

the Indian Océan. Byulleten Moskovskogo obshchesr-

va ispytatelei Prirody Otdel Biologicheskii 98: 33-40

[in Russian],

— 1995. — Two new species of shrimps of the genus

Pasiphaea and new records of other shrimps.

Zoologichesky Zhumal 74 (12): 121-126 [in

Russian],

— 1996. — Shrimps of genus Pasiphaea : systematics

and some remarks ou new findings (Decapoda,

Caridea) Zoologichesky Zhumal 75: 841-847 [in

Russian).

Burukovsky R, N. Ht Romensky L. L. 1979. — On

some deep-water shrimps, new for the fauna of

South-East Atlantic. Zoologichesky Zhumal 58:

328-331 [in Russian].

— 1980. — A new species of shrimp from the genus

Pasiphaea. Zoologichesky Zhumal 59: 1096-1097 [in

Russian 1.

-— 1987. — Description of Pasiphaea halssi sp.n., a

new species of shrimp from South Atlantic

(Crustacea, Decapoda, Pasiphaeidae) and poLyto-

mous key for identification of the shrimps in the

genus. Byulleten Moskovskogo obshchestva ispytatelei

Prirody Otdel Biologicheskii 92; 51-60 [in Russian],

Butler T. H. 1980. — Shrimps of the Pacific coast of

Canada. Canadian Bulletin of Fiducies and Aquatic

Sciences 202, 280 p.

Esmark L. 1866. — Carcinoloeiske bidrag tdden

skandinaviske Fauna. Forhandlinger Vtdenskabssel-

skabet Christiania 1865: 259-260, 314-316.

Haie H. M. 1941. — Dccapod Crustacea. B.A.N.Z.

Antarctir Research Expédition 1929-11131. Report

Séries B4 (9): 257-285.

Hayashi K.-l. 1990. — I’rawns, shrimps and lobsters

from Japan (55) Family Pasiphaeidae - Genus

Pasiphaea. 2. Aquabiulogy 12: 400-403 [in

Japanese],

Hayashi K.-l. & Miyaké S. 1971. — A new species of

the genus Pasiphaea from the East China Sea

(Crustacea, Decapoda, Pasiphaeidae). Proceeding of

rite Japanese Society of 'Systernatic Zoology 7'. 39-44.

Kensley B. 1977. — The South African Museum’s

Meiring Naude ctuises. Part 5. Crustacea,

Decapoda, Reptantia and Narantîa. Annals of the

South African Muséum? 4: 13-44.

Kensley B., Tramer H. A. & Griffin D. J. G.

1987. — Deepwater dccapod Crustacea from east-

em Australia (Penaeklea and Caridea). Records of

the Australian Muséum 39: 263-331

Kroyer H. 1845. — Karcmolngiske Bidrag

(Fortsaettelse), Naturhistoi'isk Tidsskrifi , new sériés

1:453-538.

Rathbun M. J. 1902. — Descriptions of new decapod

crustaceans from the we.sr coast of North America.

Proceedings of the United States National Muséum

24: 885-905.

— 1904. — Dccapod crustaceans of the northwest

coast of Nortn America. Harriman Alaska

Expédition (Harriman Alaska sériés) 10: 1-190.

Swensen E. & Holthuis L. B. 1956. — Crustacea

Decapoda (the Penaeidea and Stenopodtdea excep-

ted). Report on the Scient ifie Results of the ‘Michael

Sars" North Atlantic Dcep-Sea Expédition 1910. 5

(12): 1-54.

Stebbing T, R. R. 1914. — Stalk-eyed Crustacea

Malaçostraca of the Scottish National Antatctic

Expédition, Transactions of the Royal Society of

Edinburgh 50: 253-307-

Siind O. 1913. — The glass shrimps ( Pasiphaea ) in

northern waters. Bergens Muséums Aarhok. 1912, 6:

1-17.

Wood Mason J. 1891. — in Wood Mason J. &C

Alcock A. 1891. Natural history notes from H. M.

Indian Marine Survey Steamer “Investigator”,

518

ZOOSYSTEMA • 1998 • 20(3)

New Pasiphaea from the South Indian Océan

Commander R. F. Hoskyn, R. N. commanding.

No. 21. Note on the results of the last season’s

deep-sea dredging. Armais and Magazine ofNatural

History 6 , 7: 186-202.

— 1893. — Natural history notes from H. M. Indian

Marine Survey Steamer “Investigator”, Com¬

mander R. F. Hoskyn, R. N. commanding. Sériés

II, No. 1, on the results of deep-sea dredging

during the season 1890-1891. Annals and

Magazine ofNatural History 6, 11: 161-172.

Yaldwyn J. C. 1971. — Preliminary descriptions of a

new genus and twelve new species of natant deca-

pod Crustacea from New Zealand. Records of

Dominion Muséum 7: 85-94.

Zariquiey Alvarez R. 1957. — Decapodos espanoles

XIII. Las Pasiphaeas del Mediterraneo occidental.

Trabajo Museo Zoologia Barcelona, new sériés 2 (5):

1-31.

— 1968. — Crustaceos decapodos ibéricos.

Investigation Pesquera 32\ 1-510.

Submitted on 23 December 1997;

accepted on 17 March 1998.

ZOOSYSTEMA • 1998 • 20(3)

519

Onychiuridae (Insecta, Collembola)

d’Herzégovine en région karstique avec

description d’une nouvelle espèce

Jean-Auguste BARRA

Laboratoire de Zoologie, 12, rue de l’Université, F-67000 Strasbourg

barra@neurochem.u-strasbg.fr

Barra J.-A. 1998. — Onychiuridae (Insecta, Collembola) d’Herzégovine en région karstique

avec description d’une nouvelle espèce. Zoosystema 20 (3) : 521-528.

MOTS CLÉS

collemboles,

Onychiuridae,

systématique,

région karstique,

Herzégovine,

Europe.

RÉSUMÉ

L’étude des Onychiuridae (Insecta, Collembola) de deux massifs calcaires

dominant le site de Medjugorje a conduit à la description d'une nouvelle

espèce : Hymenaphorura nicolae n.sp. Celle-ci se caractérise par un organe

postantennaire prolongé dorsalement sur la tête, des macrochètes nettement

différenciés sur le corps -, le tergite abdominal V porte 3 + 3 macrochètes,

griffe sans dent interne. De plus, les genres Doutnacia et Fissuraphorura sont

cités pour la première fois dans cette région.

KEY WORDS

Collembola,

Onychiuridae,

systematics,

karstic mountains,

Herzegovina,

Europe.

ABSTRACT

Onychiuridae (Insecta, Collembola) from Herzegovina with description of a new

species from a karstic area. This paper deals with study of a new species

Hymenaphorura nicolae n.sp. collected in two karstic mountains near

Medjugorje (Herzegovina). This species is caracterized by an elongated post-

antennal organ, well differentiatied macrochetae on thoracic and abdominal

segments ; tergite abdominal V has 3 + 3 macrochaetae and internai teethless

claw. Moreover, the généra Doutnacia and Fissuraphorura are cited for the

first time from this area.

ZOOSYSTEMA • 1998 • 20(3)

521

Barra J.-A.

INTRODUCTION

Les travaux sur la faune collembologique de

Yougoslavie nous donnent une bonne connais¬

sance de ce groupe (ef Dunger & Zivadinovic

1989 pour la bibliographie). Les catalogues dres¬

sés témoignent de la répartition des espèces à tra¬

vers les différentes légions du pays. La province

d’Herzégovine et plus spécialement les massifs

calcaires à climat chaud et sec semblent moins

bien connus. Les collemboies Onychiuridae

décrits ou cités dans cette note proviennent de

deux massifs calcaires qui dominent le site de

Medjugorje (Barra 1993). Les récoltes ont été

faites au printemps (mai 1991) et en automne

(octobre 1994, 1995 et 1997). Tous les échan¬

tillons ont été extraits à sec.

Abréviations

aO soie médiane antérieure

c soies de l'avant-dernière rangée postérieure

g soies latéro-postérieures

oc soies centro-latérales

p 1-5 soies de la rangée postérieure

sd soies parallèles à la série paramédiane d

SYSTÉMATIQUE

Hymenaphorura nicolae n.sp.

(Fig. 1A-K)

Matériel-type. — Holorype d et 4 paratypes

(3 9 2 er I dj plus 7 paratypes juvéniles de 1 mm de

longueur). L’holotype, un paratype 9 et 2 paratypes

juvéniles ont été déposés au Muséum national

d’Histoire naturelle, Paris.

Localité-type. — Medjugorje, massif du Podbrdo

(altitude 400 m), sol brun léger couvert de lichens

blanchâtres, le 8,XJ 995. Station de récolte : Med. 17.

Étymologie. — Espèce dédiée à mon épouse qui a

récolté ce matériel.

Diagnose

Gouttière de l’organe postantennaire prolongée

dorsalement sur la tête. Grains tégumentaires dis¬

tincts. Chétotaxie asymétrique, macrochètes dis¬

tincts. Sur les tergites abdominaux de I à IV p2

plus petits que p3, sur abdomen V : 3 + 3 macro¬

chètes. Griffe sans dent interne.

Description

Longueur totale : 1,9 mm. Coloration blanche.

Les grains tégumentaires sont bien marqués sur

toute la (ace dorsale et nettement plus développés

sur le dernier segment abdominal (2,5-4 pm).

Sur les antennes et sur la tête, ceux-ci mesurent

entre 1,7-2 pm, sur les tergites : 2-2,5 pm. Les

grains autour des pseudocelles et ceux qui les

avoisinent atteignent 2,5 pm et jusqu'à 3,5 pm

sur l'abdomen V , ces grains forment des amas

plus denses en relation avec leur structure et leur

hauteur (Fig. 1E, F).

Antennes légèrement inférieures à la diagonale

céphalique. Article antennaire IV avec des soies

ordinaires, sans vésicule apicale, avec une micro-

sensille subapicale et une seconde au tiers infé¬

rieur au-dessus de l’organe antennaire III

(Fig. IB), L'organe sensoriel d'antennaire 111 est

constitué par deux masses allongées granuleuses

et deux sensilles rectilignes ; cinq papilles tégu¬

mentaires coniques les protègent ainsi que quatre

longues soies (Fig. IB). Les papilles sont bosse¬

lées, exceptionnellement l’une d’elle peut-être

bifide (Fig. IC).

L’organe postantennaire en position latéro-

ventrale c-st forme de douze à quatorze vésicules

parallèles (Fig. 1D). La dépression profonde qui

contient les vésicules se prolonge dorsalement sur

la tête en s’incurvant vers l'arrière (Fig. 1 A).

Les macrochètes dorsaux sont bien distincts. La

formule des pseudocelles dorsaux par demi-

tergite est : 10/011/11112 ; les pseudocelles ves-

tigaux mal visibles ne peuvent être localisés avec

certitude. Aulout de chaque pseudocelle on

dénombre dix à douze à 12 grains tégumentaires.

Sur les aires denses circonscrites aux pseudocelles

du corps, les soies p2 sont inférieures à p3, rap¬

port p3/p2 = 1,2-1,4 sauf sur l’abdomen IV où

p3/p2 = 2,3. Le tergite abdominal V porte 3 + 3

macrochètes (a4, p2, p5). al sont mésochètes

comme pi et p3, rapport : pl/al = 1, p2/al =

2,3. Abdomen VI avec deux épines anales,

chaque épine est insérée sur une papille à trois

couronnes de grains tégumentaires ; épines +

couronnes égales aux griffes (Fig. 11).

Les sternites thoraciques de I à III portent nor¬

malement 0, 1,1 paires de soies, mais les combi¬

naisons suivantes sont fréquentes : 0, 1 + 2,

1+2, voire 0, 2 + 2, 2 + 2, les juvéniles de 1 mm

522

ZOOSYSTEMA • 1998 • 20(3)

Onychiuridae d’Herzégovine en région karstique

Fig. 1. — Hymertaphorura nicolae n.sp. ; A, chétotaxie dorsale et répartition des pseudopores ; B, organe antennaire III et microsen-

silles (ms) latéro-ventrales ; C, organe antennaire III avec une papille bifide ; D, organe postantennaire ; E, granulation autour du

pseudocelle sur l'abdomen IV ; F, granulation autour du pseudocelle du thorax II ; G, papille génitale femelle ; H, papille génitale

mâle ; I, épine anale ; J, tibiotarse III et griffe ; K, sternite IV : aire furcale, Ps : pseudopore. Echelles : A, 200 pm ; B, C, 40 pm ;

D, 30 pm ; E, F, 27 pm ; G, H, 100 pm ; l-K, 30 pm.

ZOOSYSTEMA • 1998 • 20(3)

523

Barra J.-A.

de longueur n’onr pas encore de soies sternales.

Pas de furca. Tube ventral avec 10-11 + 10-11

soies. Plaque génitale femelle avec dix à quatorze

soies (Fig. IG), plaque génitale mâle : 16 + 16

soies (Fig. 1H), organe ventral mâle absent.

Tibiotarses sans soies particulières, de 1 à III on

dénombre 20, 20, 19 soies. Grilles sans dent

interne. Empodium terminé en aiguille avec une

courte lamelle basale, empodium égal ou légère¬

ment inférieur à la griffe (Fig. 1 J), À l’emplace¬

ment de la furca, on note dans les deux sexes une

aire à granulations fines avec deux soies, en arriè¬

re et entre les deux macrochètes latéraux huit à

neuf soies manubriales disposées le plus souvent

d’une manière irrégulière (Fig. 1 K). Sur la ligne

médiane des sternites II à IV, on peut observer

un pseudopore atypique par segment (observa¬

tion non constante).

Variabilité

La chétotaxie dorsale n’est pas symétrique : perte

de soies ou soies en surnombre. Les pseudocelles

peuvent faire défaut d’un seul côté ou des deux

côtés. La deuxième papille externe de l’organe

antennaire 111 peur être fourchue (deux observa¬

tions, mâle et femelle). Les griffes sont normale¬

ment sans dent interne, mais deux exemplaires

présentaient une minuscule dent interne. À

signaler aussi la variabilité des soies sternales tho¬

raciques.

Discussion

Les espèces d’ Hymenaphonira du « groupe sibiri-

ca » ont été redéfinies et discutées par Pomotski

(1990b). Sur la base de ce travail, Arbea &t

Jordana (1994) ont reconnu deux nouvelles

espèces de la péninsule Ibérique. Hymenaphonira

nicolae n.sp. répond aux critères définis par

Pomorski. La répartition des zones â grains torts

et contrastés place cette nouvelle espèce dans le

type «A» (sensu Arbea &C Jordana). Les soies p2

sont inférieures à p3 sur les tergites thoraciques

et abdominaux comme chez H. creatricis

Pomorski, 1990, mais cette dernière a toujours

une dent à la griffe et 4 + 4 macrochètes sur

l’abdomen V (al, a4, p2 et p5), alors que

H. nicolae n’en comporte que 3+3 (a4, p2 et

p5). L’une des papilles de l’organe antennaire III

est occasionnellement fourchue ; ce caractère,

non lié au sexe, se retrouve exceptionnellement

aussi chez H. rca Chrisdansen et Bellinger, 1980

et, d’une manière plus constante, chez H. nova

Pomorski, 1990.

Les autres Onychiuridae rencontrés sur ces deux

massifs calcaires sont :

Protaphorura du groupe fimata (Gisin, 1952)

Localité. — Les onze exemplaires proviennent tous

du même prélèvement Med. 02, récolté le 6,VJ 991.

Medjugorje colline du Podbrdo (altitude 4Ü0 m).

Petit massif couvert d'arbustes et d’épineux, laissant ça

et là des plages dénudées, sol argileux rouge entre les

blocs rocheux. Échantillon de soi (0 à —5 cm de pro¬

fondeur) contre un rocher, J.-A. Barra Jcg.

Deux adultes des deux sexes et neuf juvéniles.

Les pseudocelles dorsaux par demi-tergites sont

répartis en 33/022/33333. Tous les caractères

concordent avec les descriptions de Pomorski

(1986 et 1990a).

Mesaphorura critica Ellis, 1976

LOCALITÉ. — Espèce présente dans tous les prélève¬

ments de sol argileux sur les deux massifs calcaires

proches de la ville de Medjugorje, la colline du

Podbrdo et le Mont Krizcvac (altitude 520 m).

Récoltés en mai 1991 et octobre 1995.

Metaphorura affinis (Borner, 1902)

LOCALITÉ. — Espèce assez fréquente dans les prélève¬

ments de sol argileux sur la colline du Podbrdo, à tous

les niveaux. Récoltés en mai 1991 et octobre 1997.

Tous les exemplaires rencontrés correspondent à

M. affinis , forme B selon Ellis, 1976 avec,

comme formule pseudocellaire dorsale par demi-

tergite, 11/111/11111. Parmi les seize exem¬

plaires montés, sept présentent un organe

postantennaire avec des vésicules bifides inégales.

Comme le font remarquer Arbea & Jordana

1991, la reconnaissance de M, bipartita ne sera

possible qu’en étudiant le matériel-type ou en

utilisant le microscope électronique à balayage.

Metaphorura detiisi Simon, 1985

LOCALITÉ. — Espèce plus rare, a toujours été trouvée

524

ZOOSYSTEMA • 1998 • 20(3)

Onychiuridae d’Herzégovine en région karstique

Fig. 2. — Fissuraphorura sp. ; A, chétotaxie dorsale ; B, antenne droite, face dorsale ; C, articles antennaires III et IV face ventrale ;

D, chétotaxie des sternites abdominaux ; E, organe postantennaire ; F, aspect des grains tégumentaires sur les tergites

abdominaux IV et V, 0,8 à 3,4 pm ; G, disposition des grains polygonaux sur l’abdomen V et VI. Échelles : A, D, 65 pm ; B, C, E, G :

30 pm.

ZOOSYSTEMA • 1998 • 20(3)

525

Barra J.-A.

Tableau 1. — Tableau comparatif des soles céphaliques des différentes espèces de Fissuraphorura (selon les dessins ou les don¬

nées des auteurs). M, macrochètes : m, mësochètes ; p, microchètes.

Soies

F. cubanica

Rusek. 1991

F. deharvengi

Rusek, 1991

F. duplex

(Gama, 1962)

F. gisini

(Selga, 1963)

F. miscellanea

Barra, 1995

Fissuraphorura

sp.

sd4

oc2

en compagnie de M. a/finis. Cinq exemplaires récoltés

en mai 1991 et une dizaine en octobre 1997 sur la

colline du Podbrdo.

Cinq spécimens femelles ou immatures présen¬

tent, par demi-tergite, la formule pseudocellaire

dorsale suivante : 11/122/22221 Selon Simon

(1985), la forme C de Ellis (1976) se réfère à

M. denisi.

Doutnacia xerophîla Ruselc, 1974

Localité. — Espèce récoltée en nombre sur les deux

massifs calcaires et dans tous les échantillons de sol en

mai 1991 et octobre 1994, 1995, 1997.

Espèce très commune, présente dans tous les pré¬

lèvements en nombre variable.

Fissuraphorura sp.

(Fig. 2A-G)

Materiel étudié. — Un seul exemplaire non adulte

déposé au Laboratoire d'Entomologie du Muséum

national d’Histoire naturelle, Paris.

LOCALITÉ. — Medjugorje, colline du Podbrdo (altitude

400 m). Petit massif couvert d’arbustes et d’épineux,

laissant çà et là des plages dénudées à sol argileux

rouge entre les blocs rocheux. Dans sol argileux (0 à

—5 cm de profondeur) contre un rocher, le 6.V.1991.

Extraction à sec. Station de récolte : Med. 2 ; J.-A.

Barra leg.

Description

Taille : 0,95 mm. Coloration blanche. Le grain

tégumentairc est bien marqué, dessinant une

sorte de pavage à éléments polygonaux (Fig. 2G)

sur la face dorsale de la tête, du corps et sur les

articles antennaires III et IV. Sur le thorax et les

segments abdominaux 1 et II, les grains mesurent

1,7 pm, sur l’abdomen III à V, 2,5 à 3,4 pm, sur

le dernier segment abdominal ceux-ci atteignent

2,5 pm, mais sont nettement arrondis. Les grains

résultent de la fusion de plusieurs grains pri¬

maires avec un halo clait au centre (Fig. 2F).

Antennes plus courtes que la tête. Les articles

antennaires I, 11, III, IV ont une taille relative de

l’ordre de 18, 24, 43, 21 pm. Les cinq senxilles

cylindriques (a-e) de l’article IV sont bien visibles

avec b et e plus développées ; les mictosensilles f

et g sont présentes. La vésicule apicale globuleuse

et granuleuse se détache du tégument par son

pédoncule L’organe antennaire III comprend

deux microsensilles tubulaires logées derrière un

repli tégumentaire, encadrées par trois sensilles

épaisses dont depx courbées l’une vers l’autre, sur

la face ventrale une sensille épaisse en position

transversale (Fig. 2B-C).

La chétotaxje dorsale de la tête et du corps

(Fig. 2A) met en évidence la répartition des diffé¬

rentes soies. Sur la tête sont macrochètes les soies

aO (19 pm), g3 (22 pm) et p5 (25 pm), sont

mésochètes sd4 (16 pm), oc2 (16 pm), c5, pl et

p3. Organe postantennaire à six vésicules allon¬

gées caractéristiques du genre (Fig. 2E), les grains

tégumentaires bordant la goutttère mesurent

2,5 pm. La répartition des soies est symétrique,

certaines d’entre elles sont épaissies : p5 d’abdo-

526

ZOOSYSTEMA • 1998 • 20 (3)

Onychiuridae d’Herzégovine en région karstique

men II, p3 d’abdomen III, p4 d'abdomen IV, p5

d’abdomen V, al d’abdomen VI et la soie anté¬

rieure latérale d’abdomen II et 111. Sur la face

ventrale l’aire génitale porte 1 t- 1 soies épaissies

(Fig. 2D) ; les stérilités thoraciques de I à III por¬

tent 0, 1, 1 paires de soies..

La répartition des pseudocelles par demi-tergite

correspond i) la formule suivante 11/122/22221 ;

de forme circulaire (8-9 pm) ils présentent un

bord en forme de croissant avec généralement

trois crêtes internes.

Pattes sans soies différenciées, griffes sans dent ni

empodium. Tube ventral avec 7 + 7 soies (Fig.

2D).

Discussion

La répartition et le nombre des soies dorsales sur

le thorax et sur l’abdomen sont remarquablement

constants parmi les espèces décrites du genre à

l’exception de F deharvengi Ruselc, 1991 où m4

fait défaut sur l’abdomen IV. Sur l’exemplaire,

l’absence des soies m3 (abdomen I et II) et p4

(abdomen l et III) ne peut être considérée

comme définitive. Lucianez &T Simon ( 1992) ont

utilisé les soies céphaliques comme critère discri¬

minatoire accessoire des espèces (Tableau 1). De

toutes les espèces, F. cubanua Rusek, 1991 est

celle qui possède le plus grand nombre de rnicro-

ebètes céphaliques ; cet exemplaire est plus

proche de F duplex (Gama, 1962).

Si l'analyse détaillée de la chétotaxie rend la

détermination difficile, les Fissuraphorura ont

toutes un tégument à granulation fine. F debar-

vengi présente un tégument plus grossier sur

l’abdomen VI et deux épines, caractères interpré¬

tés par fauteur comme une possible écomorpho-

se. Ce spécimen se distingue aisément par son

tégument grossier sur l’ensemble de la face dorsa¬

le et la soie al du tergite VI seulement épaissie à

la base. Le tissu adipeux est normal s le tube

digestif est visible, rempli de nourriture, faits qui

ne plaident pas en faveur d’une forme écomor-

phique.

CONCLUSIONS

Le climat sec et chaud qui règne sur ces deux

massifs conditionne les populations de la faune

du sol dépendantes des précipitations. Les prélè¬

vements de mai et d’octobre ne sont pas très

favorables, les formes juvéniles sont nombreuses

et les adultes limités à quelques exemplaires.

Compte tenu des variations individuelles chez les

Onychiuridae, l’identification de certaines

espèces s’avère difficile. Cette note complète nos

connaissances sur les collernboles de Yougoslavie

et les genres Doutrutciel et Fi s s u rapho ru ra soni

cités pour la première fois dans cette région.

RÉFÉRENCES

Arbea J. I. & Jordana R. 1991. — Colémboles de

Navarra (Norte de la Peninsula Ibérica) I. Orden

Poduromorpha (Collenabola). Publicaciones de

Biologie Universidad de Navarra , sérié Zoologica

22: 1-149.

— 1994, — Guano nuevas especies de la Familia

Onychiuridae de la Peninsula Ibérica (Collembola,

Poduromorpha). Publicaciones de Biologia

Universidad de Navarra , sérié Zoologica 24: 39-59.

Barra J.-A. 1993. — Trois nouvelles espèces de

Neanutidae d’Herzégovine en région karstique

(Insecta, Collemboia). Bulletin du Muséum national

d’Histoire naturelle, Paris, série 4, 15 : 69-78.

— 1995. — Nouveaux Collernboles Podurontotphes

des sables littoraux (partie terrestre) de la Province

du Natal (Rép. Sud Africaine) (Insecta,

Collembola). Journal of African Zoology 109:

125-139.

Dunger W. & Zivadinovic J. 1989. — Taxonomie

und Verbreitung der Gattung Pohoinia Willem,

1902 (Hexapoda, Collernboîà) in Bosnien und

Hercegovina (Jugoslawicn). Abhandlungen und

Beriehte Nu tu rku ndeMuséums (Jêrlitz 63:1-12.

Ellis W. N. 1976. — Aummn fauna of Collembola

front Central Crete. Tijdschrift l)oor Entomologie

119:221-326.

Lucianez M. J. &c Simon J. C. 1992. — Lin nuevo

genero y dos nuevas tribus de Tullbergiinae

(Collembola, Onychiuridae) de la Peninsula

Ibérica. Eos 68: 105-114.

Pomorski R. ]. 1986. — Morphological-systematic

studies on the variability of pseudocellae and saine

morphologicai characters in "arm a tus group’’

(Collembola, Onychiuridae). Part I. Onychiurus

(Prntapborura) Jtmatus Gisin, 1952. Bulletin

Entomologique de Pologne 56: 531 -556.

— 1990a. — Ibidem. Part II. On synonyms within

the “annatus- group”, with spécial référencé to dia¬

gnostic characters. Annales Zoologici Warszawa 43:

536-576.

— 1990b. — New data on the genus Hymenapborura

(Collembola, Onychiuridae) front Europe. Bulletin

ZOOSYSTEMA • 1998 • 20(3)

527

Barra J.-A.

de la Société entomologique suisse 63 : 209-225. bohemoslovaca 88: 145-155.

Rusek J. 1991. — New tropical Tullbergiinae Simon J. C. 1985. — Colémbolos de suelos de sabinar

(Collembola, Onychiuridae). Acta entomologica en la provincia de Guadalajara. Eos6l: 293-318.

Soumis le 29 août 1997 ;

accepté le 20 février 1998.

528

ZOOSYSTEMA • 1998 • 20(3)

Review of Giraud’s types of the species of

Synergus Hartig, 1840 (Hymenoptera, Cynipidae)

Juli PUJADE-VILLAR & Palmira ROS-FARRÉ

University of Barcelona, Department of Animal Biology,

645, Diagonal Avenue, E-08028 Barcelona (Espagne)

pujade@porthos.bio.ub.es

palmira@porthos.bio.ub.es

Pujade-Villar J. & Ros-Farré P. 1998. — Review of Giraud's types of the species of

Synergus Hartig, 1840 (Hymenoptera, Cynipidae). Zoosystema 20 (3): 529-540.

KEYWORDS

Hymenoptera,

Cynipidae,

synergini,

Synergus,

Giraud,

lectotypes,

systematics,

synonymies,

hosts.

ABSTRACT

In this work the typical sériés of the remaining Synergus species described by

Giraud are studied. The lectotypes of each one are designated. Synergus

consobrinus is redescribed, S. variabilis Mayr, 1872 (= S, cerridis n.syn.)

( = S. confirmas n.syn.) and S. diaphanus are characterized. The respective

synonymies for the rest of the studied species are cstablishcd. Finally, biologi-

cal data of some of the species are given and the distribution area of some of

them is widened.

MOTS CLÉS

Hymenoptera,

Cynipidae,

synergini,

Synergus,

Giraud,

lectotypes,

systématique,

synonymies,

hôtes.

RÉSUMÉ

Révision des espèces-types du genre Synergus Hartig, 1840 (Hymenoptera,

Cynipidae). Dans ce travail, nous étudions le matériel des séries de types

décrites par Giraud concernant le genre Synergus. Les lectotypes de ces séries

sont désignés. Nous redécrivons l’espèce Synergus consobrinus ; les espèces

S. variabilis Mayr, 1872 {= S. cerridis n.syn.) (= S. confomiis n.syn.) et S. dia¬

phanus sont caractérisées. Nous établissons les synonymies concernant les

autres espèces du genre. Pour chaque espèce nous mentionnons les princi¬

pales caractéristiques biologiques et la répartition.

ZOOSYSTEMA • 1998 • 20(3)

529

Pujade-Villar J. & Ros-Farré P.

INTRODUCTION

Joseph-Étienne Giraud (1808-1877) was one of

the greatest cynipidologists of the last century.

His Cynipidae studies, though few in number

(Giraud 1859. 1866. 1868a, b, 1871; Darboux

&C Houard 1907; Laboulbène 1877), greatly lead

to the récognition of a large number of cynipid

gall-forming species. He described more

than forty species as well as the new genus

Dryocosmus. A part of the author’s research was

compiled in an unpublished manuscript deposi-

ted in the Muséum national d’Histoire naturelle,

Paris (MNHN). Houard (1911) published a ver¬

sion of Giraud’s manuscript. In this publication

eleven new Synergus species are described:

S. apertns , S. cerridis, S. clavatus, S. confortais,

S. consobrinus, S. diaphanus , S. hartigi, S. inflaitis ,

S. longiventris, S, subterraneus and S, vesiculosus.

The study also comains descriptions of four new

cynipids not belonging to the Synergtts species.

The only species of Synergus thar lias previously

been examined is S. itpertus, which was synoni-

myzed with Saphonecrus undulatus (Mayr 1872)

by Pujade-Villar & Nieves-Aldrey (1990).

Giraud’s material was drawn largely from Central

Europe. In his manuscript there is no mention

either of the origin of specimens.

The study of the Synergus species type material is

especially interesting in the Palaearctic zone,

since some doubt has been cast on thirty-one of

the fifty-nine species described.

Abbreviations

NMW Nacurhistorisches Muséum, Wien

MNCNM Museo Nacional de Ciencias Naturales,

Madrid

MNHN Muséum national d’Histoire naturelle, Paris

ZSBS Zoologische Staatssammlung, München

SEM METHODS

The SEM photographs of the type material

Synergus consobrinus were taken without any coa-

ting. The voltage used ranged becween 200 and

1000 V, depcnding on the specimens. In this way

we were able to obtain pictures while preserving

the specimen types without any manipulation.

The pictures of S. variabilis were obtained by

gold coating of one of the dissected S. cerridis

specimens.

LIST OF SPECIES

Giraud’s nominal species described in Synergus

genus by Houard (1911) are dealt with below.

The heading for each species treatment is the

currently accepted name. For each species the

lectotypes ate designated. The synonymous

liâmes, hosts, distribution and information

concerning type material are given. The main

taxonomie aspects are discussed. Girâud’s accep¬

ted species and S. variabilis Mayr are characreri-

zed and differentiated from the closely related

species using a taxonomical key.

Saphonecrus haimi (Mayr, 1872)

Sapholitus haimi Mayr, 1872. Verh. Ges. Wien 22:

723. Types in NMW.

Synergus clavatus Giraud (in Houard 1911: 333),

n.syn.

Saphonecrus haimi (Mayr), in Pujade-Villar et Nieves-

Aldrey, 1990: 53 (misspelling), n.syn.

TYPE MATERIAL. — Lectotype 9, and paralectotypes:

1 d and 4 9 9 of Saphonecrus haimi hâve been desi¬

gnated ni Pujade-Villar tk Nieves-Aldrey (1990).

Lectotype 9 of Synergus clavatus Giraud, here desi¬

gnated, mounced on micro-pin; paralectotypes,

32 6 6 and 5 9 9 , ail of them mounted on micro¬

pins.

DISTRIBUTION. — Central Europe and Israël in

Cynipid galls of Quercus cerris and Q. ithaburensis.

HOSTS. — Hymenoptera, Cynipidae, Cynipini:

Cbilaspis nitidus (Giraud, 1859) ô, Neurotenu lanugi-

nosiu (Giraud, 1859) ô, N. salions (Kollar, 1857) ô and

N. minutas (Giraud. 1859) ô. Diptera., Cccido-

myiidac : Janetia cerris (Kollar, 1850).'

Communes

This species and Saphonecrus ondulants (Mayr,

1872) are close to Synergus genus because they

hâve pronatal canna as do most of the species of

Synergus Pujade-Villar & Nieves-Aldrey (1990).

The Neuroterus minutus galls is a new host record

of Saphonecrus haimi.

530

ZOOSYSTEMA • 1998 • 20(3)

Giraud’s types species of Synergus

Synergus variabilis Mayr, 1872

(Figs 1-3)

Synergus variabilis Mayr, 1872. Ver h. Ges. Wien 22:

702. Types in NMW.

Synergus cerridis Giraud (in Houard, 1911: 331-332),

n.syn.

Synergus conformis Giraud (in Houard, 1911: 333-

334), n.syn.

Synergus «m'ctt/tw Vasileva-Samnalieva, 1986, n.syn.

Tyi'E MATERIAI. — The paralectorypes of Synergus

variabilis Mayr (1 d and 2 2 9), mounted on micro-

pins, hâve been exami ned. Lectotype S ol Synergus

cerridis Giraud, here designated, mounted on micro-

pin; paralectorypes: 15 dd and 26 2 9 , ail of them

mounted on micro-pins. Lectotype d of Synergus

conformis Giraud, here designated, mounted on

micro-pin; paralectotypcs: 17 tî d and 16 9 9,

mounted on micro-pins; 1 2, on card.

Distribution. — Central Europe and Israël in

Cynipid galls ot Qucrcus cerris and Q. itbaburensis.

Hosts. — Hymenoptera, Cynipidae, Cynipini:

Andricus grossitlariac Giraud, 1859 d 2, Aphelonyx

cerricola (Giraud, 1859) ô, Chilaspis nitidus (Giraud,

1859) ô, Dryocosmus ccrriphilus Giraud, 1859 ô,

Neuraicnis lanuginosus (Giraud, 1859) 6 and

N. macrnpierus (J lartig, 1843) ô; Synergini: Synophrus

politus Hartig, 1843. Diptera, Cecidomyiidae:

Dryomyia circinnans (Giraud, 1861) and Janelia cerris

(Kollar, 1850).

COMMENTS

This specics is a cynipid galls inquiline in Q. cer¬

ris belonging to Section 11 of Synergus. Kieffer

(1902) dilïereruiated four chromatic variations.

Quinlan (1978) redescribed this species but he

made a number of errors. Firsrly, rhe tarsal claws

are not simple, radier they hâve a small rooth,

nor pointed and usually hidden b)’ the arolium;

secondly, and more imponantly: ail type material

examined does not hâve a ptonotal carina

(Fig. IA) although when viewed dorsally the pro-

notum is angular at the latéral corners (Fig. IB).

Only the species Synergus plagiotrochi does not

hâve a latéral ptonotal caritiae. This species is

typical in the Mediterratiean région on cvnipid-

galls ol Pbtgiotrochus. This species is closely rela-

ted to 5. apicalis and S. romndivemris.

In order to distinguish these morphologically

closely related species a dichotomie key is provi-

ded:

1. Latéral pronotal carina présent. Punctures on upper face usually distinct. Inquilines

frequently associated with shoots Quercus , usually deciduous Quercus .*

— Latéral pronotal carina absent. Punctures on upper face usually obsolète.2

2. Radial cell open on margin. In dorsal view latéral pronotum is rotund. Punctuation

on upper face and frontal carina always obsolète. Inquiline in galls of species of

Plagiotrochus genus on evergreen oaks Q. ilex and Q, coccifera .

. S. plagiotrochi Nieves-Aldrey et Pujade-Villar, 1986

— Radial cell closed. In dorsal vision the pronotum is angular in the latéral corners.

Punctuation on upper face and frontal carina présent in larger specimens. Inquiline

in several galls of Neuroterus associated to Q. cerris, mainly N. macropterus and

N. lanuginosus ... S. variabilis Mayr, 1872

* A complex of two closely related species: fî. apicalis Hartig, 1841 and S. rotundiventris Mayr, 1872, that cannot

be satisfactorlly distingulahed by external morphological characteristics. The Intraspecific variation is great and the

limits are not clear,

Other notable features include: vertex not punc-

tuated in small specimens (Fig. 2A) and with

weak punctuation in large specimens (Fig. 2B);

latéral frontal carinae présent but not very clear

(Fig. 2); second antennomere very long in large

specimens, always longer than they are broad

and larger than third antennomere; male with

first flagellomere weakly expanded basally and

apically (Fig. 3); mesonotum with interrupted,

sharp and widely-spaced transverse rugae

ZOOSYSTEMA • 1998 • 20(3)

531

Pujade-Villar J. & Ros-Farré P.

(Fig. 1); notauli shailow and short, faint or

absent at least in anterior one third of mesoscu-

tum; notauli shailow and short, faint or absent at

least in the anterior third of mesoscurum, and

médial impression ver y short or absent (Fig. 1 B);

mesopleuron usually ventrally smooth and shiny

(Fig. IA); radial cell closedon margin.

The materia! of Giraud belonging to S. eerridis

and S. cünjomns is the same as that belonging to

S. variabilis. This sériés has ail of the chromatic

variations described by Kieffer (£902). Even

though the type sériés of Synergus cerne oins

Vassileva-Samnalîeva were not examined, after

the examination of the type material of S. varia'

bilis Mayr, 1872 and Giraud’s type sériés as well

as the original description of the former species,

we conclude that Synergies cerricolus is a junior

synonym of S. variabilis.

The mention of Neuroterus saliens and Janetia

cerris as a Synergus variabilis host is interesting

because they are new for 5. variabilis . Finaily, the

specimens of S. variabilis collected in Janetia cer¬

ris , deposited in Giraud's collection, are impor¬

tant as this species was mentioned in

Cecidomyiidae galls' (Dryomyia circinnans

Giraud, 1861, by Mayr, 1872) and it has been

considered dubious as it has not been confirmed

by subséquent authors.

Synergus thaumacerus (Dalman, 1823)

Cynips tbaumacera Dalman, 1823. Analecta Ente. 96.

Location of types unknown.

Synergus luteus Hartig, 1840. Z. Ent. Germar 2\ 199.—

Mayr 1872.

Synergus klugii Hartig, 1840. Z. Ent. Germar 2: 199. —

Mayr 1872.

Synergus cannants Marri g, 1841. Ent. Germar 3: 348.—

Mayr 1872.

Synergus thaumatocerus Dalla Torre, 1893. Cat. Hym.

2: 114 (unjustified emendation).

Synergus vesiculosus Giraud (in Houard 1911:

323-234), n.syn.

Synergus inflatus Giraud (in Houard 1911: 324-325),

n.syn.

TYPE MATERIAL. — Giraud’s collection. Lectotype d

of Synergus vesiculosus Giraud, here designated, moun-

ted on micro-pin; paralectotypes: 5 do and 10 9$,

Fig. 1. — Thorax of S. variabilis Mayr; A, latéral view. B, dorsal

view. PR, pronotum. See text for additional discussion. Scale

bars: a, 200 pm; B, 500 pm.

ail of them mounted on micro-pins. Lectotype 6 of

Synergus inflatus Giraud, here designated, mounted on

micro-pin; paralectotypes: 22 d d and 27 9 9, ail of

them mounted on micro-pins.

Distribution. — Western Palaearctic in Cynipid

galls of Quercus cerris, Q. faginea, Q. humilis ,

Q. petraca, Q. pyrenaica and Q. robur.

HOSTS. — Only Cynipini: Andrtcus anthracinus

(Curtis, 1838) ô, À. curvator Hartig, 1840 3 9 ,

A. cydoniae Giraud, 1859 d 9, A. singulus Mayr, 1870

532

ZOOSYSTEMA • 1998 • 20(3)

Fig. 2. — Head of S. varlabilis Mayr, dorsal visw; A. small specimen: B, large specimen. A, antennomere; FC, facial carinae. See

text for addltional discussion. Scale bars: 200 pm.

ô, A. galhmmuufornm (Boyer de Fonscolombe,1832) D. nervosus (Giraud,! 859) 3 9, Neuroterus salions

(= suflator Mayr, 1882) 3 9 . A. gallaetinctoriae (Kollar, 1857), 3 9 , N. saliens (= glandiformis

(Olivier, 1791) ô, Aphelonyx cenicola (Giraud,1859) Kaltenbach, 1867) ô, N. quercusbaccarum (L., 1758)

ô,Cynips disticha Hartig, î 840 ô, C. quercusfolii L., 3 9, N. quercusbaccarum (= lenticularis Olivier, 1791)

1758 ô, Dryocosmus cerriphilus Giraud, 1859 ô, ô, N. tricolor (Hartig, 1841) 3 9, N. tricolor (= fitmi-

ZOOSYSTEMA • 1998 • 20(3)

533

Pujade-Villar J. & Ros-Farré P.

Fig. 3. — First male antennomeres of S. variabilis Mayr. Scale bar: 200 pm.

permis Hartig, 1841) ô, Trygonaspis megaptera (Panzer,

1801) S 9, T. megaptera (= renum Harrig, 1840) ô,

T. mendesi Tavares, 1901 ô and T. synaspis (Hartig,

1841) (?$, Synophrus politus Hartig, 1843.

COMMENTS

The characters for recognizing S. thaumacerus are

described in Nieves-Aldrey & Pujade-Villar

(1986). Giraud’s material of S. vesiculosus and

S. inflatus do not differ from that of S. thaumace¬

rus. The hosts Andricus gallaetinctoriae and

Dryocosmus ccrriphilus and Synophrus politus are

new to 5. thaumacerus.

Synergies stibterraneus Giraud

(in Houard, 1911:335)

Synergus latifrons Nieves-Aldrey et Martin-Chicote,

1985. Bol. Asoc. esp. Entom. 9: 151. n.syn. Types in

MNCNM.

Type material. — Of Giraud’s collection.

Lectotype 2, here designated, mounted on micro-pin;

paralectotypes; 8 9 2 mounted on micro-pins; 3 V?

on carts.

Distribution AND hosts. — Only Cynipini: Spain

in Andricus sieboldi (Hartig, 1843) ô on Quercus pyre-

naica and in central Europe (country unknown) in

Andricus rhyzomae (plant-host unknow, probably

Q. humilis (= pubescens) because Giraud (in Houard,

1911) collected A. rhyzomae in Q. humilis).

COMMENTS

The width ol the head in frontal vision is a very

spécifie character of this species, since the trans¬

facial line is at least twice as long as the height of

the compound eye. The biology is associated

with subterraneous galls of Andricus.

Synergus albipes Hartig, 1841

Synergus albipes Hartig, 1841. Z. Ent. Gerrnar 3: 349.

Types in ZSM.

Synergus erythrocerus Hartig, 1841. Z. Ent. Gemiar 3:

349. - Mayr 1872.

Synerçnts varias Hartig, 1841. Z. Ent. Gerrnar 3: 349. -

Eady 1952.

Synergus tristis Mayr, 1873. Verh. Ges. Wienll: 715. -

Eady 1952.

Synergus nervosus Hartig, sensu Ross, 1951. Trans Soc.

Brir. h'mom. 1 (3): 92. - Eady 1952.

Synergus nervosus f. albipes Ross, 1951. Trans Soc. Brit.

Entom. 2 (3): 91. - Eady 1952.

Synergus nervosus f. tristis Ross, 1951. Trans Soc. Brit.

Ëntorn. 2 (3): 93. - Eady 1952.

Synergus mutabilis Dettmer, 1924. Natuurbist.

Maandbl. 13:147. — Wiebes-Rijks 1979.

Synergus hartigi Giraud (in Houard 1911: 332) n.syn.

TYPE MATERIAL — Of Giraud’s collection.

Lectotype <$, here designated, mounted on micro-pin;

paralectotypes: 10 3 S and 11 9 9 , ail of them

mounted on micro-pins.

DISTRIBUTION. — West Palaearctic in Cynipid galls

of Quercus cerris , Q. canariensis, Q. faginea ,

Q. humilis , Q. petraea, Q. pyrenaica, Q. suber and

Q. robur.

534

ZOOSYSTEMA • 1998 • 20(3)

Giraud’s types species of Synergus

Hosts. — Only Cynipini: Andricus antbracinus

(Curtis, 1838) ô, A■ amblycerus (Giraud, 1859) ô,

A. caliciformis (Giraud, 1859) ô, A. callidoma Hartig,

1841 ô, A. capudmedusae (Harrig, 1843) ô. A, clemen-

tinae (Giraud, 1859) ô, A. congloméra tus (Giraud,

1859) ô, A. coriaritn Harrfg, 1S43 ô, A. corruptrix

(Schleclnendai, I 8~0) ô A. curvator Hartig, 1840

S 9, A. fecundator (Hartig, 1840) ô. A. gallaetincto-

riae (Olivier, 1791) ô, A. gallaeurnaeforints (Boyer de

Fonscolombc, 18.32) ô, A. glutinosus (Giraud, 1859) ô,

A. hartigi (Harrig 1843) ô, A. hungaricus (Hartig,

1843) 6, A. glohuli (Hartig, 1840) ô, A. kolltlri

(Hartig, 1843) 6, A. Itgnirolus (Hartig, 1840) 2,

A. htcidm (Hartig, 1843) ô, A. nudtts (Adler, 1881) ô

G malpighii Adler, 1881), A polyctrus (Giraud, IS59)

ô, A. quadrilt néants Hartig, 1840 ô, A. qucrcuscalicis

(Burgsdorf, 1679) ô. A quenusramuli ( L., 1761)

(= autumnalis Harrig, 1840) ô, ,4, quercustozae (Bosc,

1792) ô, A. umimmnis (Giraud, 1859) 6, A. salitarius

(Boyer de Fonscolombe, 1832) ô, Cynips a garnit

Hartig, 1840 6, C disneba Hartig, 184Ô ô, C. longi-

ventris Harrig, 1840 ô. C. quercus (Fourcroy, 1785) ô.

C. quercusfoftrl... 17586. Neuroterus atbipes (Schcnck.

1863) ô (= la evùculus Schcnck, 1863). N. lanugmosus

Giraud, 1859 ô, N. numtsmalts (Fourcroy. 1 T 85) 6,

N. nuntismalis (= vesicator Seblccbtendal, 1870) t?2,

N. sa/iens (Kollar, 1857) ô, H. quercusbaccarunt (L.,

1758) 6 9, iV. quercusbilccuntm (= lenticularis Olivier,

1791) ô, N, triçolor (Hartig, 1841) <5$, N. tricvlvr

(= fittuipcnnis Harrig, 1841) 6, Plagia troc ht es «menti

Tavares, 1902 9, Trygonaspis bruneiemtis Tavares,

1902 ô, T. megapcera (Pauzer, 1801) (= renum Harrig,

1840) ô, 71 mendesi Tavares, 1901 ô and T. synaspis

(Hartig, 184l) ci 9.

COMMENTS

This species has a large number of hosts, most of

whicb are agamie in form. Nevertkeless, it seems

not to be a spécifie feature. This species is very

closely related to S. nervosus Hartig, 1840, and

sometimes there are no clear différences bervveen

them. The first flagellomerc segment (of ail the

males of S. hartigi bave) is moderately expanded

apically and, usually the second antennomere in

both sexes are as long as they are broad or slighc-

ly longer. For rhese reasons, we believe thar

S. hartigi is synonymous of S. alhipes. Andricus

hartigi is a new host of this species.

Synergus gallaepomiformis

(Boyer de Fonscolombe, 1832)

Diplolepis gallac-pomifhrmis Boyer de Fonscolombe,

1832. Ann. Sc. Nat. 26: 195. Location of types un-

known.

Synergusfiicialis. Hartig, 1840, Z F.ni. Germar !: 199. -

Dalla Torre & Kieffer 1910.

Synergus vulgam Hartig, 1840. Z. Ent. Germarl: 198. —

Ross 1951.

Synergus bispintts. Hartig, 1841. Z. Ent. Germar 3:

347. - Mayr 1872.

Synergus erythrocerus var. 1 et 2 Hartig, 1844. Z. Ent.

Germar 3: 349.

Attlax albinervis Snellen van Vollenhoven, 1869:

Tijdschr. Ent. 12: 126.- KiefFer 1902.

Synergus pmmformts Kieffer, 1898. Wien Ent. Zeit. 17:

264 (not Ashmead, 1885), unjustified emendation.

Synergus gallaepomiformis gallaepomiformis Dalla

Torre et Kieffer, 1910. Das Tierreich 24: 621.

Synergus gallaepomiformis galUcus Dalla Torre et

Kieffer, 1910. Dits Tierreich 24: 622.

Synergus gallaepomiformis var minima Kieffer, 1899.

André. Spec. llym. Europ. 7 (I): 358.

Synergus gallaepomiformis minimus Kieffer, Dalla

Torre et Kieffer, 1910. Das Tierreich 24: 622.

Synergus maculants Tavares, 1920. Mern. Port. Sc. Nat.

ZooL 4: 47-49. — Nieves-Aldrey et Pujade-Villar,

1986.

Synergus longiventris Giraud (in Houard, 1911: 318-

319). n.syn.

Type MATERIAE. — Giraud’s collection. Lectotype 2,

here designated, mounted on micro-pin; paralecto-

types: 9 <? â and 17 2 2, ail of them mounted on

micro-pins.

DISTRIBUTION. — West Palaearctic in Cynipid galls

of Quercus terris, Q. faginea , Q. humilis, Q petraea.

Q. pyrenaica. and Q. robur.

HOSTS. — Only Cynipini: Andricus antbracinus

(Curtis, 1838) 6, A. callidoma Hartig, 1841 ô,

A. capudmedusae (Hartig. 1843) ô. A. clementinae

(Giraud, 1859) o, A. conglomeratus (Giraud, 1859) ô.

A. codantes Hartig, 1843 6. A. curonatus (Giraud in

Houard 1911)6, A. rormptrix (Schlechtendal, 1870)

ô, A. curvator Hartig. 1840 6 9, A. curvator (- collant

Hartig, 1840) ô, A. fecundator (Hartig, 1840) 6,

A. gdlfactinçtoriae (Olivier, 1791) ô, A. gcdlaeumaefor-

tnis (Boyer de Fonscolombe, 1832) o, A glandulae

(Hartig, 1840) 6, A glutinosus (Giraud, 1859) 6,

A. hartigi (Hartig, 1843) ô, A hungaricus (Hartig,

1843), 6, A globali (Harrig, 1840) ô, A. knllari

(Hartig, 1843*) ô, A. légitimas Wiebes-Rijks, 19R0 ô,

A. lignicolus (Hartig, 1840) V. A. nudus (Adler, 1881)

ô (= mmlptgldi Adler, 1881), A. paradoxus (Rad, 1866).

A. quaarilineatus Hartig, 1840 ô, A. quercuscalicis

(Burgsdorf, 1679) ô, A. quercuscorticis (L., 1761) ô,

ZOOSYSTEMA • 1998 • 20(3)

535

Pujade-Villar J. & Ros-Farré P.

A. quercusrodkh (Fabricius, 1798) ô, A. quercusradicis

(= trilincatns Harrig, 1840) 5 9, A, quercusramuli

(Linnaeus, 176!) 5 5, A. quercusramuli (= autumnalis

Hartig, 1840) ô, A, quercustozae (Bosc, 1792) ô,

A. seminationis (Giraud, 1859) 6, A. soUtarius (Boyer

de Fonscolomhe, 1832) ô, A sieba/di (Harrig,) ô,

Callirhytis gla.nd.ium (Giraud, 1859) 6, Cynips disticba

Hartig, 1840 ô, C. divisa Hartig, 1840 ô, C. lu agi ven¬

ir is Hartig. 1840 ô, C. longiventris Hartig, 1840

(= similis Adler, 1881) 5 2, C. qucrcusfolti Linnaeus,

1758 Ô, C. quercusfolii (= Mschenbergi Selechtendal,

1870) 5 9, Neuroterus a/bipes ( Scbenek, 1863) ô

(= iaevisculus 8’chetuk, 1863), N, hmugimsus Giraud,

1859 ô, N, nutnism/tlis (Fourcroy, 1785) ô N. quercus-

baccarum (Linnaeus, 1758) 5 2, N, qucrcusbaccantm

(= Icnticularis Olivier 1791) ô, A L erkolor (Hartig,

1841) 52, Al irkalor (= fumipcnnis Hanig, 1841) ô,

Trygonsspis megapma (Pan/er, 1801), A megaptera

(= renum Hartig, 1840) ô, and f. synaspts (Hartig,

1841) 5 2.

COMMENTS

This species has a very large number of hosrs,

most of them belong to the agamie form.

Neverthelcss, it seèms not to be a specifîcity in

some gall models. Moreovei, this species shows a

major morphological variability. Gicaud descri-

bed S. longiventris according to gasrral length.

Certainly, Giraud was right, but the males and

females (wichout gaster) arc dre saine as the lar¬

ges! specimens of S. gallaepornifortnis. For this

reason, we believe that gaster length is a new

variable caracter of this species. Furthermore, ail

the hosts of S. longiventris ntentioned by Giraud

hâve been cited in S. gafJaepomiformis.

Synergtis diaphanus Giraud

(in Houard, 1911:317-318)

TYPE MATERIAL — Girauds collection. Lectotype 5,

here designated, mounted on micro-pin; paralecto-

types: 2 5 5 and 7 2 2, ail of them mounted on

micro-pins.

Distribution and hosts. — Only Cynipini: in

Central Europe (counrry unknown) in Andricus gal-

laetinaoriac (Olivier, 1791) ô ( Qnercm caducifolius

host unknown; Giraud (in Houard 1911) mentioned

chat A. gallactinctoriae tvas collected in Q. petraea,

Q. fahur\ rnd Q, httmihs] .

COMMF.NTS

Section I species with the médial mesoscutal

impression are easy to differentiate. Synergus dia-

pbanus is closely related to S. ibericus because it

has the same motphology and similar eCology;

S. ibericus is a species inquiline of A. ko tb ri 6 and

S. diaphonies to A. ga llaei in cto riae, both host spe¬

cies make spherical galls (10-20 mm) with super-

ftcial protubérances (sometimes absent in

A, kollan, ) but the galls of A. gallactinctoriae are

harder, Probablv they are sister species. In order

to distinguish these species the following charac-

ters may be used:

1. Radial cell 3.0 cimes longer than wide; R2 vein straight line. Antenna light, orange;

last flagellomere segments almost 2X1. Inquiline in galls of Andricus kollari (in

Iberian Peninsula). S. ibericus Tavares, 1920

— Radial cell shorter, 2.5 to 2.7 tîntes longer than wide; R2 vein curved. Antenna

colour dark, brown; last flagellomere segments shorter, almost 1.5 times longer than

wide. Inquiline in galls of Andricus gallactinctoriae (not présent in Iberian

Peninsula). S. diaphanus Giraud (in Houard 1911)

Synergus consobrinus Giraud

(/«Houard 1911)

Type MATERIAL. — Giraud’s collection. Lectotype 5,

here designated, mounted on micro-pin; paralecto-

types: 1 3 and 9 2 2 , ail of them mounted on micro-

pins.

Distribution and hosts. — Only Cynipini: in

Central Europe (country unknown) in Andricus gros-

sulariae Giraud, 1859 probably in Q. cerris according

to Giraud, 1859.

Redescription

Male

Length 1.5-2.1 mm. Head with a yellow colora¬

tion more or less expanded; vertex black and

536

ZOOSYSTEMA • 1998 • 20(3)

Giraud’s types species of Synergus

Fig. 4. — Synergus consobrinus Giraud, dorsal view. A, head; B, thorax. FC, facial carinae. Scale bars: A, 200 pm; B, 500 pm.

ZOOSYSTEMA • 1998 • 20(3)

537

Pujade-Villar J. & Ros-Farré P.

Fig. 5. — A. head and first female antennomeres of S. consobrinus Giraud; B, first male antennomeres of S. consobrinus Giraud.

A, antennomeres. Scale bar: 200 pm.

cheeks from yellow to red; thorax black; gaster

red; legs light between yellow and brown; wing

venation light brown. Head, in dorsal view,

about two tintes broader titan long; in frontal

view, triangular and slightiy broader than long;

face radiating striate witl strias to thc toruli;

front coriarious without punctures in small spe-

cintens and with punctures in large specimens;

latéral frontal carinae strong and not branched

near ocelli (Fig. 4); vertex coriarious-rugose not

punctuated or with scattered shallow punctures

without subparallel carinae running transvetsely

between posteriot ocelli; the diameter of the laté¬

ral ocelli is 2 and POL; OOL: OCÜ relation is

6: 3: 3; antenna with fifteen antennomeres (4: 2:

7: 4: 4: 4: 4; 4: 3.5: 3: 3: 3: 3: 3: 3); A2 is longer

than broad; A3 is strongly expanded apically

(Fig. 5B); rhose from A7 to Al 5 are thicker.

Pronotum with coriarious sculpture and some

longitudinal striae laterally; latéral pronotal cari¬

nae présent; mesoscutum with coriarious sculp¬

ture; notauli (Fig. 4B) percurrent, not very deep

and posteriorly not very btoad; médial mesoscu-

tal impression short; scutelar foveae ovate and

sculptured with blurred posterior margins; meso-

pleuron longitudinally striate; parallel propodeal

carinae, pubescent area between them. Radial

cell of forewiltg closed, length from 2.5 to

2.7 tintes breadth. Gaster without punctuation,

one smailer than thorax; abdominal terga 3 + 4

without pictures posteriorly or limited to an

apical dorsal patch. Tarsal claws with a tooth.

Female

Diflfers from male as follows; length 1.5-2.5 mm;

antenna with fourteen segments (5: 3: 7: 6; 6: 6:

5: 5: 4: 4: 3: 3: 3: 5); A2 longer than broad (Fig.

5A); A3 not raodified. Face black and yellowish

around mouth.

After examining the rypical sériés of tins species,

we concluded that it is related closely to the

S. pallicornis, S. albipes and S. nervosus group.

Moreover, S. consobrinus is a cynipid-hosr in galls

of Andricus grossulariae. Not one species of this

species group has this host. Andricus grossulariae

fiorms galls in catkins on Q. terris (in Europe

except the Iberian Peninsula) or on Q. su ber.

We summariy.e the morphological différences of

this group of species in the following dichotomi-

cal key to identification:

538

ZOOSYSTEMA • 1998 • 20(3)

Giraud’s types species of Synergus

1. Latéral frontal earinae strong and often much branched near ocelli; subparallel cari-

nae ninning transversely between posterior ocelli, and obliquely From each posterior

ocellus to margjn oFocciput. First flagellomere long in both sexes; in females at least

1.5 times longer than second; in males curved in the middle and weakly expanded

distally. Face alvvays black.5. pallicomis Hartig 1841

— Latéral frontal earinae weak (where strong die face is not black) and branched or

not near ocelli; vertex coriarious, usually without subparallel earinae running trans¬

versely between posterior ocelli. First flagellomere relatively shorter; in females less

than 1.5 times longer than second; in males slightly expanded apically or curved in

the middle and not expanded (where it is weakly expanded distally the face is yellow

or yellowish red)........2

2. Latéral frontal earinae strong and complété. Males with First flagellomere weakly

expanded distally. Face in males yellowish color and red in females. Inquilines in

galls of Andricusgmsulariae in Q. cerris .... S. consobrinus Giraud (in Houard, 1911)

— Latéral frontal earinae weak. Males with the first flagellomere segment different and

face black. Females with the face black. Inquilines in a large number of Cynipid

galls in Quercus caducifolious ..

. S. albipes Hartig, 1841 and S. nervosus Harcig, 1840

TAXONOMICAL SUMMARY AFTER THE

GIRAUD’S TATES REVIEW OF Synergus

Synergini

Saphonecrus Dalla Torre et Kieffer, 1910

S. baimi (Mayr, 1872)

S. clavatus Giraud (in Houard 1911) n.syn.

S. haymi (Mayr); Pujade-Villar et Nieves-

Aldrey, 1990: (misspelling), n.syn.

S. ondulants (Mayr, 1872)

5. apertus Giraud (in Houard 1911)

S. gimudi Pu jade-Villar, 1985

Synergus Hartig, 1840

S. albipes Hartig, 1841

S. hartigi Giraud (in Houard 1911) n.syn.

S. consobrinus Giraud (/'«Houard 1911)

5. diaphanus Giraud (in Houard 1911)

S. gallaepomiforrnis ( Fonscolombe, 1832)

S. longiventris Giraud (in Houard 1911) n.syn.

S. subterraneus Giraud (/'«Houard, 1911)

S. latifrons Nieves-Aldrey et Martin-Chicote,

1985 n.syn.

S. tbaumacerus (Dalman, 1823)

S. tnflatus Giraud (/«Houard 1911) n.syn.

5. vesiculasus Giraud (in Houard 1911) n.syn.

S. variabilis Mayr, 1872

S. cerridis Giraud (in Houard 1911) n.syn.

S. confirmis Giraud (in Houard 1911) n.syn.

5. cerricolus Vassileva-Samnalieva, 1986 n.syn.

Acknowledgements

We are very grateful to the late Dr. S. Kelner-

Pillaut from the Muséum national d’Histoire

naturelle of Paris who kindly loaned ail Giraud’s

types of Synergus on which the présent study is

based. We also thank Dr. Casevitz-Weulersse

from the Muséum national d’Histoire naturelle

of Paris for her patience. I am grateful to Dr

R. R. Askew for pointing out the spelling mista-

ke of Saphonecrus haimi.

ZOOSYSTEMA • 1998 • 20(3)

539

Pujade-Villar J. & Ros-Farré P.

REFERENCES

Darboux G. &c Houard H. 1907. — Galles de

Cynipides. Recueil de figures originales exécutées

sous la direction de feu le Dr Jules Giraud.

Nouvelles Archives du Muséum national dHistoire

naturelle de Paris 9 (4) : 173-263.

Giraud J. 1859. — Signalements de quelques espèces

nouvelles de Cynipides et de leurs galles. Verhand-

lungen der zoolocisch-botanischen Gesellscbaft in

Wien 9: 337-374.

— 1866. — Comunications sur diverses galles du

chêne et sur les insectes qui les forment. Annales de

la Société entomologique de France , Paris, série 4, 6 :

197-200.

— 1868a. — Bulletin entomologique : Quelques

galles d’Hyménoptères. Annales de la Société ento¬

mologique de France, Paris, série 4, 8 : 52-55.

— 1868b. — Notes qui suivent relativement à divers

Cynips. Annales de la Société entomologique de

France , Paris, série 4, volume 8 : 109-112.

— 1871. Miscellanées Hyménoptérologiques III : des¬

cription d’Hyménoptères nouveaux avec l’indica¬

tion des mœurs ae la plupart d’entre eux et

remarques sur quelques espèces déjà connues.

Bulletin de la Société entomologique de France,

série 5, volume 1 : 389-419.

Houard C. 1911. — Les Cynipides et leurs galles

d’après le cahier de notes du Docteur Jules Giraud.

Nouvelles Archives du Muséum national d'Histoire

naturelle de Paris h. III : 199-341.

Kieffer J. J. 1902. — Les Cynipides, volume I, in

André E. (ed.). Espèces des Hyménoptères d'Europe et

dAlgérie. Paris, volume 7, 685 p.

Laboulbcnc A. 1877. — Liste des éclosions d’insectes

observées par le Dr Joseph-Étienne Giraud,

membre honoraire. Annales de la Société entomolo¬

gique de France, Paris 7 : 397-436.

Nieves-Aldrey J. L. & Pujade-Villar J. 1986. — Sobre

las especies ibéricas de la secciôn II (Mayr, 1872)

del género Synergus Hartig, con description de una

especie nueva (Hymenoptera, Cynipidae,

Cynipinae). Eos, 62: 137-165.

Pujade-Villar J. & Nieves-Aldrey J. L. 1990. —

Révision de las especies europeas del género

Sapbonecrus Dalla Torre et Kieffer, 1910 (Llymeno-

ptera, Cynipidae, Cynipinae). Bulleti de la Institucio

Catalana de Histdroa Natural ( ser. Zool.) 8: 45-55.

Quinlan J. 1978. — Xystus testaceus Hartig, 1841 and

Synergus variabilis Mayr, 1873 redefined and redes-

cribed (Hymenoptera: Cynipoidca). Zeitschrift der

Arbeitseemeinschaft Osterreichischer Entomoloven 30:

(1/2): 71-74.

Submitted on 17 November 1997;

accepted on 2 March 1998.

540

ZOOSYSTEMA • 1998 • 20(3)

Instructions aux auteurs

La ligne éditoriale

Zoosystema est une revue consacrée à l'inventaire,

1 analyse et l'interprétation de la biodiversité anima¬

le. F.llc publie des résultats originaux de recherches

en zoologie, particulièrement en systématique et

domaines associés : morphologie comparative,

fonctionnelle et évolutive ; phylogénie ; biogéogra¬

phie ; taxinomie et nomenclature...

Un numéro de Zoosystema pourra être consacré à un

thème particulier sous la responsabilité d'un éditeur

invité.

Les auteurs devront suivre le Code International de

Nomenclature Zoologique. Il est recommandé que le

matériel-type soit, au moins en partie, déposé dans

les collections du MNFIN.

Les manuscrits, dont le nombre de pages n’est pas

limité a priori devront suivre rigoureusement les

recommandations aux auteurs (voir ci-dessous), et

seront adressés à la revue:

Service des Publications Scientifiques du Muséum

Zoosystema

57 rue Cuvier

F-75231 Paris cedex 05

Tel : (33) 01 40 79 34 38

Fax : (33) 01 40 79 38 58

e.mail : bulletin@mnhn.fr

Tout manuscrit non conforme à ces instructions

sera retourné pour mise au point. Chaque manus¬

crit est évalué par deux rapporteurs, ou plus.

Instructions aux auteurs

Chaque manuscrit soumis (y compris les illustra¬

tions) doit être présenté en trois exemplaires au for¬

mat A4, avec un double interligne et des marges

d’au moins 3 cm ; chaque page sera numérotée. Les

illustrations originales seront jointes au manuscrit

définitif, ainsi qu’une disquette 3.5" de format

Apple Macintosh ou compatible IBM (traitement

de texte Word de préférence), qui devra contenir

également les tableaux et éventuellement les illus¬

trations (Adobe lllustrator, Photoshop ; Deneba

Canvas).

Le format

Les manuscrits, écrits en français ou en anglais, doi¬

vent être structurés comme suit :

- titre, si possible bref, en français et traduit en

anglais ; un titre coûtant doit être proposé ;

- nom (s) et prénom(s) de(s) auteur(s) suivis de

leur(s) adresse(s) professionnelle(s), en indiquant si

possible le numéro de télécopie et l’adresse électro¬

nique ;

- résumés écrits en français et en anglais (800 signes

au maximum chacun), suivis des mots clés et « key

words » ;

- dans le texte courant, utiliser les italiques pour

tous les noms en latin : taxons de rangs générique

et spécifique (ex : Pseudolaureula Kwon, Ferrara et

Taiti, 1992), étal., eg. ... -,

- dans le texte courant, les références aux auteurs

seront en minuscules, ex. Dupont (2001), Dupont

(2001, 2002), (Dupont 2001 ; Durand 2002),

(Dupont & Durand 2003, 2005), Dupont (2001 :

1), Dupont (2001, fig. 2) ;

- dans le texte courant, les références aux illustra¬

tions et aux tableaux de l’article seront présentées

ainsi : (Fig. 1), (Fig. 2A, D), (Fig. 2A-C), (Figs 3,

6), (Figs 3-5) ; (Tableau 1) ;

- les remerciements seront placés à la fin du texte,

avant les références bibliographiques ;

- les références bibliographiques doivent suivre les

exemples donnés ci-dessous ;

- indiquer dans la marge l’emplacement des illustra¬

tions ;

- donner les légendes des figures sur une feuille

séparée.

Les illustrations

Les illustrations au trait doivent être réalisées à

l’encre de Chine ou être fournies en impression

laser, Les photographies, bien contrastées, seront

sur fond noir ou blanc. Elles pourront être regrou¬

pées, et dans ce cas, identifiées par une lettre en

capitales (A, B, C...). Les planches photogra¬

phiques, de préférence placées dans le corps de

l’article et non regroupées à la fin de celui-ci, doi¬

vent être traitées et numérotées comme des figures.

ZOOSYSTEMA • 1998 • 20(3)

541

Instructions aux auteurs

Les illustrations pourront être assemblées sur une

colonne (70 X 190 mm) ou sur toute la largeur de la

justification (144 X 190 mm). Si possible, les

légendes (et lettrages) ne devraient pas figurer sur les

originaux. Ils seront disposés, alors, sur un calque

joint à chaque figure, la rédaction se chargeant de

les placer. Chaque figure doit comporter une échelle

métrique, sans aucun coefficient multiplicateur. Les

tableaux et graphiques, à inclure dans le manuscrit,

doivent pouvoir erre imprimés sur une page er resrer

lisibles après réduction éventuelle. Des photogra¬

phies en couleur pourront être publiées moyennant

une participation financière de ou des auteurs.

Références bibliographiques

Hoeg J. T. & Lüçzen J. 1985. — Comparative

morphology and phylogeny of che Family

Thompsoniidae (Cirripedia: Rhizocephala:

Akentrogonida) with description of three new

généra and seven new species. Zoologiea Scripta

22: 363-386.

Rôckel D., Korn W. & Kohn A. J, 1995. —

Manual of the living Conidae, volume 1: Indo-

Pacific région. Chrïsta Hemmen, Wiesbaden,

517 p.

Schwaner T. D 1985. — Population structure of

black tiger snakes, Notechis ater niger, on offsho¬

re islands of South Australîa: 35-46, in Grîgg G.,

Shine R. & F.hmann H. (eds), Bio/ogy of

Australasian Frogs and Reptiles. Surrey Beatty and

Sons, Sydney.

Épreuves et tirés à part

Les épreuves seront adressées à fauteur ou au pre¬

mier auteur (sauf indication contraire) et devront

être retournées corrigées sous huitaine. Les correc¬

tions, autres que celles imputables à la rédaction ou à

l’imprimeur, seront à la charge des auteurs. Le(s)

auteur(s) recevront gratuitement vingt-cinq tirés à

part pour les articles jusqu'à cinquante pages (au-

delà, consulter la rédaction) ; les tirés à part supplé¬

mentaires seronr à commander sur un formulaire

joint aux épreuves.

Soumettre un article pour publication dans

Zoosystema implique que celui-ci n’ait pas été sou¬

mis dans une aurre revue. Les droits de reproduc¬

tion de l'article, y compris les illustrations, sont

réservés à la revue. La reproduction de tout ou par¬

tie de l'article doit faire l'objet d'une demande écrire

préalable, adressée à la rédaction.

Scope ofthe Journal

Zoosystema is a journal devored to the inventory,

analysis and interprétation of animal biodiversity. It

publishes original résulta of zoological research,

particularly in systematics and related ftelds: com¬

parative, functional and evolutionary morphology,

phylogeny, biogeography, taxonomy and nomen¬

clature..

A complété issue of Zoosystema may be devoted to

several papers on a single copie under the responsa-

bility of an invited editor.

Papers should follow the International Code of

Zoological Nomenclature. We reeommend that the

authors should deposit in the MNHN collections,

ar least a part of the type material.

Manuscripts, without limitation of the number of

pages, must conform srrictly with the instructions

to authors and will be sent to the Editor:

Service des Publications Scientifiques du Muséum

Zoosystema

57 rue Cuvier

F-75231 Paris cedex 05

Tel : (33) 01 40 79 34 38

Fax: (33) 01 40 79 38 58

e.mail: bullerin@mnhn.fr

Instructions to authors

Manuscripts must be submitted in triplicate in A4

format, double spaced, with margins of at least

3 cm and ail pages numbered. The original figures

should be senr wirh the revised manuscript, as well

as a 3.5” diskette Apple Macintosh or lBM-compa-

rible (Word, Word Perfect... ) format, which will

also contain tables and possibly figures (Adobe

Illustrator, Photoshop ; Deneba Canvas).

Format

Papers are ro be written in simple and concise

French or English. They should be organized as fol-

lows:

- a brief tirle in F.nglish and French;

- a suggested running head;

- name(s) of author(s), followed by their full profes-

sionaJ address(es) and, if possible, Fax number and

e-mail;

- abstracts (in English and French) not exceeding

800 signs' eue h, with key words and "mots dés";

- text with italicizcd words for Latin: taxa of generic

and spécifie ranks (e.g. Pseudolaureola Kwon,

Ferrara erTaiti, 1992), et al., ...;

542

ZOOSYSTEMA • 1998 • 20(3)

Instructions aux auteurs

- references to authors in main text should be pré¬

sentée! as follows: Smith (2001), Smith (2001,

2002), (Smith 2001), (Smith 2001; Cary 2002),

(Smith & Cary 2003, 2005), Smith (2001: 1),

Smith (2001, fig. 2);

- references to illustrations and tables should be

indicated as follows: (Fig. 1), (Fig. 2A, D), (Fig.

2A-C), (Figs 3, 6), (Figs 3-5); (Table 1);

- keep acknowledgements short and place them at

the end of the text;

- give captions to illustrations on a separate sheet;

Illustrations

The éditorial board will pay spécial attention to the

quality and relevance of illustration.

Line drawings must be in Indian ink or high quality

laser printouts; high contrast photographs, placed

on white or black backgrounds, are required. These

can be grouped into Figures and their éléments and

identified by letters A, B, C... Plates are not placed

at the end of the article: they will be considered as

figures and numbered as such. Arrange Figures to fit

one or two columns (70 X 190 mm or 144 X

190 mm). No diagram or table is to exceed one

page. Letters, numbers, etc., for each figure, are to

be indicated on an accompanying overlay, not on

the original figure. A scale bar is needed for each

figure (without magnification factor).

References

Hoeg J. T. & Lützen J. 1985. — Comparative

morphology and phylogeny of the family

Thompsoniidae (Cirripedia: Rhizocephala:

Akentrogonida) with description of three new

généra and seven new species. Zoologica Scripta

22: 363-386.

Rôckel D., Korn W. & Kohn A. J. 1995. —

Manual of the living Conidae, volume 1: Indo-

Paciftc région. Christa Flemmen, Wiesbaden,

517 p.

Schwaner T. D. 1985. — Population structure of

black tiger snakes, Notechis ater niger, on offsho¬

re islands of South Australia: 35-46, in Grigg G.,

Shine R. & Ehmann H. (eds), Biology of

Australasian Frogs and Reptiles. Surrey Beatty and

Sons, Sydney.

Proofs and reprints

Proofs will be sent to the first author for correction

and must be returned within eight days by express

mail. Author(s) will receive twenty-five offprints

free of charge (for paper up to 50 pages; for paper

exceeding 50 pages, consult the editors); further

offprints can be ordered on a form supplied with

the proofs.

The submission of a manuscript to Zoosystema

implies that the paper is not being offered for

publication elsewhere. Copyright of published

paper, including illustrations, becomes the property

of the journal. Requcsts to reproduce material from

Zoosystema should be addressed to the editor.

ZOOSYSTEMA • 1998 • 20(3)

543

Mise en page

Noémie de la Selle

Packaging Editorial

Achevé d’imprimer

sur les presses de l’Imprimerie Durand

28600 Luisant (France)

Octobre 1998

Dépôt légal n° 10147

Printed on acid-free paper

Imprimé sur papier non acide

Date de distribution du fascicule 2 :

13 juillet 1998

Couverture : Branchinecta minuta Smirnov, 1948 (Crustacea, Anostraca),

détail de l’enveloppe de l’œuf (MEB)

Photographie A. Thiéry (Université d’Avignon)

1998 • 20 ( 3 )

zoosystema

429 •

Monniot F.

An enigmatic filiform organism in the epicardium cavity of a new polycitorid ascidian

■r i

Litvinova N. M.

439 • Two new species of the genus Ophiuraster (Ophiurinae, Ophiuroidea, Echinodermata) from French

collections and some remarks on the genus g£ w mh

Durette-Desset M.-C. & Corvione M.

445 • Une nouvelle espèce de Molineus (Nematoda, Trichostrongylina, Molineoidea), parasite d'un Primate

sud-américain

Sawyer R. T., Hechtel F. O. P., Hagy J. W. & Scacheri E.

451 • A study in medical history: introduction of médicinal leeches into the West Indies in the nineteenth

century

Vermeij G. J. & Bouchet P.

471 • New Pisaniinae (Mollusca, Gastropoda, Buccinidae) from New Caledonia, with remarks on Cantharus

and related généra '"■■yçfm

Lourenço W. R.

487 • a new species of Tityus C. L. Koch, 1836 (Scorpiones, Buthidae) in Colombia, with a check list and

key to the Colombian species of the genus

Maréchal P. & Marty C.

499 • Réhabilitation du genre Harmonicon (Pickard-Cambridge, 1896) et description d'une nouvelle espèce

de Guyane française (Araneae, Mygalomorphae, Dipluridae)

505 •

Carriol R.-P.

A new pedunculate cirripede (Thoracica, Heteralepas ) from the Northeast Atlantic Océan

A. '

511

Hayashi K.-l. & Yaldwin J. C.

A new species of the genus Pasiphaea from the South Indian Océan (Crustacea, Decapoda,

Pasiphaeidae) r

* inÊÈtk. «k

Barra J.-A.

521 • Onychiuridae (Insecta, Collembola) d'Herzégovine en région karstique avec description d'une

nouvelle espèce ’-tÜ

. J—, înii

529 ^ Pujade-Villar J. 6r Ros-Farré P.

Review of Giraud's types of the species of Synergus Hartig, 1840 (Hymenoptera, Cynipidae)

Conception Graphique : Isabel Gautray

ISSN : 1280-9551

Vente en France

Sales Office (France exduded)

Muséum national d'Histoire naturelle

Universal Book Services

Diffusion Delphine Henry

Dr. W. Backhuys

57, rue Cuvier, 75005 Paris,

P.O. Box 321 2300 AH Leiden

France

The Netherlands

Tel. : 33 - 01 40 79 37 00

Tel. : 31 -71 517 02 08

Fax : 33 - 01 40 79 38 40

Fax : 31 -71-517 18 56

e.mail : dhenry@mnhn.fr

e.mail : backhuys@euronet.nl