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Two new généra and species of sponges

(Porifera, Demospongiae) without skeleton

from a Mediterranean cave

Jean VACELET & Thierry PEREZ

Centre d’Océanologie de Marseille (CNRS-Université de la Méditerranée, UMR 6540 DIMAR),

Station Marine d’Endoume, F-13007 Marseille (France)

Vacelet J. & Perez T. 1998. — Two new généra and species of sponges (Porifera,

Demospongiae) without skeleton from a Mediterranean cave. Zoosystema 20 (1) : 5-22.

KEYWORDS

Porifera,

taxonomy,

Mediterranean,

new généra and species,

cave.

ABSTRACT

Two new généra and species of Demospongiae are described from a north-

western Mediterranean littoral cave characterized by cold homothermy,

which shelters deep-sea invertebrates. The two new sponges hâve neither

minerai nor fibrous skeleton. Their cytology is described using transmission

électron microscopy. Thymosiopsis cuticuldtus n.g. n.sp. (Chondrillidae)

shares sonie characters with Thymosia Topsent. but lacks the diagnostic

spongin fibres. Myceliospongta araneosa n.g, n.sp. has unusual anatomy, cyto¬

logy, and mode of growth. No clear relationship with any order of the

Demospongiae is indicated and the sponge is classified as incertae sedis within

the Demospongiae.

MOTS CLÉS

Porifera,

taxonomie,

Méditerranée,

nouveaux genres,

nouvelles espèces,

grotte.

RÉSUMÉ

Deux nouveaux genres et espèces d'éponges (Porifera, Demospongiae) sans sque¬

lette d’une grotte méditerranéenne, Deux nouveaux genres et espèces de

Demospongiae sont décrits d’une grotte littorale de la Méditerranée nord-

occidentale, qui est caractérisée par une homothermie froide er abrite des

invertébrés de mers profondes. Les deux nouvelles éponges sont dépourvues

de squelette minéral ou fibreux. Leur cytologie est décrite en microscopie

électronique. Thymosiopsis cuticulatus n.g. n.sp. (Chondrillidae) a des affinités

avec Thymosia Topsent, mais ne possède pas les fibres de spongînc distinc¬

tives. Myceliospongia araneosa n.g. n.sp. est très inhabituelle par son anato¬

mie, sa cytologie et son mode de croissance. Aucune relation ne peut être

établie avec un ordre de Demospongiae et 1 éponge est classée comme incer¬

tae sedis dans les Demospongiae.

ZOOSYSTEMA • 1998 • 20(1)

Vacelet J. & Perez T.

INTRODUCTION

Sponges without skeleton, whose taxonomy is

especially difficult in the absence of the conven-

tional diagnostic characters, display a remarkable

abundance and variety in Mediterranean subma-

rine caves. Représentatives of ali the described

généra: Oscarella Vosmaer, 1884 and

Pseudocorticium Boury-Esnault et al., 1995

(Homoscleromorpha, Plakinidae), Chondrosia

Nardo. 1847 (Tetractinomorpha, Chondrillidae),

Hexadella Topsent, 1896 (Ceractinomorpha,

Darwinellidae), and Halisarca Dujardin, 1838

(Ceractinomorpha, Halisarcidae) are présent in

Mediterranean caves, either in semi-obscure

zones near the cave entrance or in darkesr

recesses. Recent studies hâve shown that at least

four different species of Oscarella could be pré¬

sent in a single cave (Muricy et al. 1996), two of

them known exclusively from dark caves. The

recently described genus Pseudocorticium is as yet

known only from caves (Boury-Esnault et al.

1995). This abundance of sponges without skele¬

ton in caves probably reflects the situation on

vertical and overhanging surfaces of steep cliffs in

bathyal environments, with which caves bave

faunistic similarities (Harmelin et al. 1985;

Vacelet et al. 1994). The spccies which are

known only from caves most likely also live on

the continental slope, especially in steep canyons

where they are as yet unrecorded due to the

obvious difficulries in observation and sampling.

These sponges usually hâve close relatives which

hâve a skeleton. so their taxonomie position is

seldom dispured. An exception, however, is rhe

genus Halisarca, with no known skeletonized

relatives and an unusual anatomy. After having

been classified lor a long rime either as incertae

sedis or in the order Dendroceratida, rhe genus

has finally been isolared in the new order

Halisarcida (Bergquist 1996).

We describe here, rwo new sponges without ske¬

leton from the dark zones of a cave which shel-

ters an unusually high number of deep-sea

invertebrates, due ro a stable homothermie régi¬

me around 13 to 14.5 U C, similar ro that of the

deep Mediterranean (Vacelet et al. 1994). The

new sponges, which are unrecorded in caves with

température variations similar to those of the lit¬

toral zone, are probably représentatives of the

undescribed sessile fauna of the deep-

Mediterranean canyons. They belong to two new

généra, one of which has uncertain affinities.

Their description includes data on ultrastructural

cytology, which are particularly important in the

absence of the conventional taxonomie charac-

ters of the skeleton.

MATERIALS AND METHODS

FlELD OBSERVATIONS AND SAMI’l ING

The specimens were observed in situ in the

"3PP" cave near La Ciotat (43°09.47’N -

05 t, 36.01 ’E) with an underwater magnilying lens

(Mladenov & Powcll 1986) and photographed

with a close-up lens. Pièces of specimens were

collected, either by scraping sponges from the

substratum or by detaching fragments of the cave

walls. Specimens, cxcept those used for detailed

microscopy (see below) were ftxed in formalin

and stored in alcohol.

A general description of the cave is given in

Vacelet et al. (1994) and Vacelet (1996).

Température recotdings were made over two

years using Deep-Sea Sealoggers thermographs

(Vacelet 1996; Harrnelin 1997).

Cytology

For light and transmission électron microscopy

(TEM), the specimens were fixed in situ in gluta-

raldehyde 2.5% in a mixture of 0.4 M cacodylate

buffer and sea water (4 vol.: 5 vol.). They were

maintained in the frxative lor 24 hours and post-

fixed 2 hours in 2% osmium tetroxide in sea

water. Specimens were decalcified in 10% RDO

(Du Page Kinetic Lab) in sea water in order to

remove the underlying substratum, dehydrated

through an alcohol sériés and embedded in

Araldite. Semi-thin sections were stained with

toluidine blue. Th in sections, contrasted with

uranyl acetate and lead citrate, were observed

under a Zeiss EM 912 transmission électron

microscope.

ZOOSYSTEMA • 1998 • 20 (1)

New sponges without skeleton

SYSTEMATICS

Order CHONDROS1DA

Boury-Esnault et Lopes, 1985

Family CHONDRILLIDAE Gray, 1872

Thymosiopsis n.g.

Type SPECIES. — Thymosiopsis cuticulatus n.sp.

Etymology. — The generic name is derived Irom

Thymosia, a genus of the same family and the suffis

-ops (in Greek: with the aspect of).

Diagnosis

Encrusting Chondrillidae. General organization

similar to thar of the genus Thymosia, having a

smooth surface, a superficial cuticle and

pore-sieves, a marked cortex enriched with fibril-

lar collagen, but lacking spongin fibres.

Thymosiopsis cuticulatus n.sp.

(Figs 1-3)

Type MATERIAL. — North-Western Mediterranean.

La Ciotat, 3PP cave. — 50 m front cave opening,

16 m in depth, 7.VU.1996: holotype, 1 fragment

(MNHN D JV 59). — 30 m from cave opening,

20 m in depth, 3.III.1997: paratype, 2 fragments

(MNHN D JV 60).

ETYMOLOGY. — The species name refers to the pré¬

sence of a cuticle (from cuticula, Latin, thin skin).

Locality AND HABITAT. — Known only from 3PP

cave, 1.2 km south-west of La Ciotat on the French

Mediterranean coast (43°09.47’N - 05°36.01’E). The

sponge has been found on vertical or overhanging

walls, 16 to 22 m deep, 30 to 80 m from the cave

opening, in a trapped body of water whose tempéra¬

ture varies from 13 to 14.5 °C year round (Vacelet et

al. 1994). The sponge is not very common in the cave

and only a few large specimens hâve been observed.

Description

Shape and size

The sponge is encrusting, up to 15/20 cm, and

3 to 5 mm rhick in the centre, thinner on the

edges which are irregular (Fig. IA). The sponge

is firmly aftached to the substratum by its whole

undersurface, and insinuâtes into small cavities

such as empty serpulid tubes.

Colour

The in situ colour is white or yellowish white. A

faint brown tinge due to a thin, inconstant depo-

sit of iron and manganèse oxides on the cuticle,

is visible in places, especially on ridges correspon-

ding to the irregularities of the substratum.

Thinner parts of the sponge, mostly on the

edges, are greyish due to the black underlying

rock seen through the transparent tissue. After

fixation in formalin, the alcohol-preserved speci¬

mens are whitish.

Surface

The surface is smooth, but irregular as the spon¬

ge closely follows the irregularities of the substra¬

tum. Small aperrures, approximately 0.6 mm in

diameter (measurements from underwater

close-up photographs of non-contracted speci¬

mens) are gathered in oval or circular dépréssions

with an elevated ourline, 5 to 15 mm in diameter

(Fig. IA). Most of the dépréssions are probably

inhalant pore-sieves. Superficial canals, visible

below the ectosome especially on the thin zones

at the periphery of the sponge, converge towards

the larger of these dépréssions, which may be

composite oscules. Single oscilles hâve not been

observed, neither in situ nor on collected speci¬

mens. The holotype, which was living relatively

near the cave entrance (30 m), was covered by

various encrusting sponges, bryozoans and

didemnids, which were loosely attached on the

surface between the pore-sieves. Most of the spe¬

cimens, however, were free of macroepibionts.

Texture

The consistency is quite cartilaginous, although

easy to tear.

Skeleton

There is neither spiculé nor spongin fibre skele¬

ton. A small amount of foreign material is fre-

quently included in the choanosome.

Anatomy and. cyto/ogy

Ectosome and choanosome are clearly distinct

(Fig. IB), although the ectosome is not déta¬

chable. The ectosome is 40-50 pm deep and is

lined on the outer surface by a thin, non-cellular

cuticle, which appears, in sections observed with

ZOOSYSTEMA • 1998 • 20(1)

Vacelet J. & Perez T.

the light microscope, as a wrinkled layer, 2 pm in

thickness. TKis curicle is covered in most places by

a mucous layer metachromatically stained by

toluidine blue. In TEM (Fig. 2A), the surface

appears to bc covcred by a wavy dense layer,

0.15 pm in thickness, in direct contact with the

collagen librils of the underlying tissue, without

any pinacoderm This thin layer is covered by a

zone of very dark, irregular granules, which are

probablv deposits of iron and manganèse oxides,

frequent in caves (Harmelin et al. 1985; Bianchi

et al. 1986). The oxide deposit, visible as a brown-

ish coloration in some areas of the sponge surface,

and the presence of epibionts on some specimens

Fig. 1. — Thymosiopsis cuticulatus r\.g. n.sp,, A, specimen in situ, arrow head: pore-sieves; top right, the sponge with superficial

canals is Diplastrella bistellata ; B, semi-thin section showing the tissue organization. b, extraceilular bacteria; c, cuticle; ca, canal;

cc, choanocyte chamber; ci, cells with inclusion Types 1 and 2; sp, spherulous cells Type 3. Scale bars: A, approximately 13 mm;

B, 16 pm.

ZOOSYSTEMA • 1998 • 20(1)

New sponges without skeleton

both indicate chat the cuticle is nor transient, but

remains stable for at least several months. T’his

layer is covered by a fibrillar area, 1.5 to 2 pm in

thickness, having the appcarance of a niucous

sheath. An enipty xpace is often présent between

the wavy dark layer and the nther layers.

The ectosome contains bundles of collagen fibrils,

most of which are parallcl to the surface. The

fibrillar bundles are thin (1-3 pm), not very dense-

ly packed, and the ecrosome is eonsiderably thin-

ner and less speciali/.ed than in sponges with a truc

cortex, such as Chondmia or Tethya. The ectosome

also contains spheruious cells (Type 3, described

below), large cells with irregular, small granules

(Types 1 and 2), and exrracellular bacteria.

In the choanosome, collagen fasctcles arc less

dense. Choanocyte chambers occur in relatively

low density. Most of the choanosome volume is

occupied by Types 1 and 2 cells, closely pressed

together in places, and by a high nutnber of

extracellular symbiotic bacteria.

Exopinacocytes were not observed, They are

absent in the areas lined by the cuticle, and are

probably found only in die pore-sieves, for which

we hâve no good sections, Endopinacocytes are

not flageUated (Eig. 2B).

Choanocyte chambers (Figs 1 B, 2C-F) are eury-

pylous and spherical, 15 to 30 pm diameter. Tvvo

aspects hâve been observed. In sorne chambers

(Fig. 2C), choanotyres are eylindrical or pyrami¬

dal, 3 to 4 pm in size with a nucléus approxima-

tely 2 pm in diameter, spherical or pyriform,

rarely nucleolated, and a cyroplasm containing

fc-w phagosomes. In other chambers (Fig. 2E, F),

choanocytes are very irregular, and their cyto-

plasrn couvains a larger number of phagosomes.

In both cases, the collât is 3-6 pm in diameter

and inade up of rhirty-two ro tliirty-fdur micro-

villi (Fig. 2D). There is no periflagcllar sleeve.

The flagellum possesses rwo thin latéral exten¬

sions, which are faintly visible on sorne transverse

sections and which may be interprered as poorly

preserved flagellât vanes (Aiâelius 1961; Mehl &

Reiswig 1991 ) Conventional apopylar (or cône)

cells (Boury-Esnaulr étal. 1984; Langenbruch et

al. 1985; De Vos et al. 1990) and central cells

(Reiswig & Brown 1977) werc not observed.

The choanocyte base is fiat or displays a few

short pseudopodia anchoring the cell in the

underlying mesohyl, usually without a pinaco-

cyte lining. However, in certain chambers with

irregularly-shaped choanocytes, very dense fusi-

form cells line the choanocyte base and extend

long, thin lamellipodia which insinuate between

the choanocyte bodies and nnnifv into the chant-

ber çavity. Lamellipodia are in close contact with

the choanocyte collar or flagellum (Fig. 2F).

These cells, which are reniarkable in the density

o( their cytoplasm, seem to be pinacocytes lining

the choanocyte hase, as observed in sorne demo-

spouges, but in this case possibly haviug a rôle in

the régulation of the water flow by extending

pseudopodia into the chamber lumen.

Fout distinct types of cells with inclusions are

présent:

Type 1. (Fig. 3 A) Large cells, 15 to 18 pm in

diameter, containing numerous granules enclo-

sed in clear vesicles delineated by a thin sheet of

cyroplasm. I he cells hâve a degeneraiing aspect,

with nucléus and organelles rarely observed. The

granules, 0.5 to 1.1 pm in diameter, hâve a lluely

granuiar content and an irregular outlinc, with a

cloudy aspect. They appear moderately dense to

the électrons in TEM and axe stained by tolui-

dine hlue in semi-thin sections. Type 1 cells are

ahundanr by places in the choanosome and occur

as dense clusters in the ectosome.

Type 2. (Fig. 3B) Large cells, approximately the

saine size as Type 1, with similar clear vesicles

containing a dispersed fibrillar material and small

rod-like inclusions, 0.1-0.2/0.45-1.1 pm, with a

clear central area surrounded by an irregular

dense zone. These rod-îike inclusions are proba¬

bly bacteria, morphologically different from the

exrracellular ones described below. These cells,

wbose cycoplasmic outline is rarely observed,

appear to be advaneed stages in degetieration of

Type 1 cells.

Type 3. (Fig. 3C) Spheruious cells, approximate¬

ly 10 pm in diameter, with two to cen large,

homogeneous spherules, 1 to 4.5 pm in diame¬

ter, which occupy most of the cell volume.

Spherules were occasionally observed free in the

mesohyl, cspecially in the ectosome, after dege-

neration of the cell. However, these cells are not

clearly secreting the intercellular matrix, as they

do in sorne Halisarca or in Chondrosia (Vacelet &

Donadey 1987).

ZOOSYSTEMA ■ 1998 • 20 (1)

Vacelet J. & Perez T.

ZOOSYSTEMA • 1998

mm.. . ♦ * ■

■jp, ■ • j

r - 4'

- ' j NBJ

n7 tM

m é jfl

New sponges without skeleton

Type 4. (Fig. 3D) Microglanulàt cells, most

often elongated (2.5-3 pm/4.5-8 pm) with

dense, ovoid inclusions, 0.3/0.9 pin in rhe cyto-

plasm. These cells are not very abundanr.

The collagen fibrils are thin, approximately

15 nm in diameter (Fig. 3B). Their periodicity

is not apparent. They scem to belong to the

smooth type of collagen fibrils (Garrone 1978),

although this character is not well defined here.

They are orgamzcd in bundles, 1 to 3 pm in dia¬

meter, especially in the ectosome. There is no

disjunction betwecn the collagen bundles and a

granulo-fibrillar matrix such as is observed in

some species of Halisarca or of Chondrillidae

(Vacelet & Donadey 1987).

Symbiotic extraceJlular bacteria (Figs 2A, B, 3D)

found in the mesohyl belong to scveral different

morphological types, varying in size from

0.2-0.4 pm to 0.8-2.5 pm. They display the

various morphologies found in defnosponges

having a high density mesohyl (Vacelet 1975;

Vacelet & Donadey 1977, Boury-F.snault et al.

1995), in which the cell walls are complex and

frequently display a remarkable enlargement of

the periplasm. An unusual type (Fig. 2A) Ls a

rod-like cell, 0.4 pm in diameter and at least

1.5 pm in length, with a contorted shape, dense

cytoplasm and reduced nuclear area, which is

rarely observed in other demosponges, although

it morphologies lly resembles the intra- and inter¬

cellular harteria described in the parenchymella

of Haliclvna tubifera (Woollacott 1993).

Reproduction

Not observed.

Remarks

This sponge appears to hâve aftiniries with

Thymosia Topsent, 1895, a monospecific genus

of “kératose” sponge from the North-East

Fig. 2. — T. cuticulatus n.sp.. A, transmission électron microgra-

ph (TEM) of the sponge surface; b t bactEri.i c. cuticle:

mi, minerai deposit: mu, mucous deposit; sp, Isolated spherula

ot a sptierulous cell B, TEM of the ctioanosorrie showing a

small canal. C. TEM of a choanocyte chamber D, TEM of a

choanocyte collar, note the tlayellai uanes E TEM of a choano

cyte chamber wvitli irregularly-shapecl ctioanocytos. F, TEM of a

choanocyte chamber showing an elongalo coll with dense cyto-

plasm extendtng lamelllpoçlla in ihe çhnrnlter lumen (arrows)

Scale bars; A, 1.6 pm; B, 2.5 pm; C, 2.0 pm; D, 1.0 pm;

E, 2.0 pm; F, 1.3 pm.

Atlantic. After various allocations, the genus is

presently classified in family Chondrillidae

(= Chondrosiidae), as originally proposed by

Topsent. The main différences beeween

Thymosia guernei Topsent, 1895 as recently

redescribed (Boury-Esnault & Lopès 1985;

Roscll 1988; Carballo 1994), and Thymosiopsis

cuticulatus n.sp are the absence of the verrucose

horny fibres which are highly diagnostic of

Thymosia , and the composite nature of the

oscules. The two sponges share the presence of a

cuticle on most of the surface, of pore-sieves and

of a specialized ecrosome constituting a weakly

developed cortex. The characters of the choano¬

cyte chambers, aquiferous systent and Type 3

cells with inclusions (sphcrulous cells) cannot

provide a diagnosis, but do not contradict the

supposed aflïnity berween the rwo généra.

These similarities conld be differently interpre-

ted. The unusual fibres of Thymosia guernei bave

been interpreted as a hydroid skeleton (Bergquist

1980). The new sponge could be considered

either as .specimen of Thymosia guernei without

the hydroid associate, or as a Medirerranean

Thymosia - a genus never rccorded from this sea

— which would hâve lost the spongin fibres in

low-energy environmencs such as the deep sea or

a cave. Both hypothèses appear unlikely. The

hydroid nature of rhe fibres, already ruled out hy

Topsent (1895) in the original description, has

been contradicted by ail recent records of the

sponge. \Ve hâve checked their genuine nature

on new spécimens of Thymosia guernei from

Portugal (unpublished observations). The ultra-

structure of these spécimens shows that,

although the general orgamzacion is rather simi-

lar. the two sponges differ clearly by the structure

of cells with inclusions Most significantly, cells

with inclusions Type 1 and Type 2, which are

very abundant and remarkable in Thymosiopsis

cuticulatus , are absent in Thymosia guernei. These

différences indicacc that the two sponges belong

to diffèrent species. The absence ol the characte-

ristic fibres, combined with these différences in

cytology, justifies the création of a new genus,

parallel to the présent classification in rhe

Darwinellidae where the genus Hexadella,

without skeleton, is considered distinct from

Aplysilla and Darwinella.

ZOOSYSTEMA • 1998 ■ 20(1)

Vacelet J. & Perez T.

The obvious différences in the development of 1988), and by the présence of a constant cuticle

cortex and of collagen bundles benveen ibe new and well organi/ed porc-sieves in Thymosiopsis.

sponge and the genus Chondrosia (Garrone et al. The anatomy and cytology of the new sponge are

1975) precludes its allocation to that genus. The certainly more similar to those of Thymosia than

distinction of Thymosiopsis from Chondrosia is of Chondrosia.

also justîfied by différences in the canal System The genus Thymosia is rather puzzling in the

organization (Schulze 1877; Bavestrello et al. Demospongiae, where it has been diverseiy allo-

Fig. 3. — T. cuticulatus n.sp., A, TEM of a granular œil (Type 1 cell); B, TEM of a granular cell (Type 2 cell) with intraceilular bacte-

ria (arrows). C T TEM oi a spherulous cell (Type 3 cell). D TEM of the mesobyl showing various symbiotic bacteria, a degenerating

spherulous cell (Type 3 cell, sp) and degenerating microgranular cells (Type 4 cell, mg). Scale bars: A, 1.3 pm; B, 0.8 pm;

C, 1.3 pm; D, 1.6 pm.

ZOOSYSTEMA • 1998

New sponges without skeleton

cated (Rosell 1988). It bears u nu.su al, nodulose

spongin fibres and bas cytological charactcrs dif¬

ferent from those of "kératose” sponges. Most

authors presendy classify tbe genus in family

Chondrillidae (= Chondtosiidae), because of

anatomical resemblances with the genus

Chondrosia. These resemblances ate not fully

convincing, especially as the ectosôme is less spe-

cialized than in Chondrosia or in Chondrilla, and

does not constitute a thick cortex with dense fas-

cicles of collagen. Furthermorc, Thymosia guer-

nei , which posse.sses pore-sieves wdtich are

considerably more organized than the cribriporai

chones of the Chondrillidae (Schutze 1877;

Bavestrello et al. 1988), docs not display the dis¬

tinction berween both cellular and collagen élé¬

ments and a granulo-fibrillar matrix as described

in Chondrillidae (Vacelet Se Dnnadey 1987). In

our opinion, the relationships of i'hymosia and

Thymosiopsis with the Chondrillidae remain to be

confirmed; tint il this can be clone, the généra

should remain in that family.

DEMOSPONG1AE, Order incertae sedis

Family incertae sedis

Genus Myceliospongia n.g.

Type species. — Myceliospongia araneosa n.sp.

ETYMOLOGY. — The generic name dérivés from

tnukês (in Greek: futigus) and refers to the shape of

the type species.

Diagnosis

Demospongiae incertae sedis without skeleton

and without cortex. Body encrusting, from

which arises a réticulation of thin filaments cove-

ring the substratum. The sponge has a reduced

canal system and a low nurnber of choanocyte

chambers. Exopinacocytes non flagellated, cove-

red by a mucous sheet. Most of the cçlls contain

granular inclusions and symbiotic bacteria.

Myceliospongia araneosa n.sp.

(Figs 4-6)

Type MATERIAE. — North-Western Mediterranean.

La Ciotat, 3PP cave 60 m Ironi cave opening, 18 m in

depth. — 7. Vil. 1 996: hnlocvpc (MNHN D fV 61). —

17. XII. 1996 : para type (MNHN D J V 62).

1 folotypc and paratype are fragmentaty specimens.

EtYMOLOCY. — The species name dérivés from ara-

neosus (in Larîn: similar to a spider web), and refers to

the shape of the margin of the sponge.

LOCAHTY AND HABITAT. — Known only from 3PP

cave, 1.2 km south-west of La Ciotat on the French

Mediterranean coasi (43"09.47'N - 05°36.01 K). The

sponge lives on vertical or overhanging walls. 18 to

21 m deep, 50 to 80 m front the cave opening, in a

trapped body of w'ater whose température varies from

13 to 14.5 ”'C year round. ll seems to be absent far-

ther in the cave, which attends up to 120 m from

opening. The sponge is not very common in the cave

and only a few large specimens hâve been observed.

However, specimens may easily be overlooked as they

are occasionally covered by other sponges.

Description

Shape and size

Sponge encrusting, approximately I mm in

maximum thickness, composcd of a “body” cove-

ring most of tbe surface of the substratum,

altbough with irregular lacunae, and of filaments

which exrend a long distance from rhe body and

are closely applied to the substratum (Fig. 4A).

The maximum size observed is 25 cm in diatne-

ter for the body, with the filaments visible in situ

extending al least 12 cm from rbe body, rhe total

diameter o( the surface eolonized by rhe sponge

thus being approximately 50 cm. The filaments

deçrease in diameter with the distance from the

body. They can he extrcmcly thin at their extre-

mities' filaments 5 pm in diameter extending to

20 pm in lengrli hâve hecn observed. They divide

diehotomopsly or anastomose, to form an irregu-

lar réticulation. The body ol some specimens is

entirely covered by other sponges, predominantly

Petchastrissa pathologica (Schmidt, 1868) or

Rhaphisia laxa Topsent, 1892 in which case only

the filaments are visible. The filaments most

often cover the rocky surfaces, foliowing the irre-

gularities of" the substratum or instnuattng into

small cavities such as empty serpulid worm

tubes. They may also run on the surface of other

sponges. They differ from the stretched filaments

described in some Chondrillidae or

Homoscleromorpha living under sledges, which

ZOOSYSTEMA • 1998 • 20(1)

Vacelet J. & Perez T.

are elongated areas of sponge tissue stretched by

the weighc of a detached piece of substratum

(Gaino & Pronzato 1983). The sponge is not

firmly attached and can be easily removcd from

the substratum.

This aspect has been observed vear round. Visual

observations over two years and the comparison

of serial underwater photographs of the same

individual during a six months period from July

to December (Fig. 4B, C) indicate a rather stable

situation ovet tirne, with a slight increase of the

surface covered by the sponge. The filaments and

body maintained the same general shape over six

months. Some of the filaments increased in dia-

meter and coalesced, and a small zone which was

colonized by a loose réticulation in July, was

wholly covered in December. Conversely, a few

filaments regressed, but this was tess common.

The spider-web like réticulation on the rock

around the sponge consec]uently does not corres¬

pond to a rapid régression or a fragmentation

process, but rather to a slow growth process of a

sponge maintaining this spécial shape over long

periods.

Colour

Colour is whitc in life. Preserved specimens are

cream coloured. Some specimens, especially the

holotype, hâve turned clear pink in alcohol after

fixation in formalin.

The surface is smooth, appearing sometimes irre-

gular because the sponge closely follows the irre-

gularities of the substratum. There is no

détachable ectosome. The oscules, 0.5 to 1 2 mm

in diameter (mcasurements from underwater

close-up photographs), are rare on the body and

absent from the filaments. There is no

pore-sieve, and the ostia are not visible. Canals,

approximately 1 mm in maximum diameter, are

visible under the ectosome in the body and in

the largest filaments, with only one canal in each

filament.

Texture

Texture is fleshy, rather soft and fragile.

Skeleton

Spiculé, spongin fibre, bundle or thick condensa¬

tion of collagen fibrils are absent.

Anatomy and cytology

The dertnal structure (Fig. 5A, B) consists of a

thin ectosome, with a layer ot T-shaped exopina-

cocytes and a thin cuticle, without any urider-

lying spécial ectosome difFcrendation. Exopina-

cocytes hâve no flagellum. Their cytoplasm

contains dense granules similar to those of the

granular cells ol the rnesohyl, although smaller

(0.4 to 1.1 pm) and less numerous. Vacuoles

with sytnbiodc bacteria are absent. The latéral

expansions of the exopinacocytes are frequently

superposed for a few micromètres, and are linked

without specialized cell junedons. The thickness

of the pinâcocyte expansions is variable, from

very thin sheets without any inclusions to swel-

lings, 3 to 4 pm thick, containing dense gra¬

nulés. The outer surface of the exopinacocytes is

covered by an extremely thin cuticle, covered by

a well organized fibrillar material, which is quite

similar to the layer termed "glycocalyx" by

Willenz (1981, 1982) in / femimyaile evlumella

Burton, 1934. This outer cover of the pinaco-

derrn, ü 12 to 0.25 pm in total thickness, is

made up of a clear zone, containing fibrils mostly

perpendicular to the outer cell membrane of the

pinâcocyte, followed by a dense zone which is

covered by thin erecr fibrils, which are denser at

their extremities. Contrary to Hemimycale colu-

mclla there are no bacteria fixed on this external

"glycocalyx’', although bacteria are occasionally

engulfed. This structure is found on the outer

surface of both body and filaments of the spon¬

ge. In places, the exopinacodertn displays invagi¬

nations, approximately 2 to 4 pm in diameter,

leading to small canals (Fig. 5A). The canals are

lined by pinacocytes which hâve a similar '‘glyco¬

calyx”, but which are more ovoid in shape and

with a nucléus 2.8 to 3 pm in diameter. These

Fig. a. — Myceliospongia araneosa n.g. nsp., A specimen in

situ, B, in situ vie w, 11 .VI 1,1996; C, same as ligure B,

3.XII.1996; although the surface covered by the sponge stightly

Increased (horizontal arrows), most filaments remamed stable,

rearrangement occutred in a few places (vertical arrows). Scale

bars: A, approximately 13 mm; B, approximately 12 mm;

C, 12 mm.

ZOOSYSTEMA • 1998 • 20(1)

New sponges without skeleton

ZOOSYSTEMA • 1998 • 20(1)

Vacelet J. & Perez T.

openings are probably ostia, although of an unu-

sually small size.

The undersurface in contact vvith the substratum

is rnade up of ovoid basopinacocytes which are

lined by a cutitle similar to that of the exopina-

coderm, although it appears devoid of a “glycoca-

lyx” layer.

The choano.some also shows a very unusual

structure. Canals and choanocyte chambers

(Fig. 6A) are rare in sections. The canals visible

in the in situ, specimens are small and difficult to

observe in preserved specimens, tvherc they are

probably contracted. They are lined by ovoid

endopinacocytes, with a nucléus of 1.9 to 2.5 pm

in diameter, vvilhout fiagellum and containing

dense granules sirnil.tr to those of the exopinaco-

cytes (Fig. 51), E). The endopinacocytes are

devoid of the granulo-fibrillar layer (“glycoca-

lyx”) présent on rhe exopinacocytes. Most of the

choanosomal tissue is rnade up ol a single cell

category, i.e. granulat cclls of highly diverse size

and shape, containing a variable amount of dense

inclusions and invracellular symbiotic bacteria.

Spherulous cells are absent.

Granulat cclls (Fig. 5A-C, £), with a frequently

nucleolated nucléus 2.4-4 pm in diameter, may

be up to 40 pm in maximum diameter when

ovoid or spherical. They contain a variable

amount of spherical inclusions which are highly

variable in size front 0.2 to 12 pm in diameter.

Inclusions, which are surrounded by a membrane,

contain a large mass made up of dense ovoid gra¬

nules, 0.1 pm in maximum size, included in a

dense matrix. This tnass, usually spherical, is

often deformed bv a lew lipid globules or irre-

gular, myelinic-liké granules, These inclusions

are intense)}' metachromatically stained purple

by toluidine hlue in senh-thin sections. The large

granular cells also contain symbiotic bacteria

included in vacuoles of variable size, which often

occupy most of the cc|! volume, and stain clear

blue with toluidine hlue (Figs 5E, 6C, D). The

bacteria belong to several different morphologi-

cal types, front rod-lilce bacteria (0.1 5/1.7 pm) to

ovoid cells (0.8/1.7 pm, rarely up to 1/3.7 pm).

They display the varions morphologies found in

demosponges having a high densiry mesohyl

(Vacelet 1975; Vacelet & Donadey 1977: Boury-

Esnault et al. 1995). Most of them hâve a com¬

plet cell wall. Cells of Type E (Vacelet 1975),

witlt an enlarged periplasm bearing an indenta¬

tion, are frequent.

The shape of rhe granular cells varies importantlv

according to rhe zone of the mesohyl, from sphe¬

rical to vert’ elongated cells. The elongated cells,

which hâve smaller and clearer granules, no bac¬

teria, and long pseudopodia, are usually in paral-

lel arrangement in tracts, especially in the

filaments, suggesting intense, orientated cell

migrations akuig trails vvith denscr and polarized

collagen librils (Figs 5C, 6B). Intermediate stages

between elongated cells and spherical cells are

numerous.

Choanocyte chambers (Fig. 6A) are présent in

low density, and only a few of them havç beeti

observed. They are ovoid, approxirnately 7.5 to

15/20 to 24 pm. The choanocytes hâve a cylin-

drical (2.5 pm in maximum height) or flattened

body, with a spherical, anucleolate nucléus 1.6 to

1.8 pm in diameter. Their base displays latéral

lamellipodia, up to 5.6 pm long, which atrach to

or cover the appendages of rhe othet choano¬

cytes. The choanocyte chamber is thus surroun¬

ded by a thin cellular sheet formed by the

choanocyte latéral appendages. The tyfoplasm

contains numerous clear vacuoles and sortie small

metachromatic inclusions, lêss fhan I pm in dia-

metcr, which hâve the saine ultrastructure as the

smaller unes observed in granular cells. The col-

lar is rnade up of thirty-eight or tliirty-nine

microvilli and is surrounded externally by a “gly-

cocalyx" consisting of a réticulation of thin

fibrils. The fiagellum bas no Yane, but displays a

sheet ot fibrillar niaterial similar to that observed

outside the collât. No âpopylar or cotre cells were

found. but their absence has to be checkcd on a

larger tiumber ol chambers. Fhe mesohyl has a

low dcnsity around the choanocyte chambers,

with a few extraçellular symbiotic bacteria and

highly dispetsed collagen fibrils.

Fie. 5. — Af. araneosa n.sp. A, TEM ot the sponge surface;

pi. exopmacocyte. os. ostium (?): B, TEM ol the sponge surface

showing thé pinacocyle layer with a cet junctïon (arrow), the

culicle (c). the 'glycocalyx 1 layer, and a granu'ar cell; C TEM of

the mesohyl showing elongate granulat oeils and a lewestracel-

lular bacteria; D, TEM ol a small canal lined by an endoplnaco-

cyte; E. TEM ol a canal lined by endopinacocytes and granular

cens,, one or which (letll contains numerous irwacetluiar bacte¬

ria. Scaie bars: A, 3.2 pm; B, 0.6 pm; C, 1.6 pm; D, 1.6 pm;

E, 2.0 pm.

ZOOSYSTEMA • 1998 • 20(1)

New sponges without skeleton

ZOOSYSTEMA • 1998

Vacelet J. & Perez T.

The intercellular matrix is made up of collagen

fibrils wirh variable densicy. Fibrils are most often

irregularly dispersed, except in zones vvitb don-

gated granular cells and beneath the pinacoderm,

where tbey form thin, poorly organized fasdcles.

Fibrils are of the rough type (Garrone 1978),

18-19 nm in diameter, with a poorly defined

striation of approxitnately 22 nm periodicity.

The matrix contains dispersed extracdlular bac-

teria (Fig. 5C) which do not display the high

morphological variety of the intracellular bacte-

ria. Most arc rod-shaped cells, 0.2-0.3/1-1.3 pm,

with a dense area in the clear central nuclear

zone.

Reproduction

No stages of reproduction has been observed.

Remarks

This sponge without skeleton is very unusual

with regard lo anatomy and cytology, with a

remarkable uniformity of cell types, a redticed

aquiferous systc-m, and an unusual morphology

and mode of growth, These peculiarities, which

are observed year round in ail specimens exami-

ned, are not related to stages of reproduction,

which rnay icmporarily change rhe anatomy and

cytology of sonie demosponges such as Halisami

drasticallv (Lévi 1956; Chen 1976; Bergquist

1996). They are also not related to a possible ri.s-

sue régression, which h-as been described in over-

wintering specimens ot in sponges undergoing

degeneration by Iragmentation under adverse

conditions, Tissue régression causes a réduction

or loss of the aquiferous System and a dedifleren-

tiation ofmosc cell types (Simpson 1984), which

frequentlv becotnc archaeocyte-like cells with

numerous residual bodies, thus resulring in fea-

tures which are retnitiiscent of those observed in

Myceliospongid. Flowever, the stabiliry of charac-

ters in ail rhe examined specimens and our obser¬

vations on rhe growth of Myceliospongid during a

six months period (Fig. 4B, C) both indicate

clearly rhat \ve hâve observed normal, non-

degenerating sponges. The relative resemblance

of the cell iéatures to those described during tis¬

sue régression only indicates- that the mesohyi

cells are herc eonstamly poorly differentiated.

The above characteristics do not permit the allo¬

cation of Myceliospongid to any known family of

Demospongiae in which non -skeleton ized gtnera

hâve been described. The simplicitv of rhe cyto¬

logy is shared with the Homoscleromorpha, in

which rvvo généra without skeleton are known

(Oscarella and Pseudocorticium). However,

Myceliospongid does not display any ot the

unique characteristics of this subclass, such as fia-

gellated exo- and endopinacocytes, large choano-

cyte chambers, and a unique basement

membrane underlying both pinacoderm and

choanoderm (Boury-Esnault et al. 1984 ; Boute

et al. 1996). The histology and anatomy difïer

extensively from those ot Chondrillidae and

Darwinellidae, familieS in which rhe généra

Chondrosia and Hexadella respectively are devoid

of skeleton. The sponge is clearly distinct from

the new genus ot Chondrillidae, Tbymosiopsis,

described earlier in this paper. h has neither the

ectosomal organizarion ot collagen fibrils nor the

rubular, branched choanocyte chambers which

are distinctive ot the order I lalisarcida (Bergquist

1996). The possibly distinctive features shared in

varying measure by species ol Halisarcidae and of

Chondrillidae, such as a granulo-frbrillar matrix

distinct from the collagen zones in which the

choanocytes are anchored by long pseudopodia

(Vacelet & Donadey 1987), arc absent. There is

no possible telationship to Bajalus Lendenfeld,

1885 which has recently been shown to be a

synonym of Halisarça (Bergquist 1996).

Furthermore, the unique characters of Mycelio¬

spongid provide no cleat évidente which would

permit affiliation with any exisring order of ske-

letonized Demospongiae. The tibrillar layer

covering the exopinacocytes resembles the “gly-

cocalyx” of the poecilosclerid Hemimycale colu-

mella (Willenz 1982). The absence of typical

spherulous cells and the large number of inclu¬

sions similar to residual bodies in most cell types

are shared with the kératose sponge Dysidea

dvd.ra (Uriz et al, 1996). However, no relation¬

ship between Myceliospongid and the généra

Fig, 6. — M. araneosa asp. A. TEM oi a choanocyte chamber;

noie lhe latéral expansions of the choanocytes, B semi-thin

section through lhe mesohyi near the uodersurface (bottom), C.

TEM ot a granular cell with numerous intracellular bacteria, D,

Intracellular bacteria in a granular cell. Scale bars: A, 1.3 pm; B,

22.5 pm; C, 1.6 pm; D, 1.0 pm.

ZOOSYSTEMA • 1998 • 20(1)

V«£H

rV** *

L *

IA a

Vacelet J. & Perez T.

Hemimycak and Dysidea should be suggested on

che basis of these fearures alone. The only évi¬

dence is che rough nature of the collagen fibrils,

which provide.s a weak indication of affiliation

with the subclass Ceractinomorpba (Garrone

1978). The relationships of the genus within the

Demo-spongiae thus canner be resolved at pré¬

sent. The absence of morphological or cytologi-

cal affinitics with any order of Demospongiae

could justif’y the distinction of a new order.

However, wc prefer to presently classify

Mycelwspongta as a genus incertae sedis within the

Demospongiae, possibly' wirhin the Ccracti-

nomorpha, pending furrher biochcrnical, repro-

ductive or genetic information.

Unlike most other sponges without 1 skeleton, the

absence of fibres or spiculés is nor countcrhalan-

ced by a development of dense fascicles of colla¬

gen fibrils, as it is in Chondrillidae or Halisar

cidae, or by a basement membrane as is the case

in Homoscleromorpha. Consequently, the spon-

ge is soft and fragile, and probably would not be

able to withstand modcrately exposed littoral

environments.

The unique body organization, wirh a reduced

aquiferous System, few choanocyte chambers, a

remarkahly developed System of expansions on

the substratum which increases the external

exchange surface, and a mucous sheet on the sur¬

face suggest.s that tire sponge has a peculiar life

strategy. Apparently, it is able to survive and even

to grow expansions when ihe whole body is cove-

red by massive sponges which smother most of

the surface. It may be supposed that filaments

hâve a rôle in the capture and ingestion of par-

ticles, with the mucous sheet possibly acting as in

Hemimycale cotumella ("Willenz 1981, 1982).

Further studies are needed to elucidate how this

sponge is funedoning.

DISCUSSION

These two sponges are known only front the 3PP

cave. This habitat is highly unusttal in having

stable température conditions, which approxi-

mate rhose of rhe deep Mediterranean which is

homéothermie at 13 °C. It is quite unlikely that

the sponges hâve been overlooked in other caves

front the Marseille area, especially Myceliospongia

araneosa n.sp. with its remarkable growtli form.

The cave has been submerged since the last

Holocene sea level rise, 7000-8000 years ago, a

relatively short titnc which would not permit

local différentiation of sucb raxa. Consequently,

die restricted known distribution to a cave which

shelters a number of deep-sea invertebrates never

recorded in other caves (Vacelet et ai, 1994)

implies thaï the two new sponges also live in the

bathyal zone, from where they hâve colonized

the cave. The 3PP cave is only 7 km distant from

the Cassidaigne canyon, which is 100 to 3000 m

deep with a poorly known invertebrate fauna.

Although the sponge fauna of the canyon has

been rhoroughly explored (Vacelet 1969), the

numerons precîpitous walis and downward-

facing surfaces certainly shelter many unrccorded

sessile invertebrates which are verv difficult or

impossible to sample and observe. It is likely that

propagules of the bathyal invertebrates présent in

the 3 PP cave hâve been advccted by an intermit¬

tent, strong upwelling current frequent in this

area (Bourcier 1978; Millot 1979). These inver¬

tebrates are absent frotn other littoral aphotic

environments because of their stenothermic cha-

racter.

The two new sponges significantly increase the

already remarkahly high number of sponges

without any skeleton which live in dark caves.

This remarkable abundance, which is probably

shared by bathyal vertical and overbanging sur¬

faces as discussed above, is not clearly related to

any chnracteristk of this environment. It has

been hypothesized that in karsric cave, a possible

reduced amount of silica in the water prevents

some demosponges from fully developing rheir

spiculé skeleton, thus explaining a relatively high

frequency of spiculé abnormalides (Bibiloni &

Cili 1982; Bibiloni et al. 1989). However, ir has

been shown (Fichez 1989) that in a karsric cave,

the amount of silica increases from the entrance

to the terminal part, as a rcsult of minerali/ation

processes during the résidence urne of rhe water

in the cave. Il appears more likely that the extre-

mely low hydrodynamic energy experienced by

most dark Caves and deep-sea environments

allows the development of fragile invertebrates,

such as sponges without skeleton or without a

ZOOSYSTEMA • 1998 • 20(1)

New sponges without skeleton

highly developed collagen cortex, which could

not withstand expo.sed littoral environments.

Acknowlcdgem en ts

We greatly acknowledge the assistance of

Ms C. Bézac for prepararion for TEM suidies and

photography, Mr C. Jalong lor diving assistance

and Mr C. Marshall for photography. We thank

Professor P Bergquist, DBE, for tare fui reading

and improvemenrs of the manuscript. ’l'his work

was supported by the ‘‘Programme National

Dynamique de la Biodiversité et Environnement”.

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Submitted on 13 J une 1997;

accepted on 11 August 1997.

ZOOSYSTEMA • 1998 • 20(1)

Deux nouveaux Neoheligmonella (Nematoda,

Trichostrongylina, Nippostrongylinae)

parasites de Mastomys erythroleucus

(Muridae) au Sénégal

Maiick DIOUF

Laboratoire Eau et Santé, Orstom, B. P. 1386, Dakar (Sénégal)

Laboratoire de Parasitologie, Département de Biologie animale,

Faculté des Sciences et Techniques de l’Université CFI. A. Diop (Sénégal)

Cheick Tidiane BÂ

Laboratoire de Parasitologie, Département de Biologie animale,

Faculté des Sciences et Techniques de l'Université CFI. A. Diop (Sénégal)

Marie-Claude DURETTE-DESSET

Laboratoire de Biologie parasitaire, Protistologie, Helminthologie, associé au CNRS,

Muséum national d'Histoire naturelle,

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

MOTS CLÉS

Neoheligmonella skirringi n.sp.,

Neoheligmonella mastomysi n .su..

Nematoda,

Trichostrongylina,

N ippostrongylinac,

rongeurs

Muridae,

Sénégal.

Diouf M., Bâ C. T. & Durette-Desset M.-C. 1998. — Deux nouveaux Neoheligmonella

(Nematoda, Trichostrongylina, Nippostrongylinae) parasites de Mastomys erythroleucus

(Muridae) au Sénégal, Zoosystema 20 (1) , 23-30.

RÉSUMÉ

Description de deux nouveaux Nippostrongylinae (Heligmosomoidea,

Heligmonellidae) Neoheligmonella skirringi n.sp. et N. mastomysi n.sp. para¬

sites de Mastomys erythroleucus dans la province de Casamance au Sénégal.

Parmi les Neoheligmonella , ces deux espèces sont les seules avec A', bainae

Durette-Desset, 1970, parasite de Steatornys opimus au Burkina Paso et

N. crânien Durette-Desset et Cassone, 1986 parasite d ' Umnomys nuldi en

Côte d’ivoire à posséder les caractères suivants ; (1) carène moyennement

développée ; (2) gradient de taille des arêtes dorsales très léger ou absent ;

(3) bourse caudale avec des côtes 2 et 3 de longueur équivalente aux côtes 5

et 6. N. tnt nier! se différencie immédiatement des deux espèces par l’absence

de carène au milieu du corps. N. bainae se distingue de N. skirringi par la

présence de six arêtes ventrales au lieu de huit et par un synlophe prévulvaire

bien développé. Il se distingue de N. mastomysi par des côtes 8 qui naissent

symétriquement sur la tôle dorsale, par des côtes 9 recourbées cl par un cône

génital bien développé. N. skirringi et N. mastomysi se différencient J’unc de

l’autre par le nombre d’arêtes tuiiculaires, la position du pote excréteur, la

naissance des côtes 8 sur la côte dorsale, le synlophe prévulvaire et le nombre

de rangées d’oeufs dans l’utérus. Les deux espèces, bien que de régions géo¬

graphiques proches, ne sont jamais trouvées ensemble chez le même individu

hôte mais, dans chaque région, leur prévalence est similaire.

ZOOSYSTEMA • 1998 • 20(1)

Diouf M., Bâ C. T. & Durette-Desset M.-C.

KEYWORDS

Neobdigniomila skirringi n.sp..

Neoheligmonelk rnastomysi n.sp.

Ncmatoda.

Trichosircmgylina,

N ippost ron gyl i lue,

Muridae,

rodents,

Sénégal.

ABSTRACT

Two new N eoheligmonella (Nematoda, Tricbostrongylimt, Nipposmmgylinae)

parasitic in Mastomys cjythroleucus fiom the Republie of Sénégal. Two new

species belonging to the Nippostrongylinae (Hcligmosomoidea,

Heligmonellidae) are dcscribed. Neubeligmunella skirringi n.sp. and N. masto-

mysi n.sp. are both parasites of Mdftontys eryihroleticus front the Republic of

Sénégal (Province of C.isamancc). Atnong the spccics belonging to the genus

Neobeligimntclla , N. bainae Dtirette-Desset ei Cassone, 1970 parasitic in

Steatamp vpimus (rom Burkina Faso, N, iranien Durette-Desset et Cassone,

1986, parasitic in Uranomys ruddi Iront the lvory Coast, and the two new

species hâve rite following chaiacters; (1)' carcne of average development;

(2) gradient in the size of the dorsal cresrs which is very slight or absent;

(3) caudal bursa with rays 2 and 3 of equal length to thaï of tuys 5 and 6,

N. tmnieri diffère front N. skîmngi and N. rnastomysi in the absence of the

caréné at the inid-body ievel. N. bainae differs front N. skirringi in the pré¬

sence of six ventral vs eight ridges and a well-developed prevulval synlophe. It

differs front N. rnastomysi in that there is a symetrical origilt oi rays 8 on the

dorsal ray, rays 9 are hook-shaped and the génital conc is well-developed.

N. skirringi and N. tjMStomysi are dilfcrentiated front each other bv the liurn-

ber of ctiticular ridges, the position of'rhe excrcrory pore, the origin of rays 8

on the dorsal ray, the prevulval synlophe and the liumber ol row.s of eggs in

the utérus.The two species, although found in neighbouring régions, do not

occur together in the same individual host but in each area the prevalences

species are similar.

INTRODUCTION

Nous poursuivons dans ce travail F identification

des Nématodes Trichostrongles, récoltés depuis

1968 par les équipes de tnammalogistes de

l’Orstom lors de leurs études sut l écologie, la

systématique et la génétique des Rongeurs du

Sénégal (Hubert 1977 ; Poulet 1982 ; Granjon

1987 ; Duplantier 1988). Dans une première

note (Diouf et al 1997). nous avons décrit deux

Nippostrongylinaê, l’un appartenant au genre

Neoheligmonella Durette-Desset, 1970 chez

Arvicdnthis niluticus (Desmarcst, 1822), dans la

province de Dielmo, et l'autre au genre

Heligmonind Baylis, 1928 chez Mastomys erythro-

leucus (Temtninck, 1853) en Casalmmce.

Dans ce travail, nous décrivons deux nouvelles

espèces de Neoheligmonella chez Mastomys ery-

throleucus en Casamance. À notre connaissante,

c’est la première fois qu’une espèce de ce genre

est décrite chez un Mastomys alors que le genre

Heligmonind est représenté chez cet hôte par

H. affinis (Baylis, 1928) au Niger, H. bignonensis

Diouf et al., 1997 en République du Sénégal et

par H. chabaudi (Desset, 1964) en République

centrafricaine et en République du Congo chez

le Mastomys à trente-deux chromosomes.

MATÉRIEL ET MÉTHODES

Les Rongeurs ont été capturés vivants à 1 aide de

pièges rigides de type Manufrance au Cap

Skirring, localité située dans le Sud du Sénégal.

Apres attropsie des hôtes, les Vers onr été fixés à

l'éthanol à 70° bouillant dans lequel ils sont

conservés. Lois de l’étude morphologique, les

spécimens sont montés entre lame et lamelle

dans du lactophénol d'Amann.

La nomenclature utilisée au-dessus du groupe

famille est celle de Durette-Desset & Chabaud

(1995). Le synlophe esc 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 (1981).

Le genre Neobeligmondlri avant une bourse cau¬

dale subsymétrique, le rapport hauteur sur lar¬

geur a été calculé sur la bourse caudale étalée.

ZOOSYSTEMA - 1998 - 20(1)

Nouveaux Nippostrongylinae de Mastomys erythroleucus au Sénégal

Pour les mensurations, le premier chiffre corres¬

pond à Phototype ou à l'allotype, les chiffres

entre parenthèses aux extrêmes des paratypes

mesurés. Pour Neoheligrnonella skirringi, tous les

types disponibles ont été mesurés. Pour

Neoheligmonella mastomysi, dix mâles et dix

femelles du matériel-type ont été mesurés.

Les spécimens-types ont été déposés soit dans les

collections de Biologie animale de PUniversité

CH. A. Diop du Sénégal (CS), soit dans les col¬

lections du Muséum National d’Histoire natu¬

relle de Paris (MNHN).

SYSTÉMATIQUE

Neoheligrnonella skirringi n.sp.

(Fig. 1)

Matériel-type. —- 6 holotype, 9 allotype CS A20,

4 SS, 5 9 9 paratypes, MNHN 558 HF, 3 d<3,

5 9 9 paratypes CS A22.

HÔTE. — Mastomys erythroleucus (Temminck, 1853),

(Muridae).

Localisation. — Tiers antérieur de l'intestin grêle.

Origine GÉOGRAPHIQUE. — Cap Skirring (Casa-

mance, Sénégal), date de récolte : 20.X1.1994.

Autre matérhu examiné- — 2 â â , 7 9 9

CS Al8 ; 1 d, 1 9 CS A2U 2 dd, 8 9 9 CS A26 ;

2 d d, 8 9 9 CS A33 ; 1 d, 9 9 9 CS A35 ; 2 9 9

CS A39. Même hôte, même origine, date de capture

encre le 18 et le 22 novembre 1994.

Description

Nématodes de petite taille enroulés en spirale de

façon senestre le long de leur ligne ventrale, selon

quatre tours de spire chez le mâle, six à sept chez

la femelle. Pore excréteur et deirides situés au

même niveau, juste en arrière de Panneau ner¬

veux. Œsophage musculaire légèrement plus long

que l’œsophage glandulaire (Fig. IA).

Tête

Vésicule céphalique présente. Ouverture buccale

triangulaire. En vue apicale, on observe deux

amphides, quatre papilles labiales externes et

quatre papilles céphaliques (Fig. 1 B).

Synlophe

(étudié en coupe transversale, chez trois mâles et

trois femelles paratypes). Dans les deux sexes,

corps parcouru longitudinalement par quinze

arctcs cuticulaires continues naissant juste en

arrière de la vésicule céphalique (Fig. 1C-1)) Les

arêtes disparaissent en avant de la bourse caudale

chez le mâJe ; chez la femelle, les arêtes ventrales

disparaissent en avant de la vulve, les dorsales

légèrement en arrière de ecllc-ci (Fig. 11., M).

Distribution des arêtes au milieu du corps : cinq

arêtes dorsales, huit ventrales et une carène bien

développée, plus grande chez le mâle que chez la

femelle. La carène est formée par les deux arêtes

latérales gauches dont la ventrale est la plus forte.

Gradient de taille décroissant de la gauche vers la

droite pour la face ventrale. Arêtes dorsales de

taille équivalente. Pointe des arêtes dirigée de la

droite vers la gauche avec un axe d'orientation

incliné à 85° sur l'axe sagittal (Fig, IC, D). Chez

la femelle, dans la région prévulvaire, les arêtes

sont orientées perpendiculairement à la surface

du corps (Fig. 1E).

Mâle

2200 (1800-2500) pm de long sur 90 (80-

95) pm de large dans sa partie moyenne.

Vésicule céphalique haute de 55 (45-60) pm sur

20 (20-20) pm de large. Anneau nerveux, pitre

excréteur et deirides situés respectivement â 175

(165-180) pm, 320 (310-330) pm et 330 (320-

345) pm de l'apex. Œsophage long de 450 (420-

470) pm (Fig. IA).

Bourse caudale subsymétrique de type 2-2-1

(Fig. IJ). Rapport hauteur sur largeur de la bour¬

se caudale de 0,30 (0,29-0,32). Extrémités des

côtes 4 coudées à angle droit vers l’avant. Côtes G

petites er éloignées des côtes 8. Côtes 8 naissant

symétriquement dans le tiers antérieur de la côre

dorsale. Dorsale divisée dans son tiers distal en

deux rameaux eux-metnes biiurqués, les rameaux

externes (côtes 9) sont recourbés vers l’avant et

plus longs que les internes (côtes 10). Cône géni¬

tal bien développé, portanl la papille zéro en

forme de languette sur sa lèvre ventrale et deux

petites papilles 7 sur sa lèvre dorsale. Spiculés fili¬

formes, ailés, longs de 350 (320-380) pm, glis¬

sant dans un gubernaculum long de 40

(40-45) pm sur 25 (20-25) pm de large à sa base.

ZOOSYSTEMA • 1998 ■ 20 (1 )

Diouf M., Bâ C. T. & Durette-Desset M.-C.

Chaque spiculé se termine par une pointe aiguë

(Fig. IJ).

Femelle

2800 (2300-3000) pm de long sur 110 (110-

115) pm dans sa partie moyenne. Vésicule

céphalique haute de 60 (55-65) pm sur 22 (20-

25) pm de large. Anneau nerveux, pore excréteur

et deirides situés respectivement à 150 ( 145-

155) pm, 185 (170-190) pm et 220 (210-

225) pm. Œsophage long de 300 (290-350) pm.

Monodelphie : vulve s’ouvrant à 120 (110-

125) pm de l'extrémité caudale. Vagina vera long

de 30 (30-35) pm. Vestibule, sphincter et trompe

longs respectivement de 125 (120-135) pm, 30

(30-35) pm et 55 (50-60) pm (Fig. IL). Utérus

long de 525 (515-610) pm contenant 11 (9-

18) œufs au stade morula, répartis sur une rangée

et hauts de 65 (55-70) pm sur 55 (40-60) pm de

large. Queue de 45 (35-50) pm de long, forte¬

ment recourbée ventralement entre la vulve et

l’anus (Fig. IL).

Discussion

Voir discussion de Neoheligmonella mastomysi.

Neoheligmonella mastomysi n.sp.

(Fig. 2)

Matériel-type. — â holotype, 9 allotype CS B6,

5 â 6 , 7 9 9 pararypes CS B7 et 3 â â, 3 9 9 para-

types MNHN 559 HF.

Hôte. — Mastomys erythroleucus (Temminck, 1853)

(Muridae).

Localisation. — Tiers antérieur de l’intestin grêle.

Origine géographique. — Cap Skirring (Casa-

mance, Sénégal), date de récolte : 19.XJ.1994.

Autre matériel examiné. — 1 d, 1 2 CS B1 -, 1 2

CS Bll ; 1 d CS B21 ; 3 6 6, 17 9 2 CS B23 ;

1 d, 1 9 CSB24:3 dd,2 22 CSB32; 1 d, 1 2

CS B38. Même hôte, même origine, date de capture

entre le 18 et le 22 novembre 1994.

Description

Nématodes de petite taille enroulés en spirale de

façon senestre le long de leur ligne ventrale, selon

un à deux tours de spire chez le mâle, trois à

quatre chez la femelle. Pore excréteur situé juste

en arrière de l’anneau nerveux ; deirides légère¬

ment postérieures. Œsophage musculaire légère¬

ment plus court que l’œsophage glandulaire

(Fig. 2A).

Tête

Vésicule céphalique présente. Ouverture buccale

triangulaire. En vue apicale, on observe deux

amphides, quatre papilles labiales externes et

quatre papilles céphaliques (Fig. 2B).

Synlophe

(étudié en coupe transversale chez deux mâles et

trois femelles). Dans les deux sexes, corps par¬

couru longitudinalement par treize arêtes cuticu-

laires continues naissant en arrière de la vésicule

céphalique sauf l’arête dorsale gauche et l’arête

latérale droite qui naissent à environ 60 (50-

70) pm en arrière de la vésicule céphalique

(Fig. 2F-I). Elles disparaissent en avant de la

bourse caudale chez le mâle et à différents

niveaux entre la vulve et l’anus chez la femelle

(Fig. 2J, K). Disrribution des arêtes au milieu du

corps : cinq arêtes dorsales, cinq ventrales, une

petite arête proche du champ latéral droit et une

carène bien développée (ormée par les deux

arêtes latérales gauches donr la ventrale est la plus

forte. À l’exception de la petite arête droite, gra¬

dient de taille des arêtes décroissant de la gauche

vêts la droite sur la face ventrale. Sur la face dor¬

sale, gradient de taille décroissant de la droite

vers la gauche chez le mâle et absence de gradient

chez la femelle. Pointe des arêtes dirigée de la

droite vers la gauche avec un axe d’orientation

sub-frontal (Fig. 2C, D). Chez la femelle, dans la

région pré-vulvaire, il riexiste pas de modifica¬

tions importantes à l’exception d’une diminution

de la taille de la carène (Fig. 2E).

Fig. 1- — Neoheligmonella skirringi n.sp.. A, 3, extrémité anté¬

rieure. vue latéiale gauche , B 7 tète, vun apicale : C. cS, syn¬

lophe au milieu du corps ; D, ", synlophe au milieu du r.orps ; E.

î. synlophe pré-vulvaire ; F l - . extrémité antérieure, naissan¬

ce des arêtes articulaires . F. vue ventrale : G, vue dorsale ; H,

vue latérale droite ; I, vue latérale gauche : J, 3, bourse cauda¬

le, vue ventrale . K. î, extrémité pastêneure, vua latérale droi¬

te ; L, M. 9, extrémité postérieure, disparition des arêtes

cutlculalret ; L, vue ta’.éralo droite ; M, vue latérale gauche,

Échelles : A, F-K, 60 pm ; B, 30 pm ; C, D, 50 pm ; E, 25 pm ; L,

M, 100 pm.

ZOOSYSTEMA • 1998 • 20(1)

Diouf M., Bâ C. T. & Durette-Desset M.-C.

Mâle

5700 (5400-6000) |im de long sur 100 (90-

110) (ira de large dans sa partie moyenne.

Vésicule céphalique haute de 60 (60-65) prn sur

26 (25-27) ptn de large. Anneau nerveux, pore

excréteur et deirides situés respectivement à 175

(165-180) pm, 195 (185-200) pm et 230 (220-

235) pm de l’apex. Œsophage long de 390 (380-

410) pm (Fig. 2A).

Bourse caudale subsymérrique de type 2-2-1

(Fig. 2L). Rapport hauteur sur largeur de la

bourse caudale de 0,040 (0,039-0,043). Extré¬

mités des côtes 4 coudées à angle droit vers

l’avant. Côtes 6 petites et éloignées des côtes 8.

Côtes 8 naissant asymétriquement sur la côte

dorsale, la côte droite naissant juste en arrière de

la racine de la dorsale, et la côte gauche, dans le

tiers médian de celle-ci. Dorsale divisée à la moi¬

tié de sa hauteur en deux rameaux eux-mêmes

bilurqués, les rameaux externes (côtes 9) étant

plus longs que les internes (côtes 10). Cône géni¬

tal portant les deux petites papilles 7 sur sa lèvre

dorsale et la papille zéro sur la lèvre ventrale.

Spiculés filiformes, ailés, longs de 300 (295-

310) pm, glissant dans un gubernaculunt long de

30 (30-35) pm sur 20 (20-25) pm de large à sa

base. Chaque spiculé se termine par une pointe

aiguë (Fig. 2L).

Femelle

6400 (6200-7000) pm de long sur 150 (145-

155) pm de large dans sa partie moyenne.

Vésicule céphalique haute de 60 (55-65) pm sur

32 (30-35) pm de large. Anneau nerveux, pore

excréteur et deirides situées respectivement à 230

(225-260) pm, 250 (240-260) pm et 380 (370-

385) pm de l’apex. Œsophage long de 450 (435-

470) pm.

Monodelphie : vulve s'ouvrant à 120 (1)5-

125) pm de l’extrémité caudale. Vagina vera long

de 60 (55-75) pm. Vestibule, sphincter et trompe

longs respectivement de 160 (155-170) pm, 40

(35-45) pm et 150 (145-150) pm (Fig. 2M).

Utérus long de 1300 (1200-1500) pm contenant

35 (28-40) œufs au stade rnorula répartis sur trois

rangées dans la partie distale de l’utérus et deux

dans la partie proximale, hauts de 60 (50-65) pm

sur 40 (35-45) pm de large. Queue arrondie à son

extrémité, longue de 50 (45-55) pm (Fig. 2M).

Discussion

Les deux espèces parasites de Mastomys erythro-

lencus décrites ci-dessus présentent les principaux

caractères du genre Neoheligmonella Durette-

Desset, 1971 (Heligmonellidae, Nippostrongyli-

nae) : synlophe avec axe d'orientation des arêtes

incliné au moins à 70° sur l’axe sagittal, arête

latéro-ventrale gauche bien développée, nombre

d’arêtes cuticulaires compris entre dix et quinze ;

bourse caudale subsymétrique de type 2-2-1,

avec un court tronc commun à la côte dorsale et

aux côtes 8.

Ce genre compte quinze espèces parasites, pour

la plupart de Muridae éthiopiens.

Seules N. bainae Durette-Desset, 1970, parasite

de Steatomys opîmus Pousargues, 1894 au

Burkina Faso et N. tmnieri Durette-Desset et

Cassone, 1986, parasite d ' Uranomys ruddi

Dollman, 1 909 en Côte d Ivoire possèdent

comme les spécimens du Mastomys : (1) un syn-

lophe caractérisé par la présence d'une carène

moyennement développée ; (2) une bourse cau¬

dale avec des côtes 2 et 3 de taille équivalente à

celles des côtes 5 et 6 ; (3) une queue femelle for¬

tement recourbée ventralement.

La première espèce parasite du Mastomys

s’éloigne de N, tmnieri par la position médiane

du pore excréteur, par la naissance symétrique

des côtes 8 sur la côte dorsale, par des côtes 9

plus longues que les côtes 10 et surtout par la

présence de la carène tout le long du corps.

Comme N. bainae , elle possède des côtes 8 nais¬

sant symétriquement sur la côte dorsale mais elle

s’en distingue par la présence de six arêtes ven¬

trales au lien de huit, par un synlophe prévulvaire

bien développé et par la présence de deux rangées

d’œufs dans l’utérus.

La seconde espèce parasite du Mastomys possède

comme N. tmnieri des côtes 8 naissant asymétri¬

quement sur la côte dorsale mais elle s’en éloigne

Fig. 2. — Neoheligmonella mastomysi n.sp., A, <È, extrémité

antérieure, vue latérale droite ; B, .. lêle, vue apicale ; C, <J,

synlophe au milieu du corps ; D. V . synlophe au milieu du

corps ; E. V. synlophe prévulvaire ; H, t, extrémité antérieure,

naissance des arêtes cuticulaires ; F, vue dorsale , G vue laté¬

rale droite : H, vu6 latérale gauche . I. vue ventrale : J. K. 9,

extremllè postérieure, disparition des arêtes cuticulaires : J, vue

latérale drolle K, vue latérale gauche ; L, bourse caudale,

vue venttale , U, '< , partie postérieure, vue latérale droite

Échelles : A, C-E, H, I, 75 pm ; B, 25 pm ; F, G, 50 pm ; J, K, M,

100 pm ; L, 125 pm.

ZOOSYSTEMA • 1998 • 20(1)

Diouf M., Bâ C. T. & Durette-Desset M.-C.

par la position antérieure du pore excréteur, par

un cône génital peu développé, par des côtes 9

rectilignes et plus longues que les côtes 10 et sur¬

tout par la présence de la carène tout le long du

corps. Elle se distingue de N. bainae par la posi¬

tion plus antérieure du pore excréteur, par des

côtes 8 naissant symétriquement sur la côte dor¬

sale, par des côtes 9 rectilignes et plus longues

que les côtes 11) et par la présence de deux ran¬

gées d’œufs dans l’utérus.

Les deux espèces parasites du Mastomys peuvent

être séparées l'une de l'autre pat le nombre

d’arêtes cuticulaires, par la présence d’un léger

gradient de taille des arêtes sur la face dorsale du

mâle de la deuxième espèce, par le synlophe pré¬

vulvaire, par la position du pore excréteur, par la

naissance des côtes 8 sur la côte dorsale, par la

forme et la longueur des côtes 9, et par le

nombre de rangées d’œufs dans l’utérus. Nous

proposons de nommer la première espèce

Neoheligmonella skrrringi n.sp., et la seconde

N. mastomysi n.sp. en référence aux hôtes chez

RÉFÉRENCES

Baylis H. A. 1928. — On a collection of nématodes

front Nigérian mammals (chiefly rodents).

Pamùtohgy 20: 284-295.

Desset M.-C. 1964. — Les systèmes d’arêtes cuticu¬

laires chez les Nématodes Héligmosomes. Étude dc-

cinq espèces parasites de Rongeurs de La Maboké.

Cahiers de La Maboké 2 : 39-78.

Diouf M., Bà C. T. H. & Durette-Desset M.-C.

1997. — Deux nouveaux Nippostrongylinae

(Nematoda. Trichostrongylina) parasites de

Muridae du Sénégal. Zonsystema 19 (2-3) : 225-233.

Duplanricr j. M. 1988. — Biologie évolutive de popu¬

lations du genre Mascorays (Rongeur, Mttridè) au

Sénégal : 1-203, Thèse de Doctorat d'État USTL

Montpellier.

Durette-Desset M.-C. 1970. — Les systèmes d’arêtes

cuticulaires chez les Nématodes Héligmosomes.

Cinq nouvelles espèces parasites de Rongeurs afri¬

cains. Cahiers de La Maboké 8 : 125-137.

— 1971. — Essai de classification des Nématodes

Héligmosomes. Corrélations avec la paléobiogco-

graphie des hôtes. Mémoires du Muséum national

d’Hhtotrc naturelle, Série A, Zoologie, 69 : 1-126.

— 1985 — Trichostrongyloid nematodes and their

vertebrate hosts: Reconstruction of the phylogeny

lesquels elles ont été trouvées. Il est intéressant de

constater que ces deux espèces très proches mor¬

phologiquement n’ont jamais été trouvées

ensemble chez le même individu-hôte. Elles

pourraient être interprétées comme des espèces

vicariantes, N. skirringi parasitant les Mastomys

capturés dans des rizières alors que N. mastomysi

a été trouvée chez les Mastomys vivant dans les

zones cultivées aux abords de la mangrove.

Remerciements

Les auteurs remercient le Dr G. Vassiliadès de

riSRA-Productions animales de Dakar et le Dr

J, M. Duplamier de l’Orstom pour leur aide et

leurs conseils lors de la réalisation de ce travail.

Us remercient également le Dr L. Granjon du

Muséum national d’Histoire naturelle de Paris

(Laboratoire de Zoologie, Mammifères et

Oiseaux) pour les renseignements concernant la

nomenclature des hôtes. Ce travail a été financé

et réalisé dans le cadre du Programme « Eau,

Santé et Développement » de l’Orstom.

of a parasitic group. Advances in Parasitology 24:

239-306.

Durette-Desset M.-C. Ht Cassone J. 1986. — Sur

deux Nérnalodes Trichostiongyloides parasites

d'un Muridé africain. I- Description des adultes.

Annales de Parasitologie Humaine et Comparée 61 :

565-574,

Durette-Desset M.-C. St Chabaud A. G. 1981. —

Nouvel essai de classification des Nématodes

I richostrongyloidea. Annales de Parasitologie

Humaine et Comparée 56 : 2.97-372.

— 1995. — Note sur la nomenclature supra-familiale

des Strongylida. Annales de Parasitologie Humaine

et Comparée 68 : 111, 112.

Granjon L. 1987. — Evolution allopatrtque chez les

Muridés : mécanismes èco-èthologiques liés au syndro¬

me d'insularité chez Mastomys et Rattus : 1-163.

Thèse de Doctorat, USTL Montpellier.

Hub ert B. 1977. — Écologie des populations de

Rongeurs de Bandia (Sénégal) en zone sahélo-souda-

nienne. Revue d'Écologie (L'erre et Vie) 31 : 33-100.

Poulet A. 1982. — Pullulation de Rongeurs dans le

Sahel : mécanismes et déterminisme du cycle d'abon¬

dance de Taterillus pygargus et z/Arvicamhis niloti-

cus (Rongeurs-Gerhiuidés et Muridés) dans le Sahel

du Sénégal de 1975 à 1977 : 1-367. Éditions

Orstom, Paris.

Soumis le 6 août 1997;

accepté le 26 novembre 1997.

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia (Crustacea) from the “Campagne

Biaçores” in the Azores région, including a

generic révision of Verrucidae

Paulo S. YOUNG

Museu Nacional, UFRJ

Quinla da Boa Vista, 20940-040, Rio de Janeiro, RJ (Brazil)

Young P. S. 1998. — Cirripedia (Crustacea) from the “Campagne Biaçores” in the Azores

région, including a generic révision of Verrucidae. Zoosystema 20 (1) : 31-92.

KEYWORDS

Cirripedia,

Verrucidae,

new genus and species,

Azores région.

ABSTRACT

The cirripeds sampled hy che N. O. Jean Charcot from the Azores région

include thirty-four species: twency lepadomorphs, eight vcrrucomorphs and

six balanomorphs. Among these are two new species: Arcoscalpellum eponkos

n.sp. and Tesseropora amoldi n.sp. and several little known species. The family

Verrucidae is revised, and a key to the généra is included. Verruca and

Metaverruca are rediagnosed, two new généra are proposed: Newmanivemruca

n.g. and Costatovetruca n.g. A list of recent species of Verrucidae is provided,

reported with keys to ail of the species. Forty-five species of cirripeds are

reported from the Azores région, of which one third are endémie.

MOTS CLÉS

Cirripedia,

Verrucidae,

nouveaux genres et espèces,

région des Açores.

RÉSUMÉ

Cirripèdes (Crustacea) de la « Campagne Biaçores » dans la région des Açores,

avec une révision des genres de Verrucidae. Les cirripèdes recueillis par le N. O.

Jean Charcot dans la région des Açores comprennent trente-quatre espèces :

vingt Lépadomorphcs, huit Verrucomorphes et six Balanomorphes. Parmi

eux, deux espèces sont nouvelles : Arcoscalpellum eponkos n.sp. et Tesseropora

amoldi n.sp., et plusieurs peu connues. Les Verrucidae sont révisés et une clé

des genres est incluse. Verruca et Metaverruca font l’objet d’une diagnose

émendée et deux nouveaux genres sont proposés : Newmanivcrruca n.g. et

Costatovetruca n.g. Une liste des espèces actuelles de Verrucidae est fournie,

ainsi que des clés pour toutes les espèces. Quarante-cinq espèces de cirripèdes

sont signalées de la région des Açores, dont le tiers est endémique.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

INTRODUCTION

The ‘Campagne Biaçores” undertaken by the

N. O. Jean Charcot and directed by Dr Jacques

Forest, sampled chc Azurés Région from the

intertidal to the abyssal zone (4700 m), especially

along the Mid-Atlantic Ridge. During the cruise

at the West European Basin deep-sea samples

were collected.

The decp water fituna of the Azotes région is bet-

ter known than rhat of shallow waters. This is

the resuit of dredging by the Challenger, Hiron¬

delle, Princesse Alice , Travailleur, Talisman , and

more reeently by the Meteor (Hoek 1883;

Aurivillus 1898; Gruvel 1900a, b. 1902a, 1920;

Young in press), The species (ound in shallow

waters were described by Pilsbry (1916),

Newman & Ross (1977) and Baker (1967). The

latter paper by Baker is based on collections from

the Chelsea Expédition at Sâo Jorge Island.

The présent study dcscribes the cirriped spccies

collected during the “Campagne Biaçores : ’ from

intertidal to abyssal depths. The verrucids collec¬

ted during this cruise form the basis for a review

and révision of the Verrncidae.

Ali the specimens are deposited in the Muséum

national d’Idistoire naturelle, Paris (MNHN),

the Museu Nacional do Rio de Janeiro (MNRJ)

and the United States National Muséum

(USNM).

Abbreviations used are as follows:

tl total length;

rc rostro-carinal diameter.

EARLY STUDIES ON THE BARNACLES

OF THEAZORES

The barnacles from the Azurés région were col¬

lected first by H. M. S. Challenger , which dred-

ged around the archipclago, Hoek (1883)

described two new species, Sca/pellum acutum

and Dichelaspis sessilis. Aurivillius (1898) descri¬

bed nineteen species from this région based on

collections from the campagnes of Hirondelle and

Princesse Alice: Scalpelluni debile, S. rigiJum,

S. mamillatum, S, anceps, S. molle, S. erectum,

S. grimaldii, S. calyculus, S. falcatum, S. incisurn ,

5. pusilum, Poecilasma unguiculus, Verruca recta,

V. costata, V, aequalis, V inermis, V. crenata,

V. co ma ta and V, sculpta.

Gruvel (1900a, b) described the species collected

by the Travailleur and Talisman later described in

detail (Gruvel 1902a). In the présent study eight

species new to the Azotes are redescribed:

S edwardsii, S. recurvitergum, S. atlanticum ,

6. striatum, S. luteutn , V. erecta, V. trisulcata and

V. linearis. In addition S. regium Thomson,

1873, and V. str&mta (Millier, 1776) were repor-

ted from tins région. Gruvel also reported

S. gigas Hoek from the Azotes région, but the

coordinates presented appear to reprexent speci¬

mens' collected off the coast of France.

Pilsbry (1916) described Megabalanus tintinna¬

bulant azoricus and recordcd Balanus trigonus

Darwin, 1854, and Chthamalus stellatus stellatus

(Poli, 1791), ail of which are shallow water spe¬

cies.

Gruvel (1920) provided the most complété study

of the barnacles from the Azores, based on a

séries of collections made by the yachts of

S A. S. le Prince de Monaco. He reported

thirty-seven shallow and deep water species, and

added new information regarding the following

species: Y. vulgarc Leach, 1824, S. velutinum

Hoek, 1883, S. alboranense Gruvel, 1920; ,Y. gra¬

cile Hoek, 1907a, S. pi/sbryi Gruvel, 1911,

V. spengleri Darwin, 1854, Il amphitrite Darwin,

1854, and B. sponguola Brown, 1844.

A collection made in the shallow waters of Sâo

Jorge Island (Baker 1967) confirmed the occur¬

rence of C. stellatus, B. trigonus, B. tintmnabulum

azoricus , and V. spengleri. Baker (1967) rqjorted

the prescuce of Tetraclita s quant osa elegans

Darwin, 1854. Howcver, Newman & Ross

(1977) indicated thar T. squamosa elegatn likely

represencs a new species, l'esseropora atlamica.

Zevina (1976) recorded S. vitreum Hoek, 1883,

and 5. michelottianum Segucnza, 1876, from the

Azotes and described 5. limpïdus from off nor-

theast of this archipclago.

Young (in press) studied the fauna of the Great

Meteor Bank and recorded Heteralepas microsto-

ma (Gruvel, 1901) from this région.

ZOOSYSTEMA • 1998 ■ 20(1)

Cirripedia from the Azores Région

THE SPECIES FROM THE “CAMPAGNES

BIAÇORES”

Order PEDUNCULATA Lamarck, 1818

Suborder HETERALEPADOMORPHA

Newman, 1987

Family Hf.TP.RA 1 EPADIDAb Nilsson-Cantell, 1921

Genus Heteralepas Pilsbry, 1907a

Heteralepas microstoma (Gruvel, 1901)

Alepas microstoma Gruvel, 1901: 259: 1902b: 282,

pl. 24, figs IB, B’, 7, 8; 1905: 162, fig. 180.

Material F.XAMINED. — Stn 73, 1 specimen,

d: 2.4 cm, on a gorgonian, MNHN Ci 2562.

Remarks

Heteralepas microstoma is a coin mon species from

the Azores and Madeira Archipelagos, and the

Great Meteor Seamouncs (Young in press). It

ranges from depths of 269 to 623 m, but more

commonly around 300 m, forming large agré¬

gations usually fixed on the stem of octocorals.

Suborder LEPADOMORPHA Pilsbry, 1916

Family OXYNASPIDIDAE Gruvel, 1905

Genus Oxynaspis Darwin, 1852

Oxynaspis patens Aurivilius, 1894

Oxynaspis païens Aurivillius, 1894: 38, pl. 3, figs 1-2,

pl. 6, figs 13-15; pl. 8, fig. 9. — Totton 1940: 476,

fig. 16. - Pilsbry 1907a: 79. Weisbord 1979: 39,

pl. 3, fig. 6-9. - Zevina 1982: 36, fig. 26.

Material EXAMINED. — Stn 89, 2 specimens, tl:

0.9-1.0 cm, MNHN Ci 2563.

Remarks

Oxynaspis patens was described by Aurivillius

(1894) based on specimens from Anguilla Island,

Antilles, between depths of 125-355 m, and later

recorded from the Bahamas (Pilsbry 1916), and

Joséphine and Great Meteor Seamounts, between

170-300 in (Young in press). The record of this

species from the Azores confirms its occurrence

in the Eastern Atlantic (Fig. 1).

Oxynaspis celata Darwin, 1852

Oxynaspis celata Darwin, 1852: 134, pl. 3,

fig. 1. - Gruvel 1905: 103, fig. 114. — Nilsson-Cantell

1921:226. fig. 37.

Material examined. — Stn P44, 1 specimen

without carina, MNHN Ci 2536.

Remarks

This specimen may be identified with certaintv

as O. celata , but lacking in the carina, the sub-

species cannoi be identified. Based on the geo-

graphical distribution (Fig. 2), it is probably

O. celata s.str.

Bacon (1976) and T.eta & Young (1996) accor-

ded species status for O. celata hirtae Totton,

including ail of the tefèrences to O. celata from

the Western Atlantic.. Figure 2 présents the gene¬

ral distribution of the O. celata group, including

the distribution of O. hirtae.

Family POECILASMATIDAE Annandale, 1909

Genus Poecilasma Darwin, 1852

Poecilasma aurantia Darwin, 1852

Poecilasma aurantia Darwin, 1852: 105, pl. 2, fig. 2.

Poecilasma Kempfen var. aurantium — Gruvel 1902a:

31, pl. 4, figs 1-2; 1905: 115, fig. 129. - Weltner

1922: 79.

Poecilasma kaempferi aurantia — Zcvina 1982: 98,

fig. 86m.

Material examined. — Stn 62, 9 specimens,

tl: 0.5-1.8 cm, MNHN Ci 2559, MNRJ 8869. —

Stn 148, 14 specimens, tl: 0.5-1.2 cm, MNHN

Ci 2575, MNRJ 8870. — Stn 157, 10 specimens,

tl: 0.4-1.2 cm, MNHN Ci 2581, MNRJ 8871. —

Stn 159, l specimen, tl: 1.5 cm, MN1 IN Ci 2583.

Stn 180, 2 specimens, il: 1.1-1.3 cm, MNHN

Ci 2665. — Stn 199, 2 specimens,

tl: 0.8-1.1 cm, MNHN Ci 2609. - - Stn 237, 8 speci¬

mens, ti; 0.4-1.0 cm, MNHN Ci 2628.

Remarks

Some ot the species assigned to Poecilasma

kaempferi group are questionable, because most

ot them do not hâve readily diagnostic characters

(Young in press). The specimens herein studied

are assigned to P. aurantia. Ail of the specimens

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

examincd agréé to Darwins (1852) description

of P. durantia , and the type locality of this species

is thc Madeira Archipelago. A révision of this

group is needed.

Genus Glyptelasma Pilsbry, 1907a

Glyptelasma hamatum (Caïman, 1919)

Megalasma ( Glyptelasma) hamatum Caïman, 1919:

370, fîgs 5-7. — Nilsson-Cantell 1927: 770, fig. 12;

1928: 23. fig. 11; 1931: 10: 1934: 49; 1955:

219. - Weisbord 1979: 48, pl.5, figs 1-2, pl. 14,

fig. 3. - Zevina 1982: 93, fig. 83.

Megalasma hamatum - Zevina 1969: 67; 1976: 1155.

Material EXAMINED. — Stn 180, 1 specimen,

tl: 0.8 cm, MNHN Ci 2666. — Stn 202, 16 spéci¬

mens, tl: 0.9-2.5 cm, MNHN Ci 2613, MNRJ

8872. — Stn 227, 1 specimen, tl: 1.8 cm, MNHN

Ci 2624.

Remarks

Glyptelasma hamatum is defined by the pair of

short, hook-like processes on the prosoma and

by the filamentarv appendage at the base of cir¬

rus I.

This species has a circumrropical distribution,

occurring bctween depths of 457 and 3778 m

(Weisbord 1979) and was recorded previously in

Lhe Eastern Atlantic from the Cape Verde Islands

(Caïman 1919) and off Sierra Leone

(Nilsson-Cantell 1927; Fig. 1).

Fig. 1. — Géographie distribution of Oxynaspis patens Aurivillius (■), Glyptelasma hamatum Caïman (*), Arcoscalpellum tritonis

(Hoek) (A) and Hexelasma americanum (Pilsbry) (•).

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Family LepadidAE Darwin, 1852

Genus Lepas Linnaeus, 1758

Lepas anatifera Linnaeus, 1758

Lepas anatifera Linnaeus, 1758: 668. - Darwin 1852:

73, pl. 1, fig. 1, la-c. - Pilsbry 1907a: 79, pl. 9,

figs 3-5.

Lepas (Anatifi) anatifera - Zevina 1981: 17, fig. 8.

Materiai. EXAMINED. — Stn 1.7, 10 specimens, tl:

0.9-2.5 cm, MNHN Ci 2539.

Remarks

Lepas anatifera is a species commonly found atta-

ched to floating objects (Pilsbry 1907a;

Weisbord 1979).

Lepas pectinata Spengler, 1792

Lepas pectinata Spengler, 1792: 106, pl. 10, fig. B,

2a-c. - Darvvin 1852; 85, pl. 1, figs 3, 3a. - Pilsbry

1907a: 81, pl. 8, figs 4-6, 8.

Lepas (Anatifi) pectinata - Zevina 1981: 15, fig. 6.

Materiai examine! >. — Stn L7, 7 specimens, tl: up

to 1.0 cm, MNHN Ci 2540. — Stn 120, 16 speci¬

mens, tl: up io 0.6 cm, MNHN Ci 2565. — Stn 131,

more than 100 specimens, up to 0.7 cm, MNHN

Ci 2569, MNRJ 8873.

Remarks

Lepas pectinata is a species commonly found atta-

ched to floating objects, including floating detâ-

ched algae (Pilsbry 1907a; Weisbord 1979). The

specimens, from depths of 2100 and 2120 m,

clearly represent situations wliere decached algae

(Sargassum ) sank to this dcpth or were erro-

neously picked up during dredging work, since

L. pectinata and Sargassum do not live at these

depths.

Genus Concboderma Olfers, 1814

Conchoderma auritum (Linnaeus, 1767)

Lepas aurita Linnaeus, 1767: 1110.

Conchoderma auritum - Darwin 1852: 141, pl- 3,

figs 4, 4a-c. - Pilsbry 1907a: 99, pl. 9, fig. 2. - Zevina

1981: 26, fig. 15.

Material EXAMINED. — Ponta Delgada, 3 specimens,

tl: 1.3-2.0 cm, MNHN Ci 2656.

Remarks

Conchoderma auritum is commonly found on

whales and ships, and also on rurtles (Pilsbry

1907a; Weisbord 1979). The specimens from

“Ponta Delgada" were collected attached to the

hul! of N. O .Jean Charcot.

Suborder SCALPE LLOMORPHA

Newman, 1987

Family CALANTiciDAE Zevina, 1978

Genus Smilium Gray, 1825

Smilium acutum (Hoek, 1883)

Scatpellum acutum Hoek, 1883: 80, pl. 3, fig. 19;

pl. 8, fig. 12; 1907a: 64, pl. 7, fig. 1. - Gmvel 1920:

12, pl. 2, fig. 7.

Smilium acutum — Newman & Ross 1971: 38,

textfïg. 12, pl. 5F (svnonymy). - Foster & Buckeridge

1995a: 166, fig. le,'

Materiai, examiner. —Stn 179, 4 specimens, tl:

0.4-1.3 cm, MNHN Ci 2593, MNRJ 8874.

Remarks

Smilium acutum is a world-wide species

(Newman & Ross 1971). In the Eastern Atlantic

it has been recorded from the West European

Basin, oflf Morocco, Cape Verde, Madeira, and

also the Azores (Hoek 1883; Caïman 1918;

Gruvel 1920; Foster & Buckeridge 1995a;

Fig. 2).

Family SCALPELLIDAE Pilsbry, 1907a

Subfamily SCALPELLINAE Pilsbry, 1907a

Genus Scalpclhwi Leach, 1817

Scalpellum scalpellum Linnaeus, 1767

Lepas scalpclluni Linnaeus, 1767: 1 100.

Scalpellum scalpellum — Darwin 1852: 222, pl. 5,

fig. 15. — Pilsbry 1907a: 16. - Nilsson-Cantell 1978:

16, figs 6-7. - Zevina 1981: 94, fig. 65.

Material examined. — Stn 260, 27 specimens, tl:

up to 1.7 cm, MNHN Ci 2654, MNRJ 8875.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Remarks

Scalpellum scalpellum is a common species along

the European and North African coasts. It ranges

from depths of 10 ro 540 m, but occurs more

commotily between 30 and 200 m

(Nilsson-Cantell 1978).

SubFamily MerosCALPELLINAE Zevina, 1978

Genus Neoscalpellum Newman et Ross, 1971

Neoscalpellum debile (Aurivillius, 1898)

Scalpellum debile Aurivillius, 1898: 189. — Gruvel

1905: 27; 1920: 31, pl. 5, figs 13-15, pl. 7, 6g. 1.

Scalpellum eduiardsü GruveL 1900a: 189; 1902a: 63,

pl. 2, figs 3B, 16; 1905: 28, fig. 27

Scalpellum dicheluplax Pilsbry, 1907a: 70,

fig. 28a-c. — Hoek 1914: 4.

Scalpellum dieheloplax benthophila Pilsbry, 1907a: 73,

fig. 28d.

Scalpellum alboranense Gruvel, 1920: 33, pl. 5,

figs 4-6.

Neoscalpellum debile — Newman & Ross 1971: 96,

figs 49-50.

MATERIAI EXAMINEE!. — Sm 131, 3 spécimens, d:

1.0-1.1 cm, MNHN Ci 2568. — Stn 202, 1 spéci¬

men, d: 0.8 cm, MNHN Ci 2614. — Stn 245, 2 spé¬

cimens, tl: 0.6-1.2 cm. MNHN Ci 2708. — Stn 249,

2 specimens, tl: 0.8-2.3 cm, MNHN Ci 2642. —

Stn 250, 1 spécimen, tl: 3.3 cm, MNHN Ci 2646.

Remarks

Neoscalpellum debile was discussed by Newman

& Ross (1971: 96) and Young (in press). It has a

Nortli Atlantic distribution, with several records

from the Adores Région (distribution map in

Young, in press).

Subfamily ARCOSCALPELLINAE Zevina, 1978

Genus Arcoscalpellum Hoek, 1907a

Arcoscalpellum michelottianum

(Seguenza, 1876)

Scalpellum michelottianum Seguenza, 1876: 381, pl. 6,

figs 15-25. pl. 10, fig. 26.

Scalpellum velutinum Hoek, 1883: 96, pl. 4,

figs 10-11, pl. 9, figs 7-9.

Arcoscalpellum michelottianum — Newman & Ross

1971: 71, texrfig. 34, pl. 9b (synonymy).

MATERJAL EXAMINE,U, — Stn 105. 1550 m, 1 spéci¬

men, tl: 1.3 cm, MNHN Ci 2564. — Stn 180,

10 specimens, tl: 1.2-4.0 cm, MNHN Ci 2594,

MM RJ 8876. — Stn 196, 6 specimens, tl: 1.9-3.1 cm,

MNHN Ci 2606, MNRJ 8877.

Remarks

This is one of the most common deep sea spe¬

cies. It occurs world-wide, and there are several

records from the Eastern Atlantic (distribution

map in Young, in press) (Hoek 1883; Gruvel

1902a, 1905, 1920; Nilsson-Cantell 1928).

Arcoscalpellum tritonis (Hoek, 1883)

(Fig. I)

Scalpellum tritonis Hoek, 1883: 122, pl. 5, figs 9-10,

pl. 10, fig. 10; 1884: 4. Pilsbrv 1907a: 34. - Broch

1953: 4.

Arcoscalpellum tritonis Newman N Ross 1971:

60.-Zevina 1978: 1350: 1981:342, fig. 342.

Materiai EXAMINED. — Stn 66, 2 specimens, tl:

1,6-1.7 cm, MNHN Ci 2663. — Stn 251, 1 spéci¬

men, t): 1.4 cm. MNHN Ci 2649.

Remarks

The are no noteworthy différences between these

specimens and those from Siboga and from

Meteor (Hoek 1883; Young in press), except the

relative length and development of the inframe-

dian latus, which is more variable. The speci¬

mens front the Meteor hâve the inframedian latus

a little higher than wide with the height almost

twice width in some specimens.

Arcoscalpellum eponkos n.sp.

(Figs 3-4)

MATERIAL EXAMINED. — Holotype: stn 249, tl:

0.9 cm, MNHN Ci 2643. Paratypes: 2 specimens

from the same locality, tl: 0.4 and 0.6 cm, MNHN

Ci 2669.

ETYMOLOGY. — From the Greek, eponkos (prégnant)

in référencé to the inflated scutum.

Dlagnosis

Plates of capitulum approximate, thin, smooth,

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Fig. 2. — Géographie distribution of Oxynaspis celata group (■) (in the Western Atlantic, O. hirtae Totton and in the Eastern Atlantic,

O. celata Darwin s.s.), Smilium acutum (Hoek) (T) and Altiverruca gibbosa (Hoek) (•).

ZOOSYSTEMA « 1998 « 20(1)

Young P. S.

except for strong longitudinal ridges at basi-late-

ral surface of carino-latus. Scutum conspicuously

inflated at basi-occludent area, with strong tergal

arm. Caudal appendage uniarticulated.

Description

Femdle

Capitulum (Fig. 3A-D) covered by thin hairless

cuticlc; length about twice width, capitulum

laterally compressed except for convex enlarge-

ment in scuta; cannai margin more convex than

occludent margin. Plates thin, smooth, except

for strong longitudinal ridges at basi-lareral area

of carino-latus and thin sparse growth lines.

Plates approximate.

Scutum (Fig. 3A-B) conspicuously inflated in

basi-occludent area, pronounced tergal arm

approximately une quarter length of tergal mar¬

gin, shallow groove ueat the apex ot upper-latus;

length twice width; ail margins convex, except

for concavity at upper portion of latéral margin;

apex recurved, extending over tergum.

Tergum (Fig. ,3A) triangular, apex conspicuously

recurved towards carina; basal margin almost

straight, occludent margin convex; surface area

larger than that of scutum.

Carina (Fig. 3A, C) regularly arched, uniforntly

broad, tectum fiat, basal portion triangular.

Upper-latus (Fig. 3A) quadrangular; scutal mar¬

gin longer than tergal; scutal margin concave,

others almost straight; carino-lateral more than

twice width of carinal margin.

Carino-latus (Fig. 3A, C) pentagonal, height

more than twice width, umbo near base of cari¬

na, projectlng slightly beyond carinal outline;

basi-lateral area with strong radial ridges. Plates

contiguous under carina, interdigitating,

Inframedian lattis (Fig. 3A) triangular, not pro-

jecting, apex at same height as upper-latus

height; height three rimes greatcr than width,

bordered by iow elevated ridge.

Rostro-latus (Fig. 3A, D) higher than wide; basal

and scutal margins diverging, latter larger, with

slight apico-basal ridge separating plate into two

triangular areas.

Rostrum (Fig. 3D) elongated, almost two third

length of rostro-latus margin, thin, rounded at

top.

Peduncle (Fig. 3A) short, length approximately

one fifth that of capitulum, covered by conspi-

cuous laterally elongated scales.

Labrum (Fig. 3E) with single row of thirty-six

sm;dl teeth. PaJp (Fig. 3F) small, distally acumi-

nate, with sparse setae on al! surfaces. Mandible

(Fig. 3F) with three teeth, lower angle denticula-

ted; distance between First and second tooch two

third that between second and third. Maxilla I

(Fig. 3G) with two steps, lower one projecting;

upper angle with two large and two smaller

spines, eigltt intermediate small spines directed

basally along cutting border Lielow. Maxilla II

(Fig. 3H) bdobed, with simple setae along mar¬

gins; rnaxiilary gland not projecting.

Cirrus 1 (Fig. 31) with anterior ramus 0.70 length

of poscenor one; lesser ramus with articles slightly

protubérant, clothed with numerous, simple

setae. Cirri 11-VI with subequal to equal rami;

intermediate anicles ot cirrus VI (Fig. 3J)

2.5 cimes as long as wide, armed with three pairs

■of simple setae and scartered small setae on ante-

rlor margin, one to two sérac on posterior angle.

Caudal appendage (Fig. 3K) uniarticulated,

short, about one fifth height of coxopodite, with

few simple setae at apex. Number of articles of

cirri J-Vl and caudal appendage i.s presented in

table 1,

Table 1. — Number of articles of cirri l-Vl, and caudal appenda¬

ge ol Arcoscalpellum eponkos n.sp. Holotype, stn 249. I-VI,

cirri I to VI, CA. caudal appendage; RC, right cirri; LC, left cirri;

+. broken ramus.

III

7/9

15/16

18/18

19+/20

23/21

21/20

6/10

15/17

18/19

21/21

20/21

22/22

Remarks

Arcoscalpellum eponkos n.sp. is a small species, the

largest specimen reaching 9 mm in overall

height. The characteristic inflated scutum with a

tergal arm, longitudinal ridges restricted to the

basi-lateral surface of the cari no-latéral, and the

uniarticulated caudal appendage distinguish this

species front ail other species of Arcoscalpellum

s.str. Small spccimens do not hâve the inflated

scutum, but the other characteristics are already

well developed.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

The lengthened inframedian latus and the high

carino-latus suggests that this species is closely

related to A. mdidtum Rao et Newman, A. gryl-

lum Zevina, A. cornpositum (Zevina), A. galupa-

ganum (Pilsbry) and A. sculptum (Hoek).

However, nonê of these species hâve a tergal arm

on the scutum, except for the A. sculptum , which

has a poorly developed arm. A. sculptum also has

strongly ridged plates.

Arcoscatpellum eponkos n.sp, is known only from

the type-locality, West European Basin, between

depths of 4620-4690m (Fig. 4).

Genus P ta noscalpellu m Zevina, 1978

Planoscalpellurn limpidus (Zevina, 1976)

Scalpellum limpidus Zevina, 1976: 1152, fig. 2.

Planosçahellujn limpidus — Zevina 1978: 1347; 1981:

187, fig. 131; 1993a: 125.

Material EXAM1NED. — Stn 245, 1 specimen, tl:

1.0 cm, MNHN Ci 2640. — Stn 249, 1 specimen, tl:

1.0 cm, MNHN Ci 2645.

Remarks

Planoscalpellurn limpidus occurs at depths between

5001-5580 m. It was recorded originally from the

Azores (Zevina 1976) and later in the Antarctic

(Zevina 1993a) and Iberian Basin (Young in

press). The présent new records extend the nor-

thern distribution to 45"50’N and indicate a shal-

lower depth of 4270 m. P. limpidus is redescribed

and discussed in detail by Young (in press).

Genus Catherinum Zevina, 1978

Catherinum recurvitergum (Gruvel, 1900a)

(Figs 4-5)

Scalpellum recurvitergum Gruvel, 1900a: 190; 1902a:

67, pl. 2, figs 3h, 21-22; 1902c: 523; 1905: 49,

fig. 54. — Welcner 1922: 72. - Nilsson-Cantell 1938:

8; 1955:218. - Zevina 1976: 1155.

Catherinum recurvitergum — Zevina 1978: 1348;

1981: 245, fig. 181.

Material examined. — Stn 202, 1 specimen and

1 scutum, tl: 1.6 cm, MNHN Ci 2616. — Srn 245,

3 specimens, tl: 1.1-1.3 cm, MNHN Ci 2641, MNR]

8878.

Description

Pemale

Capitulum (Fig. 5A-C) covered by thin hairless

cuticle; length twice its width, width uniform;

cannai and occludent margins convex, with same

curvature. Plates with thin, sparse growth lines,

with longitudinal ridges variously developed,

ridgcs absent in specimens up to 10 mm in

length, slightly elevated in specimens of about

13 mm, conspicuous and strong in specimens of

15 mm. Plates approxintate, cuticle scparating

carina from other plates.

Scutum (Fig. 5A) with occludcnt margin convex,

with shallovv groovc near apex of upper-latus;

scutal length twice width; tergal arm small, about

one fîtrh length of tergal margin; tergal margin

et]liai to length of basal margin, with small upper

rim; carinal and occludent margins convex,

except for concavity at upper portion of former;

apex cutved, extendirig slightly over tergum.

Tergum (Fig. 5A) triangular, with longitudinal

ridges more developed at occludent area; basal

margin aJmost straighr, occludent margin

convex. carinal irregnlar; apex recurved in small

specimens, acute or eroded and obtuse in large

specimens, slightly recurved toward carina; surface

area ol tergum equal to that of scutum.

Carina (Fig. 5A, B) regularly arclied, broaderapi-

calv, tectum slightly concave, bordered by two

low longitudinal ribs, basa! portion rounded.

Upper-latus (Fig. 5 A) pentagonal. Length of scu¬

tal margin greater titan tergal one, former conca¬

ve. AU other margins essenttally stratght,

carino-lateral lortgest, followed in size by tergal

and inframedian margins. Apex with small upper

rim.

Rostro-latus (Fig. 5A, C) wider than high; basal

and scutal margins diverging, scutal margin grea-

rer in length than basal margin, apico-basal ridge

separating plate into two triangular surfaces.

Carino-latus (Fig. 5A, B) pentagonal, higher

than wide, untho near base of carina, umbo pro-

jecLirtg slightly beyond carinal margin. Plates

contiguous under carina.

ZOOSYSTEMA • 1998 ■ 20(1)

Cirripedia from the Azores Région

30° 20° 10°

Fig. 4. — Géographie distribution of Arcoscalpellum eponkos n.sp. (▼). Calherinum recurvitergum (Gruvel) (•) and Teloscalpellum

tuteum (Gruvel) (■).

Inframedian latus (Fig. 5A) vase-shaped, umbo

subcentral, apical portion higher and wider than

basal portion.

Rostrum (Fig. 5C) elongate; length almost two

third that of rostro-latns margin; thin, apically

rounded,

Peduncle (Fig. 5A) about one quarter length of

capitulum, covered by conspicuous lengthened

scales.

Labrum (Fig. 5D) with row ot forry-four small

teeth. Palp (Fig. 5D) small, paddle-like, with

setae on ail surfaces.

Mandible (Fig. 5E) with three teeth, lower angle

denticulated; distance between ftrst and second

teeth less than lwice that between second and

third one.

Maxilla 1 (Fig. 5F) with .straight border, two large

and thick spines at upper angle, nine rnoderate

to small spines difected downwards below.

Maxilla II (Fig. 5G) triangular with finely pinnate

setae along margins; maxillary gland projecting

prominently.

Cirrus 1 (Fig. 5H) with shorter ramus 0,75 length

of longer; articles little protubérant, clorhed with

numerous simple setae.

Cirri Il-VT with subequal to equal rami, interme-

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

diate articles of cirrus VI (Fig. 51, J) twice as long

as wide, vvith three or four pairs of simple setae

and scattered small setae on anterior niargin,

pairs of setae more developed along the posterior

ramus; one ro two setae on posterior angle.

Caudal appendage (Fig 5K) mulriarticulated

(7-8), almost rwice length of protopod, simple

setae at distal point, a tew setae on distal margins

of articles.

Number of articles of cïrri I-VI and caudal

appendage is presented in table 2.

Table 2. — Numbet of articles of cirri l-VI, and caudal appenda¬

ge of Catherinum recurvitergum (Gruvel), stn 202. I-VI, cirri I to

VI; CA, caudal appendage; RC, rlght cirri; LC, left cirri; +, broken

ramus.

III IV

8/10

17/19

14+/15+ 22/14+

22/23

23/16+

8/10

16/16+

20/22 12+/23

15+/22

23/23

Remarks

The specimen described by Gruvel (1902a) is

11 mm in capirulum length and was illustrated

withour longitudinal ridges. Howcvcr, in his des¬

cription Gruvel stated that the plates had conspi-

cuous radial lines.

The présent specintens examined range from 10

to 16 mm in capirulum length, and it is apparent

that the development of the longitudinal ridges

increases during ontogeny.

The smallest specimen (10 mm) has only a few

longitudinal ridges on the plates and the latgest

(16 mm) has such ridges on ail plates except for

the carina (Fig. 5A). Otherwise, the apex of the

recurved tergum uoted by Gruvel ( 1902a) is not

as recurved in the specimens examined.

Catherinum recurvitergum was originally descri¬

bed from rhe Azores from 3175 m (Gruvel

1900a; 1902a) and subsequently only recorded

from eastern Africa from 1289 m (Weltner

1922). Weltner did not describe or figure his

specimens. The specimens sampled are from the

type localiry, Azores région (Fig. 4), between

2900 and 4270 m, which extends the depth

range of this species.

Gcn us Amigdoscalpellum Zevina, 1978

Amigdoscalpellum rigidum (Aurivillius, 1898)

Scalpe.llum rigidlim Aurivillius, 1898: 189. — Gruvel

1905; 86.-Zevina 1976: 1155.

Scalpellum striatum Gruvel, 1900a: 191; 1902a: 77,

pi. 2. fig. 31; 1905: 72, fig. 81; 1920: 23, pl. 2,

figs 4-6, 9-11; pl. 7, fig. 11. — Nilsson-Cantell 1955:

219. - Belle,c 1959:3.

Asnigdvscalpe!litm rigidum - Zevina 1978: 1349;

1981: 277, fig. 209.

MATËRMl- rXAM.INIT). — Stn 174, 2 specimens, tl: 2.0

and 2.1 cm, MNHN Ci 2590. — Stn 176, 1 speci¬

men, tl: 2.7 cm, MNIIN Ci 2592. — Stn 202,

17 specimens, tl: 1.5-3.7 cm. MNIIN Ci 2615,

MNRJ 8879. — Stn 206, I specimen, tl: 2.5 cm,

MNHN Ci 261 8. — Stn 227, 1 specimen, il: 4.0 cm,

MNHN Ci 2625. — Stn 245, 1 specimen, tl: 2.6 cm,

MNHN Ci 2637. — Stn 251, 2 specimens, tl:

2.1-2.4 cm, MNHN Ci 2647.

Remarks

Amigdoscalpellum rigidum appears to be a com-

nton deep-sea species from rhe Azores, Cape

Verde, and the Iberian Basin, occurring between

depths of 1267-4400 m (Aurivillius 1898;

Gruvel 1905; Young in press), lt was also recor¬

ded once from the Newfoundland Basin, from

1267 m (Aurivillius 1898). The new samples

extern! its distribution to the West European

Basin. A. rigidum is redeseribed and discussed by

Young (in press).

Genus Trianguloscalpellum Zevina, 1978

Trianguloscalpellum ovale (Hoek, 1883)

Scalpellum regium var. ovale Hoek, 1883: 109, pl. 5,

figs 5-6.

Trianguloscalpellum regium ovale — Zevina 1981: 311,

fig. 235.

MATERIAL examined. — Stn 249, 2 specimens, tl: 3.3

and 4.9 cm, MNHN Ci 2644.

Remarks

T. ovale is redeseribed and discussed in detail by

Young (in press) who présents a key for the rela-

ted species. This new sample does not add any

ZOOSYSTEMA • 1998 • 20(1)

information about the North Atlantic deep sea

distribution of this species.

Trianguloscalpellum regium (Thomson, 1873)

Scalpellum regium Thomson, 1873: 347 (part): 1877:

4 (part). - Hoek 1883: 106, pl. 4, figs 3-6, pl. 9,

fig. 12, pl. 10, figs 1-2; 1884: 10. - Pilsbry 1907a: 28,

pl. 3, fig. 5 (part). - Gruvel 1912a: 2; 1920: 30, pl. 1,

fig. 7.

Scalpellum molle Aurivillius, 1898: 191. — Gruvel

1905: 76; 1920:29, pl. 5, figs 10-12. - Nilsson-

Cantell 1955: 218.-Zevina 1976: 1 155; 1981:309.

Trianguloscalpellum regium — Zevina 1978: 1350.

- Foster &: Buckeridgc 1995a: 167, figs 2A-D, 3.

Ir tango hncalpellum regium regium — Zevina 1981:

309, fig. 234.

Materiai EXAMINED. — Stn 129, 3 specimens, tl:

1.3-4.3 cm, MNHN Ci 2566. — Stn 202, 27 speci¬

mens, tl: up to 6.4 cm, MNHN Ci 2617, MNRJ

8880. — Stn 245, 9 specimens, cl: 0.4-7.0 cm,

MNHN Ci 2638. — Stn 251, 3 specimens, tl:

1.7-5.4 cm, MNHN Ci 2648.

Remarks

In the smaller specimens the upper flange of the

carino-latus does not appear to be as well develo-

ped as in the larger specimens described by

Young (in press). The roof of the carina changes

in shapc front fiat with a central élévation in

smaller specimens to fiat with latéral ridges in

larger ones.

T. regium is redescribed in detail and discussed

by Young (in press). The présent samples do not

add new information regarding its North

Atlantic dcep-sea distribution.

Genus Teloscalpellum Zevina, 1978

Teloscalpellum luteum (Gruvel, 1900a)

(Figs 4, 6)

Scalpellum luteum Gruvel, 1900a: 192; 1902a: 80,

pl. 2, fig. 11 ; 1905; 84, fig. 93. - Nilsson-Cantell

1955:218. - Zevina 1976: 1155.

Teloscalpellum luteum — Zevina 1978: 1350; 1981:

363, fig. 280.

Material EXAMINED. — Stn 129, 1 specimen, tl:

3.0 cm, MNHN Ci 2567.

Description

Female

Capitulum (Fig. 6A, B) covered by thin hairless

curicle; length less than twice its width, breadth

tiniform; carinal margin more convex than occlu-

dent. Plates with strong longitudinal ridges and

thin, sparse growrh lines. Narrow curicle separa-

ring mosr of plates.

Scutum (Fig. 6A) convex oniy at occludent area;

height less than twice width; latéral margin lon¬

ger than basal, sinuous; basal and occludent mar¬

gins convex, tergal slightly concave; apex curved,

extendmg over tergum.

Tergum (Fig. 6A) triangular, with longitudinal

ridges developed only ai occludent side; basal

margin almost straight, occludent margin

convex, carinal with depressed area near the cari¬

nal apex; apex obtuse, slightly turned roward

carina; surface area greater than scutum.

Carina (Fig. 6A) regularlv arched, rectum concave,

bordered by high longirudinal rib, basal portion

absent in specimen, therefore I could not observe

its shape.

Upper-larus (Fig. 6A) pentagonal. Scutal and ter¬

gal margins equal in length, former concave. Ail

other margins almost straigbc, carino-lateral lon-

gest. followed in si/.e by rergal and infraniedian

margins. Umbo subapical. Apex rhickened, for-

ming upper ridged rim.

Rostro-latus (Fig. 6A, B) wider than high; basal

and scutal margins parallel, latter longer; apices

of plates overlapping.

Carino-latus (Fig. 6A) pentagonal, higher than

wide, umbo near base of carina, not projecting

from carinal oudine. Plates contiguous under

carina.

Inframedian latus (Fig. 6A) triangular, length

twice its width, apex slightly turned anteriorly,

with rim along carinal side.

Rostrum (Fig. 6B) absent.

Peduncle (Fig. 6A) about onc hall length of capi¬

tulum, covered by conspicuous lengthened scales

and thick curicle.

Labrum (Fig. 6C, D) with row of small teeth.

Palp (Fig. 6C) small, acuminate, with few setae

on upper margin and terminally. Mandible

(Fig. 6£, F) with three teeth, with or without

large tooth berween first and second teeth, denti-

culated lower angle with few teeth; distance bet-

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

ween first and second teeth nvice disrance bet-

ween second and rhird one; serulation absent,

only small seules on lower margin. Maxilla I

(Fig. 6G) witb straight anterior border or upper

portion projccting slightly, with two large and

thick spines at upper angle, nitie to rwelve mode-

rate to small spines directed basally below.

Maxilla II (Fig. 6H) nearly quadrangular with

finely pinnate setae along margins; maxillary

gland projccting prominently.

Cirrus I (Fig. 61) with anterior ramus 0.65 of

length of posterior one; articles of shorter ramus

protubérant, both rami clorhed with numerous

simple setae. Cirri II-VI with equal rami, inter¬

médiare articles (Fig. 6J) about twice as long as

wide, witb threc or four pairs of simple setae and

scattered small setae on anterior margin, one to

two setae on posterior angle, one or two simple

setae on posterior margin, Caudal appendage

(Fig. 6K) multiarticulatcd (R), slightly longer

than protopod, with simple setae at distal point,

few setae on distal margins of articles. Number

of articles of cirri 1-VI and caudal appendage is

presented in table 3.

Table 3. — Number of articles of cirri l-VI, and caudal appenda¬

ge of Teloscalpellum luteum (Gruvel), stn 129. I VI, cirri I to VI;

CA, caudal appendage; RC, right cirri; LC, left cirri; +. broken

ramus.

III

9/10

12+/20

22/22

28/30

30/31

33/28

9/10

17/19

24/25

28/31

28/33

34/29

Remarks

Teloscalpellum luteum (Gruvel) is known only by

one spécimen, câpitular length 20 mm, front the

type locâiity — Azotes - at 3175 m (Gruvel

1900a, 1902a). Gruvel did not illustrate the lon¬

gitudinal ridges of the plates, but did describe

this character. These longitudinal ridges are

conspicuous in (he specimen studied.

The general shape of the capitular plates, espe-

cially the coronate upper-latus, agréé with the

description of Gruvel (1902a) The basal portion

of the carina, where ir meets the carinal latera, is

absent in rite specimen and, therefore, 1 could

not observe the relationship of the carinal-latera.

T. luteum was collected in the same area as the

type specimens (Azores), and from a similar

depth (3056-3000 m; Fig. 4).

Order SESSILIA Lamarck, 1818

Suborder V F R R U C O M O R P H A Pilsbry, 1916

Family VERRUCIDAE Darwin, 1854

Genus Altiverruca Pilsbry, 1916

Altivemica obliqua (Hoek, 1883)

(Figs 7-9)

Verruca obliqua Hock, 1883; 143, pl. 12,

figs 15-17. - Weltner 1897: 274. - Gruvel 1905: 173,

fig. 191.-Hoek 1907b: 9.

Verruca obliqua , section D - Altiverruca - Pilsbry

1916:40.

Altiverruca obliqua - Buckeridge 1994: 93.

Altiverruca vertka — Foster & Buckeridge 1995a: 180,

«g. 14.

MaTF.RIAI. KXAMINED. — Stn 180, 1 specimen

without eggs or larvae, rc: 0.6 cm, MNHN Ci 2595.

Description

Shell (Fig. 7A-C) white, smooth, growth Fines

widely spaced, not projccting. Opetcular valves

angularly placed, approximately 45" to basis

(Fig. 7D), Base calcareous, Rostrum and carina

(Fig, 7C) convex with apïces projecting, latter

higher; suture sinuose. Fixed-tergum (Fig. 7B)

trianguiar, wider in middle, higher than

fixed'Scutum, with two well developed alate pro¬

jections, little covered by radii-like projections of

adjoining plates. Fixed-scutum (Fig, 7B) triangu-

lar, with well developed alate projection on to

rostrum, covered by small radius-like projection

of this plate; internally smooth.

Scutum smaller than tergum, basal margin one

hall length ol tergum margin. Scutum (Fig. 7E)

with two articular ridges, and fiat upper triangu¬

iar projection at occiudent margin; axial ridge

barcly developed and curved; rostral area smooth.

Internally, with deep dépréssion for adductor

muscle; occiudent margin with small tooth at

lower portion lormed by second articular ridge.

Tergum (Fig. 7F) quadrangular, with rliree arti¬

cular ridges; axial ridge narrowest, intermédiare

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

ridge broad, separated from upper ridge by

conspicuous groovc. Int er ri al ly, surface slightly

concave; occludcnc margin nearly straight, cxcept

for upper convex pordon.

Labrum (Fig. SA) witb one sériés of forry teerh.

Palp (Fig. 8A) long, few simple setae at upper

margin and distal area. Mandible (Fig. 8B) with

three teeth, distance between the ftrst and second

one twice that between the second and third one;

lower angle denticulatcd. Maxilla I (Fig. 8C, D)

straight or witb the lower part projecting, with

10 to 13 unpaired spines. Maxilla II triangülar,

anterior margin concave medially, posterior mar¬

gin convex; covcred by long simple setae.

E - F

Fig. 7. — Alliverruca obliqua (Hoek). A, top view; B, carino-rostral view; C, fixed-tergum and fixed-scutum view; D, fixed-scutum and

rostral view; E, F, tergum and scutum, internai view. Scale bars: 2 mm.

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Cirrus I (Fig. 8E) wirh anrerior ramus a little

longer than posterior, articles of boch rami cover-

ed by nuraerous long, simple setae. Cirrus II

(Fig. 8F) wirh unequal rami, anterior rwo third

length of posterior, wirh protubérant articles;

anterior ramus covered by numerous long,

simple setae. posrerior wirh setation similar to

articles of cirri III-VI. Cirrus 111 with subequal

rami, cirri 1V-V1 with equal rami. Length of

intermédiare article of cirrus VI (Fig, SG) twicè

width, with two pairs ol simple setae on anterior

margrn, one or iwo short, stout setae on poste¬

rior angle, mulrifid scalcs at distal margins.

Caudal appendage (Fig. 8H) multiarriculated,

with six to seven articles, onc half length ofcoxo-

podite. Pénis (Fig. 811) short with few setulae.

Number of articles of cirri I-VI and caudal

appendage is presented in table 4.

Table 4. — Number of articles of cirri l-VI, and caudal appenda¬

ge of AHiverruca obliqua (Hoek), stn 180. I VI, cirri I to VI; CA,

caudal appendage; RC, right cirri; LC, leftcirril.

lit

12/10

9/11

15/16

16/16

17/18

20/20

10/10

8/9

13/13

17/17

17/19

19/20

Remarks

Hoek (1883) described A. obliqua based on four

small specimens collected off southwestern Spain

at a depth of 2782 meters. Tins species has not

been collected since.

This species was recently considered a synonym

of A. quadningulans (Hoek, 1883) by Foster &

Buckeridge (1995b), but this conclusion seems

to be prématuré. Altiverruca quadrangularis , First

reported from the Southern Atlantic, is now

30 J 20° 10°

Fig. 9. — Géographie distribution of Altiverruca obliqua (Hoek) (▼), Metaverruca aequalis (Aurivillius) (•) and Metaverruca trisulcata

(Gruvel) (■).

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

known from -ail the océans (Lahille 1910; Gruvel

1920; Zevina 1988; Rosell 1989; Foster &

Buckeridge 1995b).

Considering tlie original descriptions of borh

species, several characrers distinguish lhem. Both

hâve a terguni with three arricular ridges, but

A. obliqua lias the jntermediate ridge (second ot

Hoek) broad. leaving a thin groove between rhis

ridge and rhe first one. Alternative!}', A. quadran-

gularis has the second ridge narrow, with a very

broad space separating it from the first ridge.

The scutum of A. obliqua is proportionately nar-

rower, its base one half the width of the base of

tergum. In A. quadrangulam the movable scu¬

tum is sigtiificanrly larger, with its width more

than two ihird of the width of the base of the

tergum. The axial ridge (third ridge of Hoek) is

promineni in A. qttadrangularis, but it is barely

discernihle in A. obliqua. Therefore, this synony-

my cannot be accepted,

The single specimen I examined is older and

hence more developed than the specimens Hoek

studied. Ail of rhe différences I observed appear

to be related to different stages of growth: the

carina projects freely as does rhe scutum; the arti¬

culation between carina and rostrum has more

than one single tooth.

Foster & Buckeridge (1995a) described

Altiverruca vertied from the Straits ot Gibraltar.

1rs opercular valves are identical to chose exami¬

ned here, differing only in the carina-rostrum

suture. This suture is straighter than the one

observed, but this may be due to on toge net ic dif¬

férences. FurLhcrmore, the relative length of the

rami of cirri I-UI and rhe caudal appendages are

similar to the unes 1 observed. Hoek (1883) did

not dcscribe the appendages of A. obliqua.

The distribution ol this species is restricted to

the Northeastern Atlantic, between 34° to 37°N

and 7° to 25°W (Fig. 9).

Altiverruca gibbosa (Hoek, 1883)

(Figs 2, 10)

Verruca gibbosa Hoek, 1883: 134, pl. 6, figs 17, 18,

pl. 11, figs 5-9, pl. 12, figs 1-5.

Verruca ( Altiverruca ) gibbosa - Newman & Ross

1971: 135, pl. 14, textfig. 68 (synonymy).

Altiverruca gibbosa - Zevina 1987a: 1813.

MatKRIAJ. EXAMINED. — Stn 6, 1 empty shell, rc:

0.3 cm, MNHN Ci 2550. -- Stn 159, 1 specimen,

rc: 0.4 cm, MNHN Ci 2584. — Stn 180, 2 spéci¬

mens, rc: 0.4-0.6 cm, MNHN Ci 2596, MNRJ

8903. — Srn 196, 1 specimen, rc: 0.5 cm, MNHN

Ci 2603. — Stn 199, 1 specimen, rc: 0.5 cm,

MNHN Ci 2610.

Remarks

The specimens studied (Fig. 10A, B) do not pré¬

sent any différences from the specimens descri¬

bed by 1 loek ( 1883).

A. gibbosa has a world-wide distribution and was

previously recorded in the Eastern Atlantic from

off the Straits of Gibraltar, Canaries and Cape

Verde Islands and Sierra Leone Rise (Fig. 2)

(Gruvel 1902a, as V. sulcata; Nilsson-Cantell

1927, as V. rathbuniana\ Nilsson-Cantell 1928;

Foster & Buckeridge 1995a).

Fig. 10. — Altiverruca gibbosa (Hoek). A, top view; B, fixed-

tergum and fixed-scutum view. Scale bar: 1 mm.

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Genus Metaverruca Pilsbry, 1916

Metaverruca aeqiuilis (Aurivillius, 1898)

(Figs 9, 11-12)

Verruca aequalis Aurivillius, 1898: 196, - Gruvel

1905, 176; 1920: 42, pl. 5, figs 28-29, pl. 6,

figs 6-7. - Hock 1907b: 9. - Belloc 1959: 4.

MATER1AL EXAMINE!). — Stn 171, I specimen, rc:

0.5 crn, MNHN Ci 2589. — Stn 1 7 4, 1 specimen,

rc: 0.4 cm, MN1JN Ci 2591. — Stn 180, 4 spéci¬

mens, rc: 0.5-0.6 cm, MNHN Ci 2598, MNRJ

8904. — Stn 196, 3 specimens, rc: 0.5-0.6 cm,

MNHN Ci 2605, MNRJ 8905.

Description

Shell (Fig, 1 lA, B) white, nearly smooth, grooves

only along the suture between rostrum and can¬

na; growth lines widely spaced, projecting; basal

margins thickened, inflected (Fig. I IC). Cuticle

hairy, persistent on shell and opercular valves.

Opercular valves parallel to base of wall.

Rostrum (Fig. 1 IA) larger than carina, vvith small

radius-like projection toward fixed scutum; arti¬

culation vvith carina with large tippcr ridge and

unduiating downwards; shallow grooves beside

ridge; apex projecting. Carina (Fig. 11 A) with

small radius-like projection roward frxed-tergum,

onc upper ridge unduiating downwards at rostral

suture, one groove under the ridge: apex reflexed.

Fixed-tergum (Fig. 11 A, B) smailer than

fixed-rostrum, approximately one half its widtb,

both s ides with well developed alar-like projec¬

tions; apex projecting backwards. Fixcd-scutum

(Fig. 11 A, B) with wide alar-like projection direc-

ted toward rostrum, small radii-like projection to

fixed-tergum; internally, with well developed

myophore, directed downwards.

Tergum (Fig. 11A, D) quadrangular, with three

Fig. 11, Metaverruca aequalis (Aurivillius). A, top view; B, fixed-tergum and fixed-scutum view; C, basal view D E tergum and

scutum, internai view. Scale bars: 2 mm.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

ardcular ridges, axial rjdge narrowest, well mar¬

ie e cl, intermédiare and upper ridges broad.

InternaJly, smooth; occludent margin slightly

undulated. Scutum (Fig. 1 IA, E) smaller than

tergum; with three articular ridges; axial ridge

barely developed, near occludent margin, inter¬

médiare ridge twice width of upper ridge, latter

poorly developed; rostral area smooth. Internally,

with conspicuous rounded scar for adductoi

muscle, espedaüy near articulât margin, forming

a vertical crest between them; occludent margin

nearly straight, except for a protubérance at

lower part.

Labrum (Fig. 1 2A) with sériés of small teeth

(47). Palp (Fig. 12B) short, acuminate, with few

simple setae on upper margin and distal région.

Mandible (Fig. 12C) with three teeth, distance

between first and second more than twice distan¬

ce between the second and third one; lower angle

denciculated. Maxilla I (Fig. 12D) with lower

part proJecting, shallow concavity apically; two

large spines at upper angle, four small spines in

concavity, seven to nine unpaired spines on lower

portion. Maxilla II (Fig. 12E) rectangular, ante-

rior margin with shallow concavity medially;

covered by long simple setae.

Cirrus T (Fig. 12F) with subequal rami, covered

with several long simple setae. Cirrus II

(Fig. !2G) with anterior ramus two third length

of posterior, articles more protubérant; both rami

covered by numerous long, simple setae, anterior

ramus also with finely bipectinate setae at distal

article (Fig. 12H). Rami of cirrus 111 unequal,

anterior shorrer than posterior. Rami of

cirri IV-VI equal in length. Intermédiare articles

of cirrus VI (Fig. 121) with widdt three quarters

of the length, two pairs of setae, longer setae

finely pinnate, on anterior margin; one or two

short, stout setae on posterior angle, mulrifid

scales at distal margin. Caudal appendage

Table 5. — Number of articles of cirri 1-111, and caudal appenda¬

ge of Metaverruca aequalis (Aurivillius), stn 195. I-III, cirri I to III;

CA, caudal appendage; RC. right cirri: LC, left cirri.

III

10/9

8/10

14/16

10/9

8/10

14/16

(Fig. 12]) with six articles, one half length of

coxopodite; long simple setae on distal margins

of articles. Pénis about same length as coxopo¬

dite, clotlicd with thin setulae. Number of

articles of cirri I-III and caudal appendage is pre-

sented in table 5.

Remarks

Metaverruca aequalis was described briefly by

Aurivillius (1898), but was subsequently descri¬

bed in detail and figured by Gruvel (1920),

based on the same material. The specimens exa-

mined herein agréé with the description by

Gruvel (1920). I eau only add that the hairy

cuticle is conspicuous in smaller specimens.

Gruvel (1920) did not dissect any specimen.

Therefore, he did not describe the internai mor-

phology of the shell, the opercular valves, and

the appendages. The présence of a developed

myophore on the fixed-scutum, the box-like

shape of the wall with îts inflecred basal tnargins

and the large opercular valves, indicate that the

species is a Metaverruca.

Metaverruca aequalis is known only frotn the

Azotes, between deprh of 1022 and 1385 m. Ail

the samples studied are frotn the same area, but

its depth distribution is increascd to 3215 m

(Fig. 9).

Metaverruca recta (Aurivillius, 1898)

Verruca recta Aurivillius, 1898; 195. — Gruvel 1905:

181; 1912a: 6; 1920: 46, pi. 2, fig. 18, pl. 3,

figs 3-4. - Hoek 1907b; 9. - Southward &

Southward 1958: 637, fig. 4. - Anderson 1980: 349,

figs 1-4.

Verruca sculpta Aurivillius, 1898: 197. - Gruvel,

1905: 175; 1920: 41, pl. 5. figs 26-27. - Hoek

1907b: 9. — Nilsson-Cantell 1929: 461, fig. 1; 1938:

12. - Kriiger 1940: 463. - Zevina 1969:

68. - Weisbord 1979: 97. - Foster 1981: 352. - Ren

1984: 166, fig. 1, pl. 1, figs 1-6; 1989: 420, fig. 10.

Verruca linearis Gruvel, 1900b: 243; 1902a: 107.

pl. 5, figs 11-12; 1905: 182. fig. 201. -Hoek 1907b:

Verruca capsula Hoek, 1907a: 130, pl. 12, figs 1-3,

pl. 13, figs 1-4. - Stubbings 1936: 38. - Weisbord

1979: 98.

Verruca marna Gruvel, 1901: 261; 1902a: 109, pl. 5,

figs 1-2; 1905: 184, figs 204-205. - Hoek 1907b:

9. - Gruvel 1920: 50. - Weisbord 1979: 98.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Verruca balotheca Pilsbry, 1907b: 188, pl. 4, figs 9-10;

1916: 46, pl. 8, figs 1-la. - Kolosvàrv 1943:

73. - Zullo 1968: 219. - Gordon 1970:

118. - Buckeridge 1975: 129. figs 5. 4-6. - Foster

1978: 69, pl. 8F. fig. 42. - Welsbord 1979: 98.

Verruca coraliopbila Pilsbry. 1916: 21, pl. 1,

figs 1-5. - Zullo 1968: 219. - Bayer et al. 1970:

A43. - Weisbord 1979:96.

Verruca ( Metaverruca) sculpta - Broch 1931:

4L - Buckeridge 1983: 59, fig. 45.

Verruca {Metaverruca) eookei - Rosell 1981: 299,

pl. 11, figs r, s, u, v; 1991: 33 (not Verruca eookei

Pilsbry, 1927).

Metaverruca recta - Buckeridge 1994: 116,

fig. 13a-f. - Foster & Buckeridge 1995a: 182, fig. 15;

1995b: 368, fig. 9C-E.

Matériel EXAMrNED. — Srn 4, 8 specimens and

10 empty shells, rc; 0.6-0.9 cm, MNFIN Ci 2549. —

Stn 16, 1 specimen, rc: 0.6 cm, MNHN Ci 2555. —

Stn 62, 3 empty shells, rc: 0.6-0.7 cm, MNHN

Ci 2560. — Stn 66, 1 empty shell, rc: 0.8 cm,

MNHN Ci 2561. — Stn 135, 1 specimen, rc:

0.7 cm, MNHN Ci 2570. — Stn 139, 1 empty shell,

rc: 0.7, MNHN Ci 2571. — Stn 148, 101 specimens

and 12 empty shelL, rc: 0.5-0.9 cm, MNHN

Ci 2576, MNRJ 8906. — Stn 150, 2 specimens and

1 empty shell, rc: 0.6-0.6 cm, MNHN Ci 2578. —

Stn 151, 27 specimens and 1 empty shell, rc:

0.5-0.9 cm, MNHN Ci 2580, MNRJ 8907. —

Stn 159, 1 specimen, rc: 0.3 cm, MNHN

Ci 2585, —Stn 161, 1 specimen, rc: 0.7 cm,

MNHN Ci 2664. — Stn 168, 9 specimens and

4 empty shells, rc; 0.5-0.8 cm, MNHN Ci 2588. —

Stn 180, 19 specimens and 5 empty shells, rc:

0.6-1.2 cm. MNHN Ci 2597, MNRJ 8908. —

Stn 181, 4 specimens and ! empty shell, rc:

0.6-0.7 cm, MNFIN Ci 2600. — Stn 196, 10 speci¬

mens and 18 empty shells, rc: 0.6-1.0 cm, MNHN

Ci 2604. — Stn 197, rc: 0.5-0.8 cm, MNHN

Ci 2607. — Stn 199, I specimen, rc: 0.4 cm,

MNHN Ci 2611. — Stn 218, 2 specimens and

1 empty shell, rr; 0.6-0.7 cm, MNFIN Ci 26 22. —

Stn 231, 1 empty shell, le: 0.7 cm. MNHN

Ci 2626. — Stn 237, 1 specimen and 5 empty shells,

rc: 0.6-0.7 cm, MNHN Ci 2629. — Stn 239.10 spe¬

cimens and 4 empty shells, rc: 0.6-0.9, MNHN

Ci 2632. — Srn 240, 78 specimens and 10 empty

shells, rc: 0.5-0.8 cm, MNHN Çi 2634, MNRJ

8909. — Stn 255, 2 specimens, rc: 0.9-1.1 cm,

MNHN Ci 2650.

Remarks

Metaverruca recta is the most common deep-sea

verrucid found in the Azores région. It was col-

lected from over several distinct substrates, such

as pumice stones, shells, corals, and urchin

spines. It has a world-wide distribution, and was

recorded many rimes from the Nortbeastern

Atlantic, including the Azotes from 240 to

2100 m (Auriviltius 1898; Grnvel 1912a, 1920;

Southward & Southward 1958; Foster &

Buckeridge 1995a; Young in press).

The specimens studied do not présent significant

différences from those observed by other authors.

The white shell is totalIy smooth, and has a

straight hinge between the opercular valves and

the rostrum and carina, characters which are dia¬

gnostic for this species.

Metaverruca trisulcata (Gruvel, 1900b)

(Figs 9, 13-14)

Verruca trisulcata Gruvel, 1900b: 243; 1902a: 96,

pl. 5, figs 9-10: 1905: 184, fig. 203; 1912b: 348;

1920: 49. - Hoek 1907b: 9.

Verruca striata Gruvel, 1900b: 244; 1902a: 98, pl, 5,

figs 5-6, rextfig. 17-18; 1905: 183, fig. 186,

202. - Hoek 1907b: 9,-Srubbmgs 1967: 251.

not Metaverruca trisulcata - Foster & Buckeridge

1995a: 177, figs 10-12: 1995b: 363. fig. 9a-b.

MATERIAL EXAMINFD, - Stn 197, 2 specimens, rc:

0.5 cm, MNHN Ci 2608, MNRJ 8910. — Stn 240,

1 specimen, rc; 0.4 cm, MNFTN Ci 2635.

Description

Shell (Fig. 13A, B) white, cuticle not persistent,

ornamented with several strong longitudinal

ridges and spaced growth fines, ridges on opercu¬

lar plates and at suture of rostrum and carina

conspicuous, Opercular plates parallcl co basis.

Basal margin ot wall not infleeted.

Fised-rergurn (Fig. 13A, B) smaller than

fixed-scutum, apex thickened, recurved ourward;

with two developed alate projections and trian-

gular central area, with four to five longitudinal

ridges,

Fixed-scutum (Fig. 13A, B) having central trian-

gular area with four to five strong longitudinal

ridges, well developed alate projection ai rostral

suture area, radius-like projection at tixed-

tergum side; apex thickened, recurved outward.

Internally, with well developed myophore.

Carina (Fig. 13A) same size as rostrum, with

about ten strong longitudinal ridges and well

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

developed ridge area at suture with rostrum, with Tergum (Fig. 13A, C) larger than scutum, nearly

four interlocking teeth, the larges! being usually quadrangular, with three articular ridges; axial

the second; smooth suture with fixed-tergum. ridge highest and as wide as second, distance bet-

Rostrum (Fig. 13A) with many strong longitudi- ween ridges equal, groove between First and

nal ridges, well developed ridged area at suture second ridges greater than between second and

with carina, with four interlocking teeth, the first third ones; no ridges at carinal area. Internally,

one being the largest, smooth suture with Ftxed- surface Hat and smooth; scutal margin slightly

scutum. concave, with médian tooth.

Fig. 13. — Metaverruca trisulcata (Gruvel). A, top view; B, fixed-tergum and fixed-scutum view; C, D, tergum and scutum, internai

view. Scale bars: 1 mm.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Scutum (Fig. 13A, D) with three articulât ridges,

third ridgc thin, sloping continuously to rostral

area, no ridges at rostral area; apex pointed,

directed toward tergum. Internally, tergal margin

nearly straighc, wich tooth at lower third; surface

with conspicuous upper dépréssion, including

adducror muscle pit.

Labrum (Fig. 14A) with sériés of irregular teeth.

Palp (Fig 14a) short, acuminated, with few

simple setae on upper margin and distal région,

Mandible (Fig. 14B) with three teeth, distance

between First and second teeth twice distance

between the second and third ones; lower angle

denticulated. Maxilla I (Fig. I4C) with lower

part projected; two large spines and five small

spines on upper edge of cutting edge and thir-

teen moderate to small unpaired spines on lower

part. Second maxilla not observed.

Cirrus 1 (Fig. I4D) with subequal rarni, anterior

slightly longer than posterior one; both with pro¬

tubérant articles covcred by long, simple setae.

Cirrus II (Fig. 14E) with anterior ramus about

two third length of posterior, with articles more

protubérant; both rami covered by long, simple

setae. Cirri III with unequal rami, anterior slior-

ter than posterior; cirri with rami of IV-VI çqual

length. Intermédiare articles oi cirrus VI with

width about two third length, three pairs ol long

setae on anterior margin, two setae on posterior

angle. Caudal appendage (Fig. 14F) multiarticu-

lated, seven articles, one hall length of coxopo-

dite; long simple setae on distal margins of

articles. Number of articles of cirri 1-VI and cau¬

dal appendage is presented in table 6. The spéci¬

mens dissected had approximately fifty eggs each

(from station 197).

Table 6. — Number of articles of ofrri l-VI, and caudal appenda¬

ge of Metaverruca irisulcata (Gruvel), stn 197 l-VI, cirri 1 1 o VI;

CA, caudal appendage: RC, right cirri: LC, left cirri.

III

11/15

8/12

16/19

21/23

23/25

25/26

15/12

8/12

16/19

12+/23

24/24

24/26

Remarks

M. trisulcata was described by Gruvel (1900b,

1902a, 1920) from the Azores, and it was later

recorded from the coast of Morocco (Gruvel

1912b). Foster & Buckeridge (1995a, b) recor¬

ded it from off the Straits of Gibraltar and La

Réunion (Indian Océan). These aurbors conside-

red Verruca imbrtcata Gruvel, 1900b, V. striata

Gruvel, 1900b and V radiata Gruvel, 1901 as

synonyms of M. trisulcata. Hercin 1 only consi-

der V. striata synonymous of V. trisulcata , inclu-

ded in the Metaverruca. Verruca imbricarn and

V. radiata are considered valid and included in

Newmaniverruca n.g.

The spccimens examined are in accordance with

the dctailed description of the exrernal characters

presented by Gruvel (1902a), except the width of

the second and third ridges of the tergum are

equal, instead of the second being wider. The

ornamentation of the rostrum and canna, the

ridges of tergum and scutum, and the apex of

scutum, serve as a diagnostic of this species.

Gruvel (1900b, 1902a) did not describe the

appendages ol V. trisulcata, which art described

herein, but ht described the appendages of

V. striata (Gruvel, 1902c), which is Itéré con.side-

red synonymous. The description of the appen¬

dages of V. striata concord with tltose I observed,

except that of the caudjl appendage, Gruvel

(1902c) observed nine articles that were longer

than coxopoditc of cirrus VI, instead of seven

articles that are one hait the length of the coxo-

podite. Verruca striata also differs from V. trisul¬

cata-. scutum with apex not curved, with three

articular ridges similar in width, and the apices

of’ the canna and rostrum not projecting.

Notwithstanding these différences. I agréé with

Foster & Buckeridge (1995a, h) attd consider

this species synonymous with V. trisulcata.

On the other hand, I do not consider V imbri¬

cata and V. radiata as synonyms of M. trisulcata,

due to several différences in the number and

form of the ridges ol the scutum and tergum,

and on the number and development of the

radial ridges of the shell. Foster & Buckeridge

(1995a, b) stated that the number of ridges bet¬

ween the tergum and the scutum increased with

growth, from one to four in specimens of about

6 mm in rostrocarinal length. The three spéci¬

mens I studied had a length between 0.44 and

0.52 mm and ail had a constant number of

ridges, as in Gruvel’s type. In the specimens from

ZOOSYSTEMA • 1998 ■ 20 (1)

Young P. S.

La Réunion (Foster & Buckeridge 1995b),

besides the variable number of ridges, the spéci¬

mens exh dur a plasticity in the torm ol the shell

from low splayed to quite upstanding . Citri I and

II hâve the anrerior ramus one half the lengrh ol

the poscerior one and the caudal appendage with

0.66 or 0.5 (sic) the length of cirrus VI. For the

specimens ftom the Straits of Gibraltar, Foster &

Buckeridge (1995a) repeated the samc descrip¬

tion as fhar lor the La Réunion samples - , only

adding new figures. Therefore, the différences

observed in the shell and appendages ol both

samples appears to be yet anorher species, or pos-

sibly more titan one species due to the high

variability described.

The distribution of M. trisu/cata is restricted to

the Azores région and off Morocco (Fig. 9).

Genus Costatoverruca n.g. 1

Costatoverruca cornuta (Aurivillius, 1898)

Verruca cornuta Aurivillius, 1898: 197. - Gruvel

1905: 174; 1912a: 5; 1920: 39, pl. 2, figs 12-13,

pl. 3, figs 9-10. - Hoek 1907b: 9. — Belloc 1959: 4.

MATERtAl EXAMINER, — Stn 62, I specimen, rc:

0.5 cm, MN1IN Ci 2662. — Srn 148. 13 specimens

and 1 empry shell, rc: 0.4-0.7 cm, MNHN Ci 2577,

MNRJ 8911. — Stn 150, 2 spécimens, rc: 0.4 cm,

MNHN Ci 25 7 9. — Stn 151, 3 spécimens, il:

0.4-0.5 cm, MNHN Ci 2670. — Stn 157, 5 spéci¬

mens, rc: 0.5, MNHN Ci 2582. — Stn 159, 1 speci¬

men, rc: 0.4 cm, MNHN Ci 2586. — Stn 161,

25 specimeus, rc: 0,2-0.4 cm, MNHN Ci 2587,

MNRJ 8912. — Srn 181, 8 specimens, rc:

0.3-0.6 cm, MNHN Ci 2601. — Stn 199, 6 speci¬

mens, rc: 0.5 cm, MNHN Ci 2612, MNRJ 8913.—

Stn 237, 2 specimens, rc: 0.4 cm, MNHN

Ci 2630. — Stn 239, 646-628 m, 1 specimen, rc:

0.4 cm, MNHN Ci 2633.

Remarks

The external shell characters C. cornuta were des¬

cribed by Aurivillius (1898), and later in greater

detail by Gruvel (1920), both of which are based

on samples from the Azores. Young (in press)

1. See the part of the révision of Verrucidae for generic diagno-

sis.

redescribes this species, including a description

of rhe appendages, based on new samples from

the Azores.

Ail of the specimens studied were collected in the

Azores and at a depth previously recorded, 450

to 1229 m.

Genus Verruca Schumacher, 1817

Verruca stroemia ( .VI ü 11er, 1776)

(Figs 15-17)

Lepas strômia Müller, 1776: 251.

Verruca strômia - Darwin 1854: 518, pl, 21,

fig. la-f. - Pilsbry 1916: 24.

Verruca stroemia - Nilsson-Cantell 1978: 48,

figs 23-24.

MA TF RI Al- EXAMINER. — Stn 14, 2 specimens on

DendrophylUa cornigera, MNHN Ci 2554. —

Stn 259, 720 specimens, MNHN Ci 2653, MNRJ

8914.

Description

Shell (Fig. 15A) white or translucent, flattened,

usually with rostrum prominent, cuticle not per¬

sistent, ornamented with sevcral longitudinal

ridges, sometimes nodose, and irregular growth

fines; ridges at suture between rostrum and cari-

na and between rostrum and fixed-scutum

conspicuous and projecting. Opercular plates

(Fig. î 5A) parallel with hasis, reduced in sîze, less

lhan one half rostrocarinal diameter. Surlace of

opercular plates and shell permeated by several

rows of tubes in small specimens, tubes parallel

to growth fines sealed in larger ones. Basal mar-

gin of wall not inflecred.

Fixed-iergum (Fig. 15A) same size as fixed-

scuiurn, marginal apex thickened, recurved out-

ward; aJate projections to carina, nearly straight

sutures to fixed-scutm.

Fixed-scutum (Fig. 15A) nearly quadrangular,

upper surface turned roward rostral margin of

free scutum; suture with fixed-tergum simple and

nearly straight; rostrum having conspicuous

ridges; apex marginal, not recurved. Imernally,

with well developed myophore parallel to basis.

Canna (Fig. 15A) smaller than rostrum, surface

with longitudinal ridges knobbed; well developed

ZOOSYSTEMA • 1998 - 20 (1)

Cirripedia from the Azores Région

ridge area at suture with rostrum, forming deep

grooves between ridges, with fîve interlocking

teeth, ridges decreasing slightly in size from apex

to basis; with radius-like projection to fixed-

tergum.

Rostrum (Fig. 15A) with well developed rïdged

area ar suture with carina, forming deep grooves

between ridges, wiLh four to five interlocking

teeth, ridges decreasing in size from apex ro basis;

.suture with fixcd-scutum also with conspicuous

ridges.

Opercular plates with conspicuous and projec-

ting growth lines. Tergum (Fig. 1 5A, B) larger

than scutum, nearly quadrangular, with three

articular ridges; axial ridge high, conspicuous at

Fig. 15. — Verruca stroemia (Muller). A, top view; B, C, tergum

and scutum, internai view. Scale bars: 1 mm.

both sides, about saine width as second; ridges

absenr at carinal area. Internally, surface fiat,

with some visible sealed tubes, scutal margin

with deep médial norch, forming articular ridge

ac upper margin.

Scutum (Fig. I SA, C) with three low articular

ridges, axial ridge low and rhin, sloping conti-

nuously ro rostral area; First ridge wider rhan

second; rostral area, slightly medially dcpressed,

ridges absenr; width of plate about ont third ÎLs

height. Internally, surface with some sealed tubes

and conspicuous upper dépréssion, which

includes the adducror muscle pir, tergal margin

sinuous.

Labium with row of simple, sharp teeth. Palp

(Fig. 16A) acuminate, simple setae at upper mar¬

gin. Mandible (Fig. 16B) with three teeth,

second and third with subsidiary cusps, denticles

on lower angle. Maxilla 1 (Fig. I6C) with lower

anterior border projecting anteriorly; upper bor¬

der with two large spines, followed by nineteen

to rwenty moderate and small spines at lower

border. Maxilla II (Fig. 16D) bilobed, numerous

simple secae along 1rs margins,

Cirrus 1 (Fig. 16E) with unequal rami, anterior

ramus rwo third lcngrh of posterior, both rami

with protubérant articles covered by numerous

simple setae. Cirrus II (Fig, 16F) with unequal

rami, anterior ramus about one quarter length of

posterior, both rami covered by numerous simple

setae; distal articles of anterior ramus with bipec-

tinate setae. Cirrus III (Fig. 16G) with unequal

rami; anterior ramus about onc third length of

posterior; both rami covered by numerous simple

setae, distal articles of anterior ramus wirh bipec-

tinate setae. Cirri IV ro VI with equal rami;

intermediate articles (Fig. 16H) with four pairs

of setae on anterior margin, one or rwo setae on

posterior angle; width about one half length.

Caudal appendage (Fig. 161) multiarticulated,

sixreen articles, three rimes length ol coxopodire

of cirrus VI; articles covered by numerous simple

setae at distal margins; basal articles with multi-

fid scales. Number of articles cirri I-VI and cau¬

dal appendage is presented in table 7.

Remarks

Verruca stroemia is a common temperate and

polar species from the Northeastern Atlantic,

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Table 7. — Number of articles of cirri l-VI, and caudal appen-

dage of Verruca slroemia (Muller), stn 14. l-VI, cirri I to VI; CA,

caudal appendage; RC, right cirri; LC, left cirri.

III

9/9

6/15

7/15

17/21

21/22

23/25

9/8

6/16

8/18

18/19

21/22

24/26

where it occurs from the intertidal zone to

548 m, with doubtful records from 960-998 and

2600 m. The lacter record was from the Azores

région, which is the most Southern record for

this species (Gruvel 1902a, 1920).

In several samplings hy the Meteor (Young in

press) and the Jean Charcot expéditions in the

Azores, not a single specimen of V. stroemia has

been collccted. However, VI spengUri, a species

similar to V. stroemia , is commun in this région,

but it does not occur at the depths cited by

Gruvel. Ir appcars that die Gruvel (1902a, 1920)

records represent othcr verrucids species.

The specimens of V. stroemia studied herein were

collected off Portugal, which is acceptcd here as

the Southern limit of its distribution.

Verruca stroemia has been recorded from the

Spitsbergen, Greenland, Iceland, the Faeroe

Islands, Great Britain, and the Norwcgian to

Portugal coasts (Nilsson-Cantell 1978), with

scattered records along die Europe-an coast of the

Mediterranean Sea, from France to the Adriatic

Sca (Fischer 1871; Kolosvary 1947, 1951; Relini

1969). There is also one doubtful record from

the Red Sea (Darwin 1854; Fig. 17). Due to the

superficial similarity between V. stroemia and

V. spengleri ail records from the Mediterranean

need to be reviewed (sce remarks under V. spen-

glerî).

Verruca spengleri Darwin, 1854

(Figs 17-19)

Verruca spengttn Darwin, 1854; 521, pl. 21,

fig. 2. - wèltner 1897; 274. - Hoek 1883:

133. - Gruvel 1905: 182, fig. 200; 1920: 48. - Hoek

1907b: 9. - Pilsbry 1916: 40. - Baker 1967:

47. - Buckeridge 1994: 90.

Verruca stroemia — Ruggieri 1953: 46 (not V. stroemia

Millier).

? Verruca spengleri - Tarasov & Zevina 1957: 151,

figs 49-50. - Zevina 1963: 73. - Ruggieri 1977: 71k,

figs 3-5.

MATF.KIAL EXAMINED. — Stn L8, 1 specimen,

MN11N Ci 2541. — Stn P3, 42 specimens, MNHN

Ci 2507. — Stn P4, 46 specimens, MNHN

Ci 2509. — Stn P6, 218 specimens, MNHN

Ci 2511. — Stn P7. more titan 100 specimens,

MNHN Ci 2513, MNRJ 8915. — Stn P8, more

than 100 specimens, MNHN Ci 2516, MNRJ

8916- — Stn P9, .50 specimens, MNHN Ci 2518. —

Stn P12, 86 specimens, MNHN Ci 2521. —

Stn P13, 1 specimen, MNHN Ci 2522. — Stn P17,

4 specimens, MNHN Ci 2658. — Stn P19, 110 spe¬

cimens, MNHN Ci 2525. — SmP23, 68 specimens-,

MNHN Ci 2526. — Stn P24, 98 specimens,

MNHN Ci 2527. — Stn P27, 280 specimens,

MNHN Ci 2328, MNRJ 8917. — Stn P29, 298 spe¬

cimens, MNHN G 2529. MNRJ 891.8, — Stn P30,

1 specimen, MNHN Ci 2660. — Stn P33, 86 spéci¬

mens, MNHN Ci 2532. — Stn 1, 52 specimens,

MNHN Ci 2547. — Stn 61, 69 specimens, MNHN

Ci 2558. — Stn 142, 22 specimens, MNHN

Ci 2572. — Stn 143, 1 specimen, MNHN

Ci 2573. — Stn 216, 62 specimens, MNHN

Ci 2619. — Stn 224, 16 specimens, MNHN

Ci 2623.

Description

Shell (Fig. 17A) whice or translucent; cutide not

persistent; flactened; usually with rostrum more

prominent, ornamented only with irregular

growth iines; ridges at suture between rostrum

and carina conspicuous and projecting.

Opercular plates (Fig. 17B) parallet to basis,

reduced in sifce, less than one half rostrocarinal

diameter. Surface of opercular plates

(Fig. 17A-D) and shell permeated by several per¬

sistent rows of tubes, parallcl to growth Iines.

Basal margin ofwall not inflected.

Fixed-tcrgum (Fig. 1 7A) smaller than fixed-

scunim, marginal apex thickencd, recurved out-

ward; alace projections absent, sutures essendally

straight ot with small ridge at carinal suture.

Fixcd-scutum (Fig. I7A) nearly quadrangtilar,

upper surface turned toward rostral margin of

scutum; suture areas simple and nearly straight,

small to large ridge at suture of rosrrum; apex

marginal, not recurved outward. Internally, with

well developed myophore parallcl to the basis.

Carina (Fig. 17A) smaller than rostrum, well

developed ridge area at suture with rostrum

ZOOSYSTEMA • 1998 ■ 20(1)

Young P. S.

forming deep groovcs between ridges, five to

seven interlocking ccech, ridges decreasing slight-

ly in size from apex to basis; suture with

fixed-tergum with low ridges.

Rostrum (Kg. 17A) witli well developcd ridged

area at suture with carina, forming dcep grooves

between ridges, four to six inferlocking teeth,

ridges decreasing slightly in size from apex to

basis; suture with fuced-scutum nearly smooth or

depressed for fixed-scutum tooth.

Opercular plates with growth lincs conspicuous,

but nor projecting. Tetgum (Fig. 17C) larger

than scutum, nearly quadranguiar, threc articulât

ridges; axial ridge elevated, conspicuous at both

sides, as wide as second, distance between ridges

equal, groove between First and second larger

than thaï between second and third; without

ridges at carinal area. InternaJIy, surlace Hat,

some sealed tubes visible; scutal margin sinuous.

Scutum (Fig. I7D) with three low articular ridges,

axial ridge low, thin. sloping cnntinuously to ros¬

irai area; First and second ridges equal in width or

latter wider, no ridges at rosirai area; greater width

of plate about one half scutum heighr. Intcrnally,

surface with sonie sealed tubes visible, adductor

ridge conspicuous; tergal margin sinuous.

Labium (Fig. 18A) wirh row of twenty-four

simple, sharp tecth. Palp (Fig, 18A) acuminate,

simple Setae at upper margin. Mandible

(Fig. 18B) with three teeth, second and third

usually with subsidiary cusps at upper margin,

low denricles on lower angle. Maxilla I

(Fig. 18C) with lower anterior border projecting

anteriorly; upper border with two or Lhree large

spines, tollowcd by fifteen to rwenty-four inter¬

médiare to small spines at lower border.

Maxilla II (Fig. 18D) bilobed, with rmmerous

simple setae along its margins,

Cirrus I (Fig. 18F) with unequal rami, anterior

ramus one half length oF posterior, anterior

ramus with protubérant articles, covered by

numerous simple setae, posterior ramus with

slightly protubérant articles.

Cirrus II (Fig. 18F) with unequal rami, anterior

ramus about one third length of posterior, both

rami covered by numerous simple setae, distal

articles of anterior ramus svith bipectinate setae

(Fig. 18G).

Cirrus III (Fig. 18H) with unequal rami, anterior

ramus about one half length of posterior, both

rami covered by numerous simple setae, distal

articles of anterior ramus wirh bipectinate setae.

Cirri IV to VI with equal rami; intermédiare

articles (Fig. 181) wirh four pairs of setae on

anterior margin, one or two setae on posterior

angle, small spinules on distal margin, length

about one half width. Caudal appendage

(Fig. 18J) inultiarticulated, with Fourteen

articles, two and one hall rimes length oF coxo-

podite ol cirrus VI; articles covered by numerous

simple setae at distal margins; basal articles

(Fig. 18K) with spinules and multifid scales.

Nurnber of articles oF cirri 1-VI and caudal

appendage is présentée! in table 8.

Table 8. — Nurnber of articles of cirri l-VI, and caudal appenda¬

ge of Verruca spengleri Darwin, stn P6. I-VI, cirri I to VI; CA,

caudal appendage; RC, right cirri; LC, left cirri; +, broken ramus.

III

8/11

6/13

7/15

17/18

16+/21 +

23 + 122 +

8/11

6/14

8/15

18/18

20/21

23/22

Remarks

Darwin (1854) described and Figured specimens

ol V. spengleri , but he only illustrated the internai

view oF the scutum. This species is similar to

V. stroemia, luit the wcll developed adductor

ridge of the movable scutum in V spengleri readily

separates these species. Otherwise, both species

can be distinguished by V. Spengleri having a rela-

tively narrower scutum, the First and second

ridges being of equal width, the shell being per-

meated by tubes, and lacking longitudinal ridges.

Verruca spengleri was First described from

Madeira Island (Darwin, 1854) and latet, from

the Azotes (Gruvel 1920; Baker 1967) and Black

Sea (Tarasov Se Zevina 1957). Darwin (1854)

nored that "from geographical considérations”

V. spengleri probably was the species fourid in the

Mediterranean. Ruggieri (1977) cited this species

in Southern Italy, and also noted chat probably

ail of rbe citations of Verruca from the Mediter¬

ranean Sea were actually V. spengleri. Ruggieri

(1977) did not describe his specimens, but his

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

figures 3 and 4 show the scutum with a develo-

ped adductor ridge, a thin first external ridge,

and a width/height ratio of one h al F, which

agréés with rhe description of V. spengleri.

Tarasov & Zevina (1957) cîtcd V. spengleri as com-

mon in the Black Sca, bctween 20 and 50 m. The

specimens dcscribcd are similar ta those I observ-

ed, but the scutum is relarively larger and che axial

ridge is placed in the mîddle of the basal margin

(Tarasov Zevina 1957, fig. 49). In addition, the

caudal appendage has only nine articles. There-

fore, 1 consider the identification of the Black Sea

specimens ol V. spengleri as dubious, and in need

of further confirmation. Due to the large diffé¬

rences between the specimens from the Azores-

Madeira Archipelagos and rhe Black Sea, I suspect

that the Mediterranean specimens may weil be an

undescribed species.

Verruat spengleri is common in rhe Madeira and

Azores Archipelagos, from between the intertidal

zone and 103 m. Gruvel (1920) also recorded it

from 130 to 440 m (Fig. 19).

40° 20° 0° 20° 40°

Fig. 19. — Géographie distribution of Verruca stroemia (Müller) (dotted) and Verruca spengleri Darwin (hatched).

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Suborder BALANOMORPHA Pilsbry, 1916

Family Chthamalidae Darwin, 1854

Genus Chthamalus Ranzani, 1817

Chthamalus stellatus (Poli, 1791)

Lepas stellatus Poli, 1791: 29, pl. 5, figs 18-20.

Chthamalus stellatus - Darwin 1854: 45, pl. 18,

figs la, c, c-f, h (part). - Southward 1976: 1009, text-

figs 1-2, 6, pl. 1, figs c-d, f-g, pl. 2, figs a-b, d (syno-

nymy).

Chthamalus stellatus stellatus - Pilsbry 1916: 302,

pl. 71, figs l-4a. — Nilsson-Cantell 1921: 281.

MATERIAL EXAMINEE». — Stn L5, 96 specimens,

MNHN Ci 2537, MNRJ 8919. — Stn Lll, 3 speci¬

mens, MNHN Ci 2657.

DiAGNOSIS AND DESCRIPTION. — See Southward

1976: 1009.

Remarks

The présent specimens conform to the descrip¬

tion of the shell and appendages formulated by

Southward (1976). Chthamalus stellatus is widely

distributed from the northern Scottish coast and

English Channel, to the Mediterranean Sea, to

the tropical coast of West Africa, Mauritania and

with some doubtful records south of the

Cameroons (Stubbings 1967; Southward 1976).

It was also recorded from Madeira, Cape Verde

and Azotes Tslands (Darwin 1854; Stubbings

1964; Baker 1967). h is abundant between the

Low Water Mark and the splasb zone on ail rocky

shorex of the Sâo Jorge lsland (Baker 1967). Fhe

Jean Charcot Expédition collected this species at

Santa Maria and Faial lsland; rhus it has a wide

distribution within the Azores Archipelago.

Family BathylaSMATIDÆ Newman et Ross, 1971

Genus Bathylasma Newman et Ross, 1971

Bathylasma hirsutum (Hoek, 1883)

Balanus hirsutus Hoek, 1883: 158, pl. 13, figs 8-15.

Bathylasma hirsutum — Newman & Ross 1971: 149,

textfig. 73, pl. 23-24 (synonymy).

MATERIAL EXAMINED. — Stn 6, eroded shell frag¬

ments, MNHN Ci 2551. — Stn 46, 9 pièces of ero¬

ded and dîsarciculated plates of the shell, MNHN

Ci 2556. — Stn 180, 1 scutum, MNHN Ci 2667. —

Stn 240. 1 eroded carina, MNHN Ci 2608. —

Stn 255, 13 specimens fragmented, te: 1.4-2.4 cm,

MNHN Ci 2651, MNRJ 8920. — Stn 260, 3 speci¬

mens, rc: 1.5-2.8 cm, MNHN Ci 2655.

Remarks

Bathylasma hirsutum appears to be one of the

ntost commoii Northern Hemisphere deep-sea

balanomorph barnacles. It has been recorded

several rimes in rhe Northeast Atlantic, from the

Faeroe Islands to the Azores, from depths of 944

to 1829 m (Hoek 1883, 1913; Pilsbry 1916;

Gruvel 1920; Southward & Southward 1958;

Utinomî 1965; Newman tk Ross 1971). The

présent samples are from 570 to 940 m, and

occur on rocks and on echinoid spines.

Bathylasma hirsutum was redescribed in detail by

Newman & Ross (1971).

Genus Hexelasma Hoek, 1913

Hexelasma americanum Pilsbry, 1916

(Figs 1,20-21)

Hexelasma americanum Pilsbry, 1916: 330,

pl. 69- - Utinomi 1965: 12,

Aaptolasma americanum - Newman & Ross 1971:

161, pl. 22a-b, 36-37; 1976: 46.

MATERIAL EXAMINED. — Stn 180, 1 specimen, with

the shell plates disarticulated (lacking one latera),

MNHN Ci 2599.

Description

Shell conic, color white or pale orange, aperture

tooched. Plates (Fig. 20A, B) with spaced and

conspicuous growth iines; curicle persistent only

on opercular places, in larger specimens, pilose

along growth lines; alae well developcd. with fine

lines, radii absent; basal margin of wall sometimes

undulated. Sheath (Fig. 20B) adpressed, wcll defi-

ned. Internally (Fig. 20B) lamina with thin longi¬

tudinal striae; spaces between striae covcced by

chitin in larger specimens. Carino-lateral narrow,

0.13 width of carina and 0.22 widrh of latéral.

Scutum (Fig. 20C) higher than wide, height 1.4

to width; apex recurved outwards, tergal segment

narrow, with wide and strong longitudinal

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

apico-basal rib near occludent margin; articular

margin straight, basal margin sinuous; occludent

margin straight; growth Unes conspicuous and

elevated. Internally (Fig. 20D), adductor ridge

absent, articular ridge rounded, médial to articu-

lar margin, forming shallow articular furrow; six

latéral depressor muscle crests poorly developed;

articular and occludent margins thickened, lea-

ving depressed triangular area medially.

Tergum (Fig. 20E, F) triangulat. with spur near

basi-scutal angle, spur furrow open; spur distally

rounded, curving continuously to basa! margin;

eight crests for depressor muscles, projecting;

articular ridge elevated. not curved.

Labrum (Fig. 21 A) hirsute, lacking teeth, with

shallow médial concavity. Palp (Fig. 21 B, C)

club-shaped, large, with numerous fine peccinate

setae. Mandible (Fig. 21D) with four reerh, dis¬

tance between first and second twice that bet-

ween second and third, second to fourth with

subsidiary cusps, lower angle obtuse. Maxilla I

(Fig. 21E) with two large spines followed by

conspicuous, large notch, twelve to thirteen

moderate and eight to nine small spines along

cutting edge. Maxilla II (Fig. 21F) triangular,

with long simple setae.

Cirrus I (Fig. 21 G) with subequal rami, anterior

ramus slightly longer than posterior; both with

protubérant articles covered by long simple setae,

distally with fine pinnate setae. Cirrus II

(Fig. 21 H) with equa! rami, articles protubérant,

with clusters of simple setae on protubérance and

distal angle; distal articles with fine pinnate setae.

Cirri 11I-VI with equal rami. Intermédiare atticle

of cirrus VI (Fig. 211) with length 0.60 width,

wirh rwo pairs of long, fine, pinnate setae on

anterior margin, small setulae below these pairs;

one or two short simple setae on posterior angle,

small scales on posterior margin. Pénis (Fig. 2IJ)

short, annulated, setuale on distal portion.

Number of articles of cirri I-VI is presented in

table 9.

Remarks

Foster (1981) considered Aaptolasma Newman &

Ross synonymous with Hexelasma Hoek and

proposed an enrended diagnosis. His diagnosis

stated that the scutum and tergum hâve no arti¬

cular ridge, but a tergal ridge is présent on the

Table 9. — Number of articles of cirri l-VI of Hexelasma ameri-

canum Pilsbry, stn 180. I-VI, cirri I to VI; RC, right cirri; LC, left

cirri.

III

12/11

15/20

29/29

43/44

47/49

46/49

13/13

13/24

16+/33

37/41

48/42

46/50

tergum of H. americanum. Therefore, Poster s

diagnosis has to be emended to include che tergal

articular ridge as absent or présent.

Hexelasma americanum is the only species of this

gel!us recorded from the Atlantic, lr was collec-

red off South Carolina, Blake Plateau, between

512 and 770 m (Pilsbry 1916; Newman & Ross

1971).

The specimen dredged by the Jean Charcot

Expédition is similar to that described by Pilsbry

(1916) and Newman & Ross (1971) except for

rhe following characters: The sheli has a pilose

cuticle, the spur of the tergum is distally roun¬

ded, the mandible has an obtuse angle and the

pénis has setulae at the distal portion. These cha-

racters may be individual variations due the very

few number of specimens described. Pilsbry

(1916) described only one complété and rhree

incomplète specimens, and herein I add another

specimen. Ail of the other diaractcrs of the sheli,

opercular plates and cirri agréé with the descrip¬

tions by Pilsbry (1916) and Newman & Ross

(1971), including the details on the growth lines

of the plates described by the latter.

Tire record of this species from the Azores greatly

expands the distribution of Hexelasma america¬

num (Fig. 1). The depth range of this species is

extended front 770 to 1235-1069 m.

Family TETRACLITIDAE Gruvel, 1903

Genus Tesseropora Pilsbry, 1916

Tesseropora arnoldi n.sp.

(Figs 22-23)

? Tetraclita squamosa var. elegans — Baker 1967: 46

(nor T. squamosa elegans Darwin,1854).

MaTERIAL EXAMINED. — Holotype: stn P9, 1 broken

specimen, rc: 0.7 mm, MNHN Ci 2520.

Paratypes: stn L9, 1 broken specimen, MNHN

ZOOSYSTEMA ■ 1998 • 20(1)

Young P. S.

Ci 2544. — Stn L)l, 6 specimens, rc: 0.2-0.4 cm,

MNHN Ci 2545. USNM 282803, MNRJ 8921. —

Stn P29, 15 specimens, rc: 1.0-4.0 min, MNHN

Ci 2531, MNRJ 8922. — Stn 216, 5 empty shells,

0.3- 0.5 mm, MNHN Ci 2621.

EtymolOCY. — Named in honor of Dr Arnold Ross,

in appréciation of his contributions to our knowledge

of the Tetraclitoidea.

Diagnosis

Shell and shcath white; radii incomplète; tubes

irregular, radially lengthened, with striae on

inner side of outer lamina. Scutum higher than

wide, adductor tidge separated from articular

ridge. I.abrum hairy; three or four conspicuous

teeth on each side of médial notch. Intermediate

article of cirrus VI with four pairs of setae on

anterior margin.

Description

Shell (Fig. 22A, B) conic, white, aperture smooth

or little toothed, cuticle usually persistent, when

présent fuiely pilose. Plates wich uniforndy thin

ribs, growth lines thin; alae developed, smooth;

radii incomplète, besr developed in latéral plate.

Sheath (Fig. 22C) adpressed, well defîned. Inner

lamina with rhin longitudinal striae in small spe¬

cimens to well developed striated ribs over septa

in larger ones. Tubes irregular, radially lenghte-

ned, with striae on inner side of outer lamina.

Base calcareous, thin.

Scutum (Fig. 22D) higher than wide, straighr,

growth lines conspicuous; articular and basal

margins convex, occludent margin straight and

toothed. lnternally, adductor ridge thin, well

developed, apical portion separated from articular

ridge; articular ridge long, almosr length of arti¬

cular margin, articular groove shallow; rostral and

latéral muscle dcpressor pi es conspicuous, shallow,

five rostral and latéral depressor cresrs, adductor

muscle pit small, rounded, supramedial.

Tergum (Fig. 22E) elongated, with spur near

basi-scutal angle, distal end rounded, curving

continuously to basal margin; longitudinal fur-

row open and broad; six crests lor depressor

muscles prominent; articular ridge slightly pro¬

minent in upper portion.

Labrum (Fig. 23A) hairy; three or four conspi¬

cuous teeth on each side of shallow médial

notch. Paip (Fig. 23B) club-shaped, large, with

numerous fine pectinate setae on upper margin.

Mandible (Fig. 23C) with five reeth, second to

fiourth with conspicuous subsidiary cusps, espe-

cially fourth; srrongly denticulated berween fourth

and fifth and at lower angle. Maxilla I (Fig. 23D)

with one or rwo large sptnes followed by a

conspicuous notch, seven to tune moderate and

five to seven small spines along cutting border; in

notch small spines seldom présent. Maxilla II

(Fig. 23E) bilobed with long simple setae.

Cirrus 1 (Fig. 23F) with subequal rami, anterior

twice length of posrerior; posterior one with pro¬

tubérant articles; articles of both ramt covered by

several long, simple, fine, pinnate setae. Cirrus II

(Fig. 23G) with equal rami, articles protubérant,

clothed with simple, fine pinnate setae; protopo¬

dite with plumose setae. Cirrus III (Fig. 23H)

with anterior ramus 1.35 length of posrerior,

articles with numerous bipectinare setae, few

simple and pinnate setae and multifid scales near

setae (Fig. 23I-K); protopodite with plumose

setae (Fig. 23L). Cirri IV-VI with rami of equal

length. Intermédiare article of cirrus VI

(Fig. 23M) equidimensional, four pairs of setae

on ancerior margin, one ro four simple setae on

posterior angle. Pénis short, annulated, with few

setuale. Number of articles of cirri I-VI is presen-

ted in table 10.

Table 10. — Number of articles of cirri l-VI of Tesseropora

ainoldi n.sp. (Holotype). I-VI, cirri I to VI; RC. right cirri; LC. left

cirri.

III

9/6

6/6

10/8

16/15

13/15

16/17

9/5

6/6

10/7

16/16

13/14

16/16

Remarks

Tesseropora atlantica was previously-the only spe-

cies of thi.s genus recorded from the Atlantic

Océan, having bcen reporred from Bermuda and

the Azores (Newman & Ross 1977) and on Saint

Paul Rocks (Edwards & Lubbock 1983a, b), The

description of Newrnan & Ross (1977) appears

to be based exclusively on the Bermuda speci¬

mens, including ail of their figures. But, in their

ZOOSYSTEMA ■ 1998 • 20(1)

Cirripedia from the Azores Région

material examined, they cite the specimens stu-

died by Baker (1967) from the Azores.

The samples from the Jean Charcot Expédition

show that the species of Tesseropora from the

Azores is conspicupusly distinct from that descri-

bed from Bermuda. The inner lamina of the

plates hâve ribs well developed continuously

from the base to the sheath, intercalated by

numerous fine ribs, instead of only small nume-

rous fine ribs The adductor ridge of the scutum

is removed from the articulai - ridge, whereas

T. atlantica has the adductor ridge continuous

with the articulai - ridge. The labi um has conspi-

cuous teeth on the crest, contrary to the lack of

teeth in T. atlantica , The articles of cirrus III

hâve multifid scales and no curved spines or den-

ticles on the anterior ntargin. The imerntediate

articles of c irrus VI hâve onlv four pairs of paired

setae, instead of five pairs as in T. atlantica.

The specimens examined are smaller than those

observed by Newman & Ross (1977); most are

about 5 mm în carino-rostral diameter and only

one specimen is 7 mm

Tesseropora arnoldi n.sp can be distinguished

from Pacific Testerapom [T, rosea (Krauss) and

T. wireni (NillsoD-Cantell)J, by the color of the

sheath, the development of the radii and the

pariétal tubes and the structure of the cirri.

Tesseropom rosea has the sheath dirty white tinted

pink; the radii are moderately wide, and the

intermediate articles of cirrus VI hâve three pairs

of setae and numerous short setae in dense

bunches below the two major pairs (Newman &

Ross 1977). 7. wireni has the sheath usually

pink; the pariétal cubes are divided into secondary

and tertiary rows basally, and the intermediate

articles' of cirrus VI hâve three pairs of setae and

a few short setae in dense bunches helow the two

major pairs (Newman & Ross 1977)•

Tesseropom arnoldi n.sp. was collcctcd intertidally

to about 25 m deprh, usually attached to Mega-

balanus azorirus and mollusc shells. Several

samples were taken in the Azores in intertida!

and shallow waterx, but the species was not com-

mon in these samples. It was collected only on

Faial and Sao Miguel Islands. Ail the dissected

specimens were incubating nauplii. Baker (1967)

observed that Tetraclita squamosa var. elegans

Darwin (? = 77 arnoldi n.sp.) was “very abundant

in a single situation at Urzelina in 1 m. of water”

at Sao Jorge Island.

Family Balanidæ Leach, 1817

Genus Balanns Da Costa, 1778

Balanus trigonus Darwin, 1854

Balanus trigonus Darwin, 1854: 223, pl.3,

fig. 7a-f. - Pilsbry 1916: 111, pl. 26, figs l-13e (syno-

nymy). - Baker 1967: 47.

Material examiner. — Stn L.8, 1 specimen,

MNF1N Ci 2542. — Stn P3, 1 specimen, MNHN

Ci 2508. — Stn P4, 1 specimen, MNHN

Ci 2510. — Srn P6, 31 specimens, MNHN

Ci 2512. - Stn P7, :> 170 specimens, MNHN

Ci 2514, MNRJ 8923. — Srn PS, shells fragments,

MNHN Ci 2517- — Stn P9, 5 empty shells, MNHN

Ci 2519. —Stn P14, 31 specimens, MNHN

Ci 2524. — Stn P15, 10 specimens, MNHN

Ci 2546. — Stn P17, 25 specimens. MNHN

Ci 2652. — Stn P29, 2 specimens, MNF1N

Ci 2530. — Stn P34, 1 specimen, MNHN

Ci 2533. — Stn P41, 6 specimens, MNHN

Ci 2661. - Stn 1, more than 160 specimens,

MNHN Ci 2548, MNRJ 8924. — Stn 10, 5 empty

shells, MNHN Ci 2552. — Stn 11, 2 empty shells,

MNHN Ci 2553. — Stn 60, 8 empty shells, MNHN

Ci 2557, — Stn 143, more than 100 specimens,

MNHN Ci 2574.

Remarks

Balanus trigonus has a world-wide distribution,

and is widespread in the Eastern Atlantic, occur-

ring from the Mediterranean African coast to

South Africa (Stubbings 1967). It was previously

recorded from the Azores by Baker (1967) and

Young (in press).

Genus Megabalanus Hoek, 1913

Megabalanus azoricus (Pilsbry, 1916)

Balanus tintinnabulum azoricus Pilsbrv, 1916: 62,

figs 8, 1 le, pi. 12, figs 2-2b. - Baker 1967: 46.

Megabalanus azoricus — Newman & Ross 1976:

67. Henry & McLaughlin 1986: 21, figs la, 3d-e,

6a-d.

Material examiner, — Stn L9, 10 specimens, rc. :

0.4-2.7 cm, MNHN Ci 2543, MNRJ 8925. —

Stn P7, 3 specimens, rc: 1.6-2.8 cm, MNHN

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Ci 2515. — Soi P38, 7 spécimens, rc: 2.8-3.8 cm,

MNHN Ci 2534, MNRJ 8926. — Stn P41,2 spéci¬

mens, rc: 2.9 cm, MNHN Ci 2535. — Stn 186, more

than 50 empty shells, rc: 0.5-3.5 cm, MNHN

Ci 2602. — Stn 216, 1 empty shell, rc: 3,1 cm.

MNHN Ci 2620.

Remarks

The wall and opercular plates of M. azoriais were

redescribed by Henry & McLaughlin (1986)

who reported it Iront the Azores and St. Helena

Island. The type localiry of rhis species is Terceira

Island (Pilsbry 1916), but it is also reported front

Sâo Jorge Island (Baker 1967). The Jean Churent

sampled tins species from the Faial and Sâo

Miguel Islands and, also, off Santa Maria Island.

Therefore, rhis species appears to he well distri-

buted in the Azores Archipelago, from the inter-

tidal to infralittora! zones. The record from

80-90 m off Santa Maria Island is based on one

empty crodcd shell, wltich probably fell down to

this depth.

THE GENERIC RTVISION OF

THE VERRUCIDAE

(Figs 24-25)

Remarks

Pilsbry (1916) proposed four sections to separate

the genus Verruca, i.e., Metaverruca, Verruca,

Cameraverruca and Altiverruca. He also described

four species groups in the Verruca section, viz:

group of V strnenùa , V. alha, V. nexa and V. calo-

theca. Ail o( the characters in this arrangement

were based alntost whollv on the structure of the

shell. He also included sorne différences on the

relative lengths o( the rami of citri 1 to III.

Ail of Pilsbry s sections hâve subsequently been

accorded generic status (Zevina 1978), and the

following généra Rostratn vernira Brocb, 1922,

Brochiverruca Zevina, 1993b and Spongoverruca

Zevina, 1987a hâve been added to the

Verrucidae (Broch 1931; Zevina 1987a, 1993b;

Buckeridge 1994). Rrntratoverruca encompasses

the V nexa group recogni/.ed by Pilsbry (1916).

Young (in press) questions the validity of the

genus Metaverruca, since ntany of its diagnostic

characters are présent in other généra.

C D

Fig. 24. — Schematic représentation of the relative position of

the opercular plates and size of the adductor ridge of selected

généra of Verrucidae, viewed from rostral side. A, Altiverruca',

B, Newmaniverrucan.g,; C, Metaverruca', D, Verruca s,s.

Additionally the presence of a myophore, the

niost used feature to characterize rhis genus, is

also well developed in V. stroemia, rhe type spe-

cres of Verruca.

Buckeridge ( 1997) recognized the problem of the

myophore as a diagnostic character, but proposed

mainraining the genus, and added new characters

for the diagnosis. Besidcs the thickcned basal

ledge, the well developed myophore, and rhe top

flattened as described by Pilsbry (1916), he

added the box-like shape ot rhe shell, and a

D-shaped orifice as diagnostic for rhis genus.

The greatest problem defining rhe gênera within

the Verrucidae is the diagnosis ol Verruca s.str.

Pilsbry (1916) diagnosed this genus as “Top flat¬

tened, the plane ol the movablc plates not far

from parallel with that ol the base; radio-alar area

between paricties ot fixed scurum and tecgum

small and linear ". Buckeridge (1994) defined this

genus as “Vertucids with apices of rostrum and

canna marginal; fixed scurum without myophore;

operculum parallel to base.” The diagnosis of

Buckeridge (1994: 90) is in error when he cites

rhe absence ot a myophore since Vstroemia, type

species of Verruca, has a well developed myopho¬

re, as do several other Verruca species.

ZOOSYSTEMA - 1998 • 20(1)

Cirripedia from the Azores Région

With the reassigment of many species of Vernira

s.l. to other généra (Metaverriua, Rostratoverruca,

Cameraverruca, etc.), the species which remain in

Verruca s.s. do not appear to be closely related. In

studying the shell characters of several species of

Verruca s.s., two transformation sériés can be

observed.

The First transformation sériés is based on the

shape of the sbell and the development of the

myophore' (Fig. 24). The First stage of develop¬

ment is attained in the species ot Alliverruca.

This is characterr/.ed by a steep shell with the

opercular plates nearly parallel to the fixed scu¬

tum and tergum (Fig. 24Â). The adductor

muscles are artached directly to the inner side of

the wall, without the scutum lacking an adductor

ridge and also with a poorly developed adductor

pit.

If we compare this pattern wirh an outgroup

référencé, the lepadotnorphs, this character can

be con.sidered plesiomorphic in the verrucids.

Both scuta of the lepadotnorphs are usually

parallel, and faii to develop an adductor ridge.

Considering the steep pattern of Altiverruca as

plesiomorphic, an evolutionary trend can be

vizuali/.ed when the shell becomcs more inclined

and flatter, which implies a change in the relative

position between the movable scutum and

fixed-scutum: The angle between Lhese plates

enlarges from narrowly acute to a right angle

(Fig. 24A, B). The change in position of these

plates also nécessitâtes a change in the attach-

menr of the adductor muscle. The development

of the adductor ridge ranges from a small low

ridge to a projectiflg tongue-myophore (e.g.

V. stroemià ; Fig. 24C, D). The development of

an adductor ridge makes it possible to maintain

the same area for muscle fixation.

The other transformation sériés is in the develop¬

ment of the secondary ridges on the rostrum and

the carina, directed toward the opercular valves.

The supposedly plesiomorphic position of the

umbos is marginal (apical), with a linear hinge

line between the movable opercular valves and

the rostrum-carina, as can be observed in basal

2. Sometimes, it Is dittlcult to detine the limlts between adductor

ridge” and ''myophore". The myophore is defined as a "tongue-

shaped adductor ridge” (Pilsbry 1916: 21).

Fig. 25. — Schematic représentation ot the rostra and carina

ridges of selected généra of Verrucidae, viewed from rostro-cari-

nal side. A, Newmaniverruca n.g.; B, Costatoverruca n.g.; C,

Rostratoverruca: D. Brochiverruca.

pedunculates (Fig. 25A). This stage is seen in

several species included in Verruca s.l. (e.g.

V. entobapta, V a/batrossiana, V. scrippsae).

The secondary ridges are ftrst developed on the

rostrum, beginniug at rhe umbo and movittg up

to the scutum basis (Fig. 25B). This stare is also

seen in soute species of Verruca s.l. (V. alba,

V. xanthia , V. floridana). The umbo of the ros¬

trum is displaced trom the margïn (Fig. 25C).

The species with this state comprise the genus

Rostratoverruca. Subsequently, the umbo of rhe

carina is also displaced from the margin

(Fig. 25D), which is représentative of the species

of Brochiverruca.

Based on these considérations, I propose dividing

Verruca into three généra: Newmaniverruca n.g.,

Costatoverruca n.g. and Verruca s.s. Newmani-

verruca n.g. contains the box-like species with

the opercular plates nearly parallel to the hasts,

with marginal umbos of roscrum and carina,

without any secondary ridges and without a

myophore. Costatoverruca n.g. encompasses the

box-like species with the opercular plates nearly

parallel to the hasts, secondary ridges developed

on rhe rostrum, and rarely with an adductor

ridge or myophore. Verruca s.s. contains the deri-

ved species with the shell flattened, the opercular

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

valves parallel, and the myophore strongly deve-

loped.

Therefore, Metaverruca is redefined to include

the box-like species with a weli developed myo¬

phore, without secondary ridges on the fixed ros-

trum. Also, the adults develop a iedge at the base

of the wall.

The genus Spongaverruca was described to isolate

V spongicola, which lives in sponges. Gruvel

(1911, 1912b) presented a short description with

no diagnostic character that allows the séparation

of this species in a new genus, appart from being

the only species occurring in sponges. The per-

pendicular position of the opercular valves in

relation to the basis mandates inclusion of this

species in Altiverruca. Although Buckeridge

(1994: 89) questioned the validity of this genus,

1 consider Spongaverruca as a synonym of

Altiverruca .

Broch (1931: 45) presented the name Eu-verruca

as a subgenus of Verruca to separate in the lollo-

wing way Verruca s.str. from the other subgenera

he described. He justified this new name due to

“The central group of the genus is beçrer ntarked

by fixing the prefïx Eu- to the genus name than

using the latter alone; this may serve to bring

confusion about.” Therefore Euverruca is an

objective synonym of Verruca.

KEY TO THE GENERA OF THE FAMILY VERRUCIDAE

la. Form erect; opercular plates perpendicular to base; without adductor ridge or myo¬

phore on fixed-scutum ..... Altiverruca

1 b. Form not erect; opercular plates at no more than 45° to base; with or without

adductor ridge or myophore on fixed-scutum .2

2a. Fixed tergum and rostrum medially expanded, forming internai partitioned cavities;

opercular plates 45° to base; with adductor ridge on movable scutum .

........... .....Cameraverruca

2b. Fixed tergum and rostrum not rnedially expanded, without internai cavities; oper¬

cular plates less than 45° to base; without adductor ridge on movable scutum .3

3a. Shell box-like; opercular plates proportionally large, their width one half or more

the width of shell; with or without myophore .4

3a. Shell flattencd; opercular plates proportionally small, their width less than one half

the width of shell; with a strong myophore. Verruca

4a. Umbo of rostrum and cardia marginal .....5

4b. Umbo of rostrum not marginal, umbo ofearina marginal or displaced from margin

5a. Rostrum without secondary ridges directed toward tergal base; without myophore;

basal margin of shell not thickened. Neivrnaniverruca n.g.

5b. Rostrum with secondary ridges; with or without myophore; basal margin of shell

not rhickened . Costatoverruca n.g.

5c. Rostrum without secondary ridges directed toward tergal base; with myophore;

basal margin of shell thickened . Metaverruca

6a. Umbo of rostrum displaced from margin . Rostratoverruca

6b. Umbo of rostrum and carina displaced from margin . Brocbiverruca

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Genus Altiverruca Pilsbry, 1916

Verruca Section D: Altiverruca Pilsbry, 1916: 40.

Verruca (Altiverruca ) — Broch 1931: 45. — Foster

1978: 68.

Altiverruca - Zevina 1987a: 1813. - Buckeridge 1994:

92.

Spongoverruca Zevina, 1987a: 1813.

Type SPECIES. — Verruca hoeki Pilsbry, 1907a, by ori¬

ginal désignation (Pilsbry, 1916: 40), Recent,

Anegada Passage, 18°30'N - 63°3l'W, 496 fathoms.

SPECIES INCLUDED. — A. gibbosa (Hock, 1883),

A. incerta (Hoek, 1883), A. nitida (Hoek, 1883),

A. quadrangularis (Hock, 1883), A. obliqua Hoek,

1883, A. sulcata (Hock. 1883), A. crenata (Aurivillius,

1898), A Costa ta (Aurivillius, 1898) n. comb.

(?= A. gibbosa ), A. inermis (Aurivillius, 1898)

n. comb., ,4. crecta (Gnivek 1900b), .4. longicannata

(Gruvel, 1900b), A. cristalhna (Gruvel, 1907),

A. plana (Gruvel, 1907), A. mitra (Hoek, 1907b)

(= A. gibbota ), A. darwin/ (Pilsbry, 1907a) (= A. gibbo-

sa), A. hoeki (Pilsbry, 1907a) (= K quadrartgularis),

A. joubini (Gruvel, 1912a), A. cassis (Hoek, 1913)

(= A. cristallin //). A , t'asula (Hock, 1913), A. m/vintla

(Hoek, 1913). A. bicomuta (Pilsbry, 1916) (= A. gib¬

bosa ), A. ra/bbunianu Pilsbry, 1916 (= A. gibbota),

A, cristallina laevis (Broch, 1922) (= 14 cristallina),

A. ornata Nilsson-Cantell, 1929, A regular.h

Nilsson-Cantell. 1929, A. gibbosa somaliensis

(Nilsson-Cantell, 1929) (- A. gibbosa), A allisoni (Rao

et Newman, 1972), A. ql/es (Zevina, 1975). A. angus-

titerga Zevina, 1987a, A . ga/apagtisa Zevina, 1987a,

A. gira (Zevina, 1987b), A. icutplurata Zevina, 1987a,

A. sublima Zevina, 1987a, A. longa Zevina, 1988,

A. tcbesunovi Zevina, 1988, A. v/lrea Zevina, 1988,

A. galkini Zevina. 1990, A. molltte Zevina, 1990,

A. bering/ana Zevina et Galkin, 1992, A, laeviscuta

Buckeridge, 1994, A. vertica Foster et Buckeridge,

1995a (= A. obliqua) and A. spongicola (Gruvel, 1911)

n. comb.

Diagnosis

Form erect; opercular plates erect, perpendicular

to base; flxcd scutum without adductor ridge or

myophore; suture between rosrrum and carlna

from linear to imbricate; bases of plates not

inflected.

Remarks

In this genus I include V. inermis Aurivillius,

1898, which is described in detail by Gruvel

(1920). Its steep pattern of growth, poorly deve-

loped imbricating suture between the rostrum

and the canna places this species in Altiverruca.

Due to rhe placement of Spongoverruca as a

junior subjective synonym of Altiverruca , I hâve

also included Spongoverruca spongicola (Gruvel,

1911).

Genus Cameraverruca Pilsbry, 1916

Verruca Section C: Cameraverruca Pilsbry, 1916: 39.

Verruca (Cameraverruca) - Foster 1978: 68.

Cameraverruca - Zevina 1987a: 1813. — Buckeridge

1994: 103.

Type SPECIES. — Verruca euglypta Pilsbry, 1907a, by

original désignation (Pilsbry, 1916: 39), Recent, off

Fernandina, Florida, 30°44 N, 79°26 , W, 440

fathoms.

Species INCLUDED. — C. euglypta Pilsbry, 1907a,

C. nodiscuta Buckeridge, 1994.

Diagnosis

Form with opercular plates forming angle of 45°

with base; apical cavities of ftxed tergum and ros¬

trum partitioned off, forming recesses of general

body cavity. Movable scutum with adductor

ridge.

Genus Newmaniverruca n.g.

Verruca Section B: Verruca, Group of Verruca alba

Pilsbry, 1916: 25 (in part),

Verruca Section B: Verruca, Group of Verruca calothe-

ca Pilsbry, 1916: 30 (in part).

Verruca ( Verruca) - Foster 1978: 68 (in part). - Zevina

1987a: 1812 (in part). - Buckeridge 1994: 90 (in

pan).

Type species. — Verruca albatrossiana Pilsbry, 1912;

Recent, east ofLuzon, Philippines, 310 fathoms.

Species included. — N. imbricata (Gruvel, 1900b),

N. radiata (Gruvel, 1901), N. multicostala (Gruvel,

1907), N. albatrossiana (Pilsbry, 1912), N. barbadensis

(Pilsbry, 1916), N, entabapia (Pilsbry, 1916), N.fldvi-

dula (Pilsbry, 1916), N. grex (Hoek, 1913)

(= N. albatrossiana) and, N. scrtppsue (Zullo, 1964).

ETYMOLOGY. — Named in honor of Dr William

A. Newman, who is largely responsible for my studies

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

on cirripeds, and for his interesting discussions about

barnacles.

Diagnosis

Form box-Iike, opercular plates parallel or almost

parallel wich base; urnbo of carina and rostrum

marginal, without secondât}' ridges and myophore.

Remarks

The species N. albatrossiana (Pilsbry, 1912) is

designated herein as the type species of

Newmaniverruca n.g. because it is the First well

recognised species. The species described by

Gruvel (N. imbricata, N, radiata , and N. rnulti-

costata) are too briefly described and hâve not

been collected again (Gruvel 1900b, 1901,

1907).

I am assigning spécifie status to the subspecies of

calothecn and alba, because rnany of these sub-

species become separated at the generic level

{Newmaniverruca n.g. and Costatoverruca n.g.) in

the classification formulated liere. The species

described by Gruvel are tentatively incJuded in

this genus by the absence of the secondary ridges

on the rostrum; no information was cited on the

internai surface of the fixed-scutum.

Key to species ;

la. Shell with conspicuous radial ridges directed toward the base of the plates.2

lb. Shell without radial ridges on the plates, only those directed to the rostrum-carina

articulation...3

2a. Rostral area of scutum smooth; carina smaller than rostrum .

........ N. multicostata (Gruvel)

2b. Rostral area of scutum transversely grooved; carina larger than rostrum .

....... N. albatrossiana (Pilsbry)

3a. Opercular plates nearly parallel with base of shell.4

3b. Opercular plates forming an angle with base of shell . N. flavidula (Pilsbry)

4a. Tergum wirh three articulât ridges ..... N barbadensis (Pilsbry)

4b. Tergum with four articulât ridges...... 5

4c. Tergum wirh five articular ridges . N. imbricata (Gruvel)

Sa. Scutum with three articular ridges... N. scrippsae (Zullo)

5b. Scutum with four atricular ridges...6

6a. Ail four articular ridges well developed . N. entobapta (Pilsbry)

6b. Only the two lower articular ridges well developed. N. radiata (Gruvel)

Genus Costatoverruca n.g.

Verruca Section B: Verruca, Group of Verruca alba

Pilsbry, 1916: 25 (in part).

Verruca Section B: Verruca, Group of Verruca calolbeca

Pilsbry, 1916: 30 (in part).

Verruca ( Verruca) - Foster 1978: 68 (in part). — Zevina

1987a: 1812 (in part). - Buckeridge 1994: 90 (in

part).

Type SPECIES. — Verruca alba Pilsbry, 1907a; Recent,

Straits of Florida, 24°25’45”N - 81°46’45”W,

45 fathoms.

Species included. — C. comuta (Aurivillius, 1898),

C. alba (Pilsbry, 1907a), C. calolbeca (Pilsbry, 1907a),

C, grimaldi (Gruvel, 1912a), C. caribbea (Pilsbry,

1916), C. heteropoma (Pilsbry, 1916), C. flaridana

(Pilsbry, 1916), C. xantbia insculpta (Pilsbry,

1916), C. xantbia (Pilsbry, 1916), C. niasiensis

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

(Nilsson-Camell, 1929), C. pacifiai (Buckeridge,

1994), and C. sinuosa (Foster et Buckeridge, 1995b).

ETVMOI.OGY. — from the Greek costata (ridge), refer-

ring to the présence of secondary ridgcs cm the ros-

truni.

DlAGNOSlS

Form box-like, opercular plates parallel or almost

parallel with base; umbo of carina and rostrum

marginal, rostrum with secondary ridges, seldom

on carina; with pit for adductor muscle, usually

adductor ridge or myophore.

Remarks

C. alba (Pilsbry, 1907a) is designated as rhe type

species of Costatoverruca n.g. because it is the

First species described and ilJustrated. Aurivillius

(1898) presented a brief description of

V. cornu ta, with no illustrations.

The spccies C. a/muta and C pacifica were pre-

viously included in MetavemiOi due to the pré¬

sence ofthe myophore (Buckeridge 1994; Youttg

in press), but both species hâve well dcveloped

secondary ridges on the rostrum, the diagnostic

feature of Costatoverruca n.g.

la.

lb.

2a.

2b.

2c.

3a.

3b.

4a.

4b.

5a.

5b.

6a.

6b.

7a.

7b.

8a.

8b.

Key to SPECIES :

Rostral area of scutum smooth .2

Rostral area of scutum with longitudinal ridges .3

Scutum with two tliin articulât ridges; tergum with four articular ridges .

... C. grintaldi (Gruvel)

Scutum with three articular ridges, only the axial thin; tergum with three articular

ridges..... C. xantbia (Pilsbry)

Scutum and tergum with only a minute axial ridge.

... C. sinuosa (Foster et Buckeridge)

Tergum with three articular ridges .....4

Tergum with four or more articular ridges ...7

Well developed myophore présent .5

Myophore absent.....6

Scutum smaller than tergum; fixed-scutum with a well developed upper triangular

area ..... C. cornuta (Aurivillius)

Scutum larger than tergum; without upper triangular area.

....C. xdnthia insculpta (Pilsbry)

Caudal appendage with twenty-three articles, more than three tintes length of pro-

topod .«. C. floridana (Pilsbry)

Caudal appendage with fourteen articles, 1.5 rimes length ofprotopod .

..... C. heteropoma (Pilsbry)

Carina without secondary ridges; caudal appendage twice or less the length of pro-

topod. ...8

Carina with secondary ridges; caudal appendage more than 2.5 rimes length of

protopod . C. alba (Pilsbry)

lnterlocking ridges of carina and rostrum subequal .9

lnterlocking ridges of carina and rostrum conspicuously larger in the upper por¬

tion . 10

ZOOSYSTEMA • 1998 ■ 20(1)

Young P. S.

9a. Cirri I and II with anterior abour one third length of posterior rami; caudal

appcndage with twenty-seven articles.G caribéen (Pilsbry)

9b. Cirrus I and II with rami of nearly equal length; caudal appendage with twenty-

one articles .... C. niasiensis (Nilsson-Cantell)

I()a. Articular margins ol scutum and tergum, in internai view, sitmous; caudal appen¬

dage with fourteen articles, slightly longer than protopod .

. •••••.,. C. calotheca (Pilsbry)

10b. Articular margins of scutum and tergum, in internai view, esscntially m raighr. with

only one tooth above; caudal appendage with 24-26 articles, more than twice

length of protopod . G pactfica (Buckeridge)

Genus Rostmtoverruca Broch, 1922

Verruca Section B: Verruca, Group of K nexa Pilsbry,

1916:29.

Verruca Section Rostratoverruca Broch, 1922: 298.

Verruca (Rostratoverruca ) - Broch 1931: 46. - Foster

1978:68.

Rostratovermca - Zevina 1987a: 1813. - Buckeridge

1994:118.

Type SPECIES, — Verruca nexa Darwin, 1854, by sub¬

séquent désignation Zevina (1987a: 1813); Recent,

West Indies, on a gorgonian,

Species inci.uded. — R. nexa (Darwin, 1854),

R. koehteri (Gruvel, 1907), R. conchula mirwr (Hoek,

1913), R. intexta (Pilsbry, 1912), R conchula (Hoek,

1913) (= R. intexta ), R. nexa multiradiata

(Nilsson-Cantell, 1921), R. kruegeri (Broch, 1922),

R. kruegeri multisculpta (Hiro, 1933), R. murrayi

(Stubbings, 1936) (- R. intexta ), R. sewelli (Stubbings,

1936), and R. malevichi Zevina, 1988.

Diagnosis

Form box-like, opercular plates parallel with

base; umbo of carina marginal; umbo of rostrum

displaced from margin; myophore absent.

Genus Brochiverruca Zevina, 1993b

Brochiverruca Zevina, 1993b: 9. - Buckeridge 1994:

105.

Type SPECIES. — Verruca dens Broch, 1931, by origi¬

nal désignation (Zevina 1993b); Recent, Key Islands,

05°46’S - 132°5TE, 348 meters, on a Madreporarian

coral.

Species inclcded. — B. dens (Broch, 1931), B. mar-

gulisae Zevina, 1993b, and B. polystriata Buckeridge,

1994.

Diagnosis

Form box-like, opercular plates parallel with

base; umbo ol carina and rostrum displaced from

margin; myophore absent.

Genus Metaverruca Pilsbry, 1916

Verruca Section A: Metaverruca Pilsbry, 1916: 21.

Verruca (Metaverruca) - Broch 1931: 41. - Foster

1978: 68.

Metaverruca - Zevina 1987a: 1812, - Buckeridge

1994:108.

Type SPECIES. — Verruca coraliophila Pilsbry, 1916,

by original désignation; Recent, between Bahamas

and Cape Fear (see Pilsbry 1916: 22, footnote).

Species tnciuded. - M, recta (Aurivillius, 1898),

M. sculpta (Aurivillius. 1898) (= M. recta ), M. aetjua-

lis (Aurivillius, 1898). M, line avis (Gruvel, 1900b)

(= M. recta), M. triiulcata (Gruvel. 1900b), M. striata

(Gruvel, 1900b) M. trisulcata), M. magna (Gruvel,

1901) (= A/, recta), M. Imkitheca (Pilsbry, 1907b)

(= M. recta), M. capsula (Hoek, 1913) (= M. recta),

M . coraliophila (Pilsbry, 1916) (- M. recta), M . corm-

gala (Broch, 1931), M. mai-uni (Stubbings, 1936),

M. taraiovi (Zevina. 1971), M. lepista (Zevina,

1987b), M. seriola (Zevina, 1987b), M. pallida

Zevina, 1990. M. defayeae Buckeridge, 1994, M. nor-

Jvtkensis Buckeridge, 1994, M. plicata Buckeridge,

1994, and M. reunioni Foster et Buckeridge, 1995b.

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

DIAGNOSES without secondary ridge; myophore well develo-

Form box-like, opercular plates parallel with ped, basal margin of plates enlarged, sometimes

base; umbo of carina and rostrum marginal, fused.

KüY TO SPECIES :

1 a. Shell with conspicuous radial ridges .2

1 b. Shell without radial ridges.4

2a. Numerous fine radial ridges, with eleven or more ridges on each plate.3

2b. Few large radial ridges, with ten or fewer ridges on each plate .

... M. trisulcata (Gruvel)

3a. Caudal appendage with seven articles, shorter than protopod .

. * . M. corrugata (Broch)

3b. Caudal appendage with iwenry-seven articles, more than twice length of protopod

.*. M. plicata Buckeridge

4a. Opercular valves with or without a barely defined articular ridge .5

4b. Opercular valves with at least one articular ridge conspicuous...6

5a. Opercular plates small in relation to the shell, canna and rostrum with a strai^ht

articulation.... M. pallida Zevina

5b. Opercular plates large in relation to the shell, carina and rostrum with a single

large articular ridge... M. reunioni Foster et Buckeridge

6a. Opercular plates nearly parallel with the base of the wall . 7

6b. Opercular plates forming a distinct angle with the base of the wall . 12

7a. Tergum with three articular ridges . 8

7b. Tergum with four articular ridges .. 9

8a. Shell smooth; carina and rostrum apical margins straight.

. M. recta (Aurivillius)

8b. Shell with grooves between the opercular valves ridges; carina and rostrum mar¬

gins concave... M. aequalis (Aurivillius)

9a. Scutum with three articular ridges . 10

9b. Scutum with four articular ridges. 11

10a. Rostrum and carina articulation with three ridges; radii-process of fixed scutum

sma M......... M. rnacani (Stubbings)

10b. Rostrum and carina articulation with four ridges; radii-process of fixed scutum

l ar g e ...*. M. tarasovi (Zevina)

1 la. Rostrum and carina articulation with as many as three ridges; inaudible with five

teet h ... M. defayeae Buckeridge

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

11b. Rostrum and carina articulation with as many as five ridges; mandible with three

teeth ...*. M. norfolkensis Buckeridge

12a. Scutum with a beaked apex; caudal appendage with six articles.

....... . . M. lepista (Zevina)

12b. Scutum with an obtuse apex; caudal appendage with eleven articles..

...... M. seriola (Zevina)

Genus Verruca Schumacher, 1817

Verruca Schumacher, 1817: 35. - Darwin 1854: 496

(in part).

Verruca Section B: Verruca, Group of V. strômia

Pilsbry, 1916: 23.

Verruca (F.u-Verruca) Broch, 1931: 45.

Verruca (Verruca ) - Foster 1978: 68 (in part). -

Zevina 1987a: 1812 (in part). - Buckeridge 1994: 90

(in part).

Type SPECIES. — Le pas stroemia Muller, 1776, by

monotypy; Recent, no locality (Schumacher, 1817:

91).

Sl’EClES INCLUDED. — V. stroemia (Müller, 1776);

V. spengleri Darwin, 1854; V. laevigata (Sowerby,

1827), and V cookei Pilsbry, 1927.

DtAGNOSIS

Form depresscd, opercular plates parallel with

base, proportionately small; umbo of carina and

rostrum marginal, without secondary ridge;

myophore weÜ developed, projecting parallel to

base.

KEY TO SPECIES :

la. Movable scutum with adductor ridge.2

lb. Movable scutum without adductor ridge ...3

2a. Caudal appendage with nine articles, one half length of cirrus VI .

.. V. spengleri Darwin

2b. Caudal appendage with eleven articles, two third length of cirrus VI .

..... V cookei Pilsbry

3a. Width of scutum less than its length, the second articulât ridge narrower than the

first articular ridge ... V stroemia Müller

3b. Width of scutum more than its length, the second articular ridge broader than the

first articular ridge . k laevigata Sowerby

THE AZORES FAUNA

Including the species reported herein and those

of previous studies on the Azotes Région (Hoek

1883; Aunvillius 1898; Gruvel 1900a, b, 1902a,

1920; Pilsbry 1916; Zevina 1976; Baker 1967;

Newman 5d Ross 1977; Young in press) there are

a total of sixty-one taxa in this région. Twelve of

these taxa are considered synonymous, viz:

S. edwardsi and S. alborauense (= Neoscalpellum

débité), S. créai/m and S. velutinum (= An'oscal-

pellum miehclottianum), S. striatum (= Arnigdo-

scapellum rigidum), S. molle (- Triangulo-

scalpcllum regiuin), S. gigas Gruvel, 1902

(= Tnanguloscalpellum ovale, noi S. gigas Hoek),

V. sculpta and V linearis (= Metaverruca recta),

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

S. vitreum Zevina, 1976 (= S. rigidum ); T squa-

mosa elegans Baker, 1967, and Tesseropora atlanti-

ca Newman et Ross, 1977 (part, specimens from

Azores) (= Tesseropora arnolcli n.sp .).

Within the rcmaining fifty-one species, four

records are probably misidenrifications by Gruvel

(1905, 1920): V stroemia, B. ampbitrite, B. spon-

gicola and B. crenatus, Verruea stroemia and

B. crenatus are northcrn shallow warer species

and were ordy recorded once from the Azores

Région. Balanus ampbitrite and B. spongicola

hâve not been recorded again since Gruvel

(1905, 1920).

The species of Poecilasma are poorly dcfined and,

therefore, tbere is the possibility that the three

species recorded axe synonymous. Two species,

P. aurantia and P. crassum , are definirely known

in this région. The lise of species fs, therefore,

reduced to forty-five, vvhich includes twenty-

eight lepadomorphs, cleven verrucomorphs and

six balanomorphs (Appendix 2).

The endemism of the Azores appears to be high,

fourteen species (31%). Three species occur in

the Azores and Madeira Islands, and Great

Meteor Bank, which gives an endemism of this

larger area to 38%. Furthemore, four species

occur at the Iberian Basin, with records also off

the Iberian Peninsula and Morocco coast, which

expands the endemism ratio to 47%. The remai-

ning species usually hâve a Northeasrern Atlantic

distribution or greater (Appendix 2). The Azores

fauna does not share any species with Tropical

Africa.

Conversely, it is interesting to note the low spe-

cies richness in shallow waters (less than 200 m).

Tesseropora arnoldi and M. dzoricus are endemic

to the Azores, V spengleri occurs also in Madeira.

Çbthamalus stellatus is recorded along the

European and North African coasts and B. trigo-

nus, which has a worid-wide distribution, may

hâve been recently introduced to the Atlantic

(Zullo 1992).

In die Northeasrern Atlantic the deep sea species

are reasonably wel| sampled along the

Azores-Macteira-Grear Meteor area (Iberian and

Canary Basins) and the North Sea (Norwegian

Basin) but they are grill poorly sampled in the

West European Basin.

The deep sea faunas of the Norwegian Basin and

of the Iberian/Ganarian Basins do not share any

common species, except for Scalpe/lum

scalpellum , a species with a wide depth range of

10-540 m, but usually between 30 and 200 m. lt

is recorded from shallow waters of the Iberian

Peninsula and scattered localities along the North

African coast (Nilsson-Cantell 1978).

Ackn owledgeme n ts

I wish to acknowledge Jacques Forest for allo-

wing me to study this collection and also for

information about the Biaçores Expédition;

William A. Newman and Arnold Ross, for

numerous discussions during this study and fheir

comments on the manuscript, which significandy

improved this paper; John S. Buckeridge, Diana

Jones and Annemarie Ohler far providing useful

comments on the manuscript; Florence

Kerdoncuff for helpful assistance with the

manuscript révision. This study was supported

by the Conselho Nacional de Desenvolvimento

Cientlfico e Tecnologico (CNPq) and additional

financial support Irom Fundaçâo Universitaria

José Bonitacio (FUJB).

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Cirripedia from the Azores Région

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ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

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species? Bulletin af Marine Science 50 (1): 66 74.

Submitted on 29 July 1997;

accepted on 15 December 1997.

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

APPENDIX 1

List of species per station. The numbers preceded by the letters L and P concern respectively interti-

dal hand collecting and scuba diving. The numbers without a letter refer to the stations from the

Jean Charcot.

Station

Date

Location

Species

2.X.1971

Santa Maria Island (South), Praia

Chthamalus stellatus.

5.X.1971

Praia de Santos. “E. Roque",

sur tube rejeté

Lepas anatitera.

L. pectinata.

8.X.1971

Terceira Island, West of Monte Brazil

Verruca spengleri,

Balanus trigonus.

11.X.1971

Faial Island, Horta

Tesseropora arnoldi n.sp.,

Megabalanus azoricus.

L11

11.X.1971

Faial Island, Caldeira do Inferno (South)

Chihamaius stellatus,

Tesseropora arnoldi n.sp.

I.X. 1971

Santa Maria Island. Ponta Malbusca, 12-25 m

Verruca spengleri,

Balanus trigonus.

2.X.1971

Santa Maria Island. Ponta do Casteleto, 12-15 m

Verruca spengleri,

Balanus trigonus.

7.X.1971

J. do Castro Bank, 28 e 13 N - 26°36'W,

25-35 m

Lepas anatifera,

Balanus trigonus.

8.X.1971

Terceira Island, Ponta de S. Diego

(West of Monte Brazil), 5-40 m

Verruca spengleri

Balanus trigonus,

Megabalanus azoricus.

10.X. 1971

Faial Island, Horta (Port),

0-18 m

Verruca spengleri,

Balanus trigonus.

11.X.1971

Faial Island, Monte da Guia (East of Caldeira

do Inferno), 5-15 m

Verruca spengleri,

Tesseropora arnoldi n.sp.,

Balanus trigonus.

P12

13.X.1971

Faial Island, Ponta Furada, 0-7 m

Verruca spengleri.

PI 3

13.X.1971

Faial Island. Horta, 3-18 m

Verruca spengleri.

P14

15.X.1971

S, Jorge Island, West of Cabo Monteiro, 40 m

Balanus triaonus.

P15

15.X.1971

S, Jorge Island, West of Cabo Monteiro, 27 m

Balanus trigonus.

P17

16.X.1971

Graciosa Island, Calle de Folga, 2 8m

Verruca spengleri,

Balanus trigonus.

P19

18.X.1971

Faial Island, Monte da Guia, 20 m

Verruca spengleri.

P23

22.X.1971

Flores Island, North of Santa Cruz, 15 m

Verruca spengleri.

P24

22.X. 1971

Flores Island, llheu da Muda, 22-30 m

Verruca spengleri.

P27

27.X.1971

Sâo Miquel Island. Caloura (Soulh), 0-10 m

Verruca spengleri.

P29

28.X.1971

Sâo Miguel Island. Ponta da Galera (South), 7-18 m

Verruca spengleri,

Tesseropora arnoldi n.sp.,

Balanus trigonus.

P30

29.X.1971

Sâo Miguel Island. Caloura (S.), 0-6 m

Verruca spengleri.

P33

30.X.1971

Sâo Miguel Island, Santa Clara,

Ponta Delgada (Sôuth), 7-15 m

Verruca spengleri.

P34

31.X.1971

Sâo Miguel Island. Ponta de Galera (Southeast),

10-12 m

Balanus trigonus.

P38

2.XI.1971

Sâo Miguel Island, llheu dos Mosteiros (VJ.), 2-37 m

Megabalanus azoricus.

P41

4.XI.1971

Sâo Miguel Island, Morro das Capelas (N.), 15-20 m

Balanus trigonus,

Megabalanus azoricus.

P44

8.XI.1971

Formigas Island. 35-45 m

Oxynaspis celata.

7.X.1971

38 n 13'N 26 C 36'W, 40-50 m

Verruca spengleri,

Balanus trigonus.

7.X.1971

38°11,5‘N -28’38.5‘W, 1200-1080 m

Metaverruca recta.

7.X.1971

38 Q 14'N - 26"38,5'W, 570 m

Altlverruca gibbosa,

Bathylasma hirsutum.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Station

Date

Location

Species

8.X.1971

38°39'N - 27°14,5'W, 28-33 m

Balanus trigonus.

8.X.1971

38°30'N - 27 C ’14,5 , W, 105-76 m

Balanus trigonus.

4.XII.1968

47=56,3'N - 07°32,S'W, 214-235 m

Verruca stroemia.

8.X.1971

38"39‘N 27=21'W. 990-880 m

Metaverruca recta.

12.X. 1971

37“34'N - 28°54'W, 784 m

Bathylasma hirsutum.

14.X.1971

38°33,5'N - 28 U 33'W, 66-70 m

Balanus trigonus.

14.X.1971

38' , 34'N 28=32,5'W 77 m

Verruca spengleri.

14.X.1971

38'’39,5'N - 28"37,5'W, 800-736 m

Poecilasma aurantia,

Metaverruca recta.

Costatoverruca comuta.

15.X.1971

38=34,5'N - 28‘"19,5'W, 1260-1225 m

Metaverruca recta,

Arcoscalpellum tritonis.

15.X.1971

38‘30'N - 275t,5'W, 245 m

Heteralepas microstoma.

17.X.1971

39°04.5'N - 28''07,5'W, 358-406 m

Oxynaspis patens.

105

20.X.1971

39 n 35'N - 31 Û 23'W, 1550 m

Arcoscalpellum michelottianum.

120

22.X.1971

39°03,5'N - 32 J 43,5’W, 2100 m

Lepas pectinata.

129

23.X.1971

38' 58'N - 33'26,5'W, 3056-3000 m

Trianguloscalpellum regium,

Teloscalpellum luteum.

131

24.X.1971

39‘04,5’N - 32M3,5'W, 2120 m

Lepas pectinata,

Neoscalpellum débité.

135

25.X.1971

39=24,5'N - 31 05,5'W, 860-760 m

Metaverruca recta.

139

26.X.1971

38 U 36,5’N - 28-17.5'W, 1260 m

Metaverruca recta.

142

30.X.1971

37’41,5'N - 25 31'W, 103 m

Verruca spengleri.

143

30.X.1971

37=42'N - 25°32’W, 69-61 m

Verruca spengleri,

Balanus trigonus.

148

30.X.1971

37=34,5'N - 25=34,5’W, 847-870 m

Poecilasma aurantia,

Metaverruca recta,

Costatoverruca comuta.

150

30.X.1971

37°37'N - 25=35’W, 600-550 m

Metaverruca recta.

Costatoverruca cornuta.

151

30.X.1971

37=37,5'N - 25=39,5'W, 788-780 m

Metaverruca recta,

Costatoverruca cornuta

157

31.X.1971

37°33,5’N - 25°43,5'W t 826-787 m

Poecilasma aurantia,

Costatoverruca cornuta.

159

31.X.1971

37°26'N - 25°51'W, 600-525 m

Poecilasma aurantia,

Metaverruca recta,

Altiverruca gibbosa,

Costatoverruca cornuta.

161

31.X.1971

37“39,5 N - 25 50,5'W, 590 m

Metaverruca recta.

Costatoverruca cornuta

168

1.XI.1971

37=48,5 N - 25 r 54'W, 800-665 m

Metaverruca recta.

171

1 .XI.1971

37-58,5 N - 26=07'W. 3215 m

Metaverruca aequalis.

174

2.XI.1971

38=06'N - 26°15'W, 3094-3038 m

Amigdoscalpellum rigidum,

Metaverruca aequalis.

176

2.XI.1971

38°00,5’N - 26°21,5'W, 2720-2440 m

Amigdoscalpellum rigidum.

179 '

3.XI.1971

38°05,5'N - 25°46'W, 1665-1590 m

Smllium acutum.

180

3.XI.1971

37°57,5’N - 25°33'W, 1235-1069 m

Glyptelasma hamatum,

Poecilasma aurantia,

Arcoscalpellum michelottianum,

Altiverruca obliqua,

A. gibbosa,

Metaverruca aequalis,

M. recta,

Bathylasma hirsutum,

Hexelasma americanum.

181

3.XI.1971

37°53’N - 25°35,5’W, 620-450 m

Metaverruca recta,

Costatoverruca cornuta.

186

4.XI.1971

37°51,5’N - 25°40'W, 455-370 m

Megabatanus azoricus.

196

5.XI.1971

37°50’N - 24°55,5’W, 1191 -1146 m

Metaverruca recta,

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Station Date

Location

Species

197

5.XI.1971

37=49,5'N - 25=01,5'W, 815 m

Arcoscalpellum michelottianum,

Altiverruca gibbosa,

Metaverruca aequalis.

Metaverruca recta,

199

5.XI.1971

37=54,5’N - 25°08'W, 800-770 m

M. thsulcata.

Poecilasma aurantia,

202

6.XI.1971

37=26,5'N - 25°00'W, 2900 m

Altiverruca gibbosa,

Metaverruca recta,

Costatoverruca cornuta.

Glyptelasma hamatum,

206

7.XI.1971

37=21’N - 25°28,5’W, 2047 m

Neoscalpellum débité,

Catherinum recurvitergum,

Amigdoscaipellum rigidum,

Trianguloscalpellum regium.

Amigdoscaipellum rigidum.

216

8.XI.1971

37°16,5'N - 24=46,5’W, 80-90 m

Verruca spengleri,

218

8.XI.1971

36=54’N - 25=08'W, 772-800 m

Tesseropora amoldi n.sp.,

Megabalanus azoricus.

Metaverruca recta.

224

9.XI.1971

37°03'N - 25=05’W. 60 m

Verruca spengleri.

227

10.XI.1971

37=1 Q'N - 25=19,5'W, 2160-2180 m

Glyptelasma hamatum.

231

10.XI.1971

36=55'N - 25“10'W, 440-380 m

Amigdoscaipellum rigidum.

Metaverruca recta.

237

11.XI.1971

37=21,5'N - 25=45,5'W, 759 m

Poecilasma aurantia,

239

11.XI.1971

37=28,5'N - 25°45’W, 646-628 m

Metaverruca recta,

Costatoverruca cornuta.

Metaverruca recta,

240

12.XI.1971

37°35’N - 25=32,5’W, 825-810 m

Costatoverruca cornuta.

Metaverruca recta,

245

14.XI.1971

40=57'N - 22°16’W, 4270 m

M. trisulcata,

Bathylasma hirsutum.

Neoscalpellum débité,

249

16.XI.1971

45=50’N - 17=32,5'W, 4620-4690 m

Planoscalpellum limpidus ,

Catherinum recurvitergum,

Amigdoscaipellum rigidum,

Trianguloscalpellum regium.

Neoscalpellum debile.

250

16.XI.1971

45=54,5'N - 17°29,5’W, 4631 m

Arcoscalpellum eponkos n.sp.,

Planoscalpellum limpidus,

Trianguloscalpellum ovale.

Neoscalpellum debile.

251

18.XI.1971

47=38'N - 08°56'W, 3600-3360 m

Arcoscalpellum tritonis,

255

18.XI.1971

47=47'N - 07=56,5'W, 900-960 m

Amigdoscaipellum rigidum,

Trianguloscalpellum regium.

Metaverruca recta,

259

19.XI.1971

48=07,5'N- 07=51,5'W, 190 m

Bathylasma hirsutum.

Verruca stroemia.

260

19.XI.1971

47°46'N - 08 '04‘W, 940-820 m

Scalpellum scalpellum,

Ponta Delgada

on the hull of the Jean Charcot

Bathylasma hirsutum.

Conchoderma auritum.

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

APPENDIX 2

Cirriped species vvith présent taxonomie status and geographical distribution and references citing

them in the Azores Région. *, herein.

Species

(Names used

by the authors)

Authors

Présent taxonomie status

Geographical distribution

Heteralepadidae

Heteralepas microstoma

Young, in press; *

same

Azores, Madeira, Great

Meteor.

OXYNASPIDIDAE

Oxynaspis celata

same

Cosmopolitan

O. patens

same

North Atlantic

POECILASMATIDAE

Dichelaspis sessilis

Hoek, 1883

same

Azores

Glyptelasma hamatum

same

Circumtropical

Poecilasma aurantia

Gruvel. 1920; *

same

Eastern Atlantic

P. carinatum

Gruvel, 1920

Glyptelasma carinatum

Cosmopolitan

P. crassum

Gruvel. 1920

same

Circumtropical

P. kaempteri

Gruvel, 1920

? = P. aurantia

P, unguiculus

Aurivillius, 1894

? = P. aurantia

—

Calanticidae

Scalpellum acutum

Hoek, 1883; Gruvel,

1920;*

Smilium acutum

Cosmopolitan

S. calyculus

Aurivillius. 1898; Gruvel,

1920

Aurivilialepas calyculus

Azores

S. falcatum

Aurivillius, 1898; Gruvel,

1920

Aur/vilialepas falcata

Azores, Great Meteor, off

Portugal

S. Grimaldii

Scalpellidae

SCALPELUNAE

Aurivillius, 1898; Gruvel,

1920

Scillaelepas grimaldi

Azores

Scalpellum vulgare

Scalpellidae

Meroscalpellinae

Gruvel, 1920

Scalpellum scalpellum

Northeastern Atlantic

Scalpellum alboranense

Gruvel, 1920

= N. débité

S. débité

Aurivillius, 1898; Gruvel,

1920;*

Neoscalpellum débité

North Atlantic

S. Edwardsii

Scalpellidae

Arcoscalpellinae

Gruvel, 1900a; 1902a

= N. débité

Arcoscalpellum

michelottianum

Zevina, 1976; *

same

Cosmopolitan

Arcoscalpellum tritonis

same

Northeastern Atlantic

Planoscalpellum limpidus Zevina, 1976;*

same

Northeastern Atlantic and

Scalpellum anceps

Aurivillius, 1898; Gruvel,

1920

Teloscalpellum anceps

Azores

S. atlanticum

Gruvel, 1900a, 1902a;

1920

Teloscalpelum atlanticum

Azores, off Portugal

ZOOSYSTEMA • 1998 • 20(1)

Cirripedia from the Azores Région

Species Authors Présent taxonomie status Geographical distribution

(Names used

by the authors)

S. erectum

Aurivillius, 1898

= A. michelottianum

S. gigas

Gruvel, 1905

= T ovale (not S. gigas Hoek)

North Atlantic

S. gracile

Gruvel, 1920

Teloscalpellum gracile

Northeastern Atlantic

S. incisum

Aurivillius, 1898; Gruvel,

1920

Teloscalpelum incisum

Azores

S. luteum

Gruvel. 1900a; 1902a; *

Teloscalpellum luteum

Azores

S. mamillatum

Aurivillius, 1898; Gruvel,

1920

Amigdoscalpellum mamillatum Azores

S. molle

Aurivillius, 1898

(= T. regium )

S. pusillum

Aurivillius. 1898; Gruvel,

1920

Weltnerium pusillum

Newfoundland and Azores

S. recurvitergum

Gruvel. 1905:’

Catherinum recurvitergum

Azores, ?East Africa

S. regium

Gruvel, 1900a; 1902a

Trianguloscalpellum regium

North Atlantic

S. rigidum

Aurivillius, 1898

Amigdoscalpellum rigidum

Northeastern Atlantic

S. striatum

Gruvel. 1905; 1920

= A. rigidum

S. velutinum

Gruvel, 1920

= A. michelottianum

S. vitreum

Zevina, 1976

= A. rigidum

(not S. vitreum Hoek)

Verrucidae

Verruca inermis

Aurivillius, 1898; Gruvel,

1920

Altiverruca inermis

Azores

V. costata

Aurivillius, 1898; Gruvel,

1920

Altiverruca costata

North Atlantic

V. crenata

Aurivillius, 1898; Gruvel,

1920

Altiverruca crenata

Azores

V. erecta

Gruvel. 1900b; 1902a

Altiverruca erecta

Azores

Altiverruca gibbosa

same

Cosmopolitan

A. obliqua

same

Azores; off Spain

Verruca recta

Aurivillius. 1898; Gruvel,

1920; ’

Metaverruca recta

Cosmopolitan

V. sculpta

Aurivillius, 1898; Gruvel.

1920

= M. recta

V. linearis

Gruvel, 1900b; 1902a;

1920

= M. recta

—

V. aequalis

Aurivillius, 1898; Gruvel,

1920;*

Metaverruca aequalis

Azores

V. trisulcata

Gruvel, 1900b; 1902a;

1920;’

Metaverruca trisulcata

Azores

V. cornuta

Aurivillius, 1898; Gruvel,

1920;*

Costatoverruca cornuta

Azores, Great Meteor

V. strômia

Gruvel, 1902a

probably a misidentification

—

V. spengteri

Gruvel, 1920; Baker,

1967;*

same

Azores, Madeira, Black Sea?

Chthamalidae

Chthamalus stellatus

Pilsbry, 1916; Gruvel,

1920; Baker, 1967;*

same

Northeastern Atlantic

Tetraclitidae

Tesseropora arnotdi n.sp.

Tetraclita squamosa

same

Azores

elegans

Baker, 1967

= T. arnotdi,

not T. elegans Darwin

—

Tesseropora atlantica

Newman and Ross, 1977

= T . arnoldi (in part,

specimens from Azores)

ZOOSYSTEMA • 1998 • 20(1)

Young P. S.

Species

(Names used

by the authors)

Authors

Présent taxonomie status

Geographical distribution

Bathylasmatidae

Balanus hirsutus

Gruvel, 1920; *

Bathylasma hirsutum

Northeastern Atlantic

Balanidae

B. crenatus

Gruvel, 1920

probably a misidentification

B. amphitrite

Gruvel, 1920

probably a misidentification

B. trigonus

Pilsbry, 1916; Gruvel,

1920; Baker, 1967;*

same

Cosmopolitan

B. spongicola

Gruvel, 1920

probably a misidentification

B. tintinnabulum azoricus Pilsbry, 1916; Gruvel,

1920; Baker, 1967;*

Megabalanus azoricus

Azores

ZOOSYSTEMA • 1998 • 20(1)

Isopodes terrestres de Nouvelle-Calédonie.

II. Sur une nouvelle espèce du genre

Pseudolaureola (Crustacea, Oniscidea)

Henri DALENS

Laboratoire d’Écologie terrestre, Université Paul Sabatier

118 route de Narbonne, F-31062 Toulouse cedex (France)

Dalens H. 1998. — Isopodes terrestres de Nouvelle-Calédonie. II. Sur une nouvelle espèce

du genre Pseudolaureola (Crustacea, Oniscidea). Zoosystema 20 (1) : 93-100.

MOTS CLÉS

Crustacea,

lsopoda,

Oniscidea,

Nouvelle-Calédo n ie,

espèce nouvelle.

RÉSUMÉ

Une espèce nouvelle, Pseudolaureola deharvengi n.sp., est décrite de Nouvelle-

Calédonie (Province Nord, Kaala-Gomen). Elle est caractérisée par des

tubercules dorsaux particulièrement développés portant de longues et très

nombreuses soies. Une diagnose émendée du genre est proposée.

KEYWORDS

Crustacea,

lsopoda,

Oniscidea,

New Caledonia,

new species.

ABSTRACT

Terrestrial lsopoda from New Caledonia. II. A new species of tbe genus

Pseudolaureola (Crustacea, Oniscidea). A new species, Pseudolaureola dehar¬

vengi n.sp., is described from New Caledonia (Northern Province, Kaala-

Gomen). It is characterized by particularly well-developed dorsal tubercules,

bearing numerous long setae. An emended diagnosis of the genus is given.

ZOOSYSTEMA • 1998 • 20(1)

Dalens H.

En 1995. L. Deharveng et A. Bedos ont mené

des recherches de terrain dans différentes régions

du sud-ouest Pacifique. Parmi le matériel collecté

en Nouvelle-Calédonie figurait une nouvelle

espèce d’isopode terrestre, assez remarquable par

son ornementation. Sa description fait l’objet de

la présente note qui s’intégre dans une étude déjà

commencée sur la tau ne des isopodes terrestres

de Nouvelle-Calédonie (Dalens 1993).

Abréviations

MNHN Muséum national d’Histoire naturelle.

Paris ;

antennule ;

antenne ;

Mxl

maxillule ;

Mxp

maxillipède.

Famille ARMAD1LUDAE Verhoefif, 1917

Sous-famille Akermaniinae Vandel, 1973

Genre Pseudolaureola

Kwon, Ferrara et Tain, 1992

Espèce-type. — Pseudolaureola atlantica (Vandel,

1977).

Espèces incluses. — P. wilsmorei (Nicholls et Barnes,

1926), P. kystrix (Barnard, 1958), P. deharvengi n.sp.

Diagnose émendée

Pleurépimères, à l’exception du premier, et par¬

fois néopieu tons, prolongés en pointe donnant

un aspect foliacé aux représentants de ce genre ;

exopodites du premier pléopodc mâle très réduit

ou même absent ; exopodire de l’uropode absent

ou très réduit quand il est présent.

Pseudolaureola deharvengi n.sp.

(Figs 1-3)

MATÉRIEL EXAMINÉ — Nouvelle-Calédonie.

Province Nord, Kaala-Gomen, altitude 250 m, forêt

sèche sur calcaire, 18.XII.1995, A. Bedos &

L. Deharveng legs, 5 d <? et 7 ? 9 .

Matf.kièi.-TYI'L. — L'bolotype (MNHN-ls 5064),

conservé intact, est représenté par une femelle de

4,1 mm, ce qui constitue la taille maximale des échan¬

tillons examinés. L’holotype (MNHN-ls 5064) et une

partie des paratypes (MNHN-ls 5065) sont déposés

au Muséum national d’Histoire naturelle, Paris.

Quelques paratypes sont conservés dans la collection

de l'aureur. Des paratypes mâles de 2,2 mm paraissent

bien différenciés pour ce qui est des pièces génitales.

ÉTYMOLOGIE — Cette espèce est dédiée à mon col¬

lègue L Deharveng qui, avec A. Bedos, a récolté ces

échantillons.

RÉPARTITION. — Bien que de nombreux prélèvements

aient été effectués dans différentes régions de file,

cette espèce n’est connue que d’une seule station.

Description

Coloration

Les animaux vivants sont jaunâtres avec, par

endroits, des taches brunes. Les pleurépimères

sont toujours plus pâles. Il existe des variations

dans l'intensité de la coloration suivant les indivi¬

dus et le réseau pigmentaire est rarement homo¬

gène. En alcool, la coloration s’atténue fortement

et, par endroits, peut même disparaître totale¬

ment.

Appareil oculaire

Il est formé de neuf ommatidies (Fig. IA). Ce

nombre se réduit toutefois à six chez les indivi¬

dus de petite taille.

Morphologie somatique

Il s'agit d’une forme volvationnelle (Fig. IB),

dont le céphalon et le péréion Sont hérissés de

tubercules plus ou moins longs qui portent sur

toute leur surface de nombreuses et longues

soies. Le pléon est très raccourci et tombe subver-

ticalement, si bien qu’en vue latérale, ou même

dorsale, il passe pratiquement inaperçu.

Sur le céphalon (Fig. IC), les tubercules sont au

nombre de huit, disposés en deux rangées trans¬

versales. La première est formée de quatre tuber¬

cules très petits situés immédiatement en arrière

de la ligne frontale ; la seconde est également for¬

mée de quatre tubercules, mais beaucoup plus

développés et situés en avant du bord postérieur

du vertex, lequel est très nettement plus large que

long. La lame frontale dépasse de très peu le ver¬

tex, On relève par ailleurs une ébauche d’écus-

Fiô, 1. — Pseudolaureola deharvengi n.sp., A, massif oculaire

gauche : B, animai in loto en vue latérale ; C, céphalon vu de

face ; D, tubercule du premier pèréionite , E, système d'engre¬

nage volvationnel des premier et second péréionites en vue

ventrale ; F. Al gauche.

ZOOSYSTEMA • 1998 • 20(1)

250 pm

' j

; n

• u

• Al-

,j /

*, . 1

Dalens H.

son, dont le revêtement écailleux se différencie

nettement de celui du reste de la plaque frontale.

Sur les péréionites, les tubercules se disposent en

treize rangées parallèles :

1. Tout d’abord, une rangée médiane de tuber¬

cules situés près du bord postérieur de chacun

des péréionites ; très petits, avec l apex recourbé

vers l’arrière du corps, ils montrent néanmoins

un léget gradient de développement croissant

suivatu l’axe antéro-postérieur.

2. Ensuite, sur chaque demi-péréionite, on trouve,

à droite comme à gauche, six rangées de tuber¬

cules de développement très inégal ; on observe :

— Flanquant la rangée médiane, une rangée para-

médiane, que nous qualifierons d’interne, formée

de tubercules situés plus’ près du bord antérieur

du péréionite, de développement prononcé er

sensiblement identique sur tous les péréionites.

— Une rangée, que nous appellerons paramédiane

externe, formée de tubercules dont le développe¬

ment est légèrement plus réduit que chez les

tubercules de la zone médiane, de positionne¬

ment identique à ceux-ci, mais dont le dévelop¬

pement diminue progressivement suivant un axe

antéro-postérieur et qui n’est plus perceptible sur

le septième péréionite.

— Plus latéralement encore et au niveau de la

zone de courbure des tergires péréiaux, une ran¬

gée de tubercules très développés et qui croissent

régulièrement en taille de l’avant vers l’arrière du

corps ; croissance particulièrement nette sur les

trois derniers péréionites et que l’on peut quali¬

fier d’hypertélique sur le septième péréionite.

— Latéralement enfin, trois rangées, dont deux

sont parallèles et formées de petits tubercules

disposés sur le pleurcpimère, alors que la plus

externe se trouve tout à fait en bordure externe et

ceinture le corps de l'animal. Les deux rangées

internes sont de développement sensiblement

identique aux tubercules de la zone médiane. La

rangée la plus interne a les tubercules situés plus

près de la bordure postérieure des tergites, ceux

de la rangée externe sont, par contre, sirués tout

à fait antérieurement sur la bordure même du

péréionite.

3. Enfin, en ce qui concerne les tubercules de

bordure du corps, ils sont plus développés et sub¬

horizontaux. Sur le premier péréionite le tuber¬

cule s’étale d’ailleurs sur la majeure partie de la

bordure épimérale er forme une lame plate. Sur

les péréionites II et III, ces tubercules sont situés

sur la partie antérieure de la zone épimérale ; sur

les péréionites suivants, ils se déplacent progressi¬

vement vers la partie arrière de la zone épimérale.

Tous ces tubercules, grands ou perirs, portent sur

toute leur surface de longues soies (Fig. 1D), en

nombre d’autant plus grand que le tubercule est

plus développé.

L’ensemble des tergites comprend une partie pro-

tergale lisse, aussi développée que la partie méta-

tergale et normalement recouverte par le

métatergite du segmenr antérieur. Protergite et

métatergite sont séparés par une très légère crête,

formée de soies-écailles redressées qui paraissent

constituer en fait le bord antérieur du segment

péréial. Le bord postérieur de chaque métatergite

est également souligné sur toute sa longueur par

une crête de même type.

Le système d'engrenage (Fig. IL) est limité aux

deux premiers péréionites avec un schisma sur le

bord latéral du péréionite 1, lequel ne porte tou¬

tefois aucune trace de sillon longitudinal. La face

interne des pleurépimères I et II ne porte aucune

différenciation en forme de dent ou de replat qui

puisse être liée à la volvation. Sur le pléon où

existent premier et deuxième pléonites, même

s'ils Sont parfois difficilement visibles, l’ornemen¬

tation est réduite à quelques touffes de longues

soies, qui paraissent signaler des tubercules dont

le développement est si réduit qu’ils passent

inaperçus. Les néopleurons 3, 4 er 5 sont recour¬

bés à l’horizontale et leur extrémité porte

quelques soies.

Appendices

Al de trois articles (Fig. 1F), le second très court

er le troisième portant quatre aesthétascs, l’un à

mi-hauteur de l'article, les trois autres à l’apex.

A2 (Fig. 2A) courtes, massives et recouvertes

d’écailles imbriquées, avec un flagelle antennaire

de deux articles dont le distal, trois lois plus long

environ que le basal, porte deux aesthétascs.

Fie. 2. — Pseudolaureola deharvengi n.sp.. A, A2 gauche ;

B, Mxl ; C, telson en vue dorsale ; D, lelson en vue ventrale ;

E, 6 , basis VII en vue caudale ; F, S, basis VII en vue rostrale.

ZOOSYSTEMA • 1998 • 20(1)

250 pm

100 pm

Dalens H.

Mandibule droite avec deux pénicillés dont un

très petit presque accolé au pénicillé molaire ;

mandibule gauche à trois pénicillés, deux inter¬

médiaires et un molaire.

Mxl (Fig. 2B) avec endite interne portant deux

pénicillés assez massifs et endite externe pourvu

de onze dents toutes entières mais dont deux

sont très petites et difficilement visibles.

Fig. 3. — Pseudolaureola deharvengi n.sp., <3, exopodite (A) et

endopodites (B) de la première paire de pléopodes.

Mxp sans particularités notables. Pléopodes au

nombre de cinq paires, la première paire étant

toutefois très réduite chez le mâle. Vous les pléo¬

podes sont pourvus de pseudotrachées qui parais¬

sent rappeler certaines structures rencontrées

chez des Kubelidae (Ferrant, Paoli & laici 1990).

lelson (Fig. 2C, D) en forme de sablier, à partie

distale nettement plus étroite que la partie basale

et portant quelques soies très courtes. Basipodites

des uropodes très développés, endopodites relati¬

vement courts et n’atteignant pas le bord posté¬

rieur du tel s on ; quant aux exopodites situés

postérieurement sur le bord interne des basipo¬

dites, ils sont particulièrement réduits.

Caractères sexuels mâles

Au niveau des péréiopodes, une différenciation

particulièrement nette affecte les péréiopodes YII

du mâle, dont le basis (Fig. 2E, F) porte des

excroissances particulièrement développées. En

ce qui concerne les pléopodes, la première paire

montre des exopodites de développement très

réduit, tandis que les endopodites présentent une

partie basale Lies élargie (Fig. 3). l es endopodites

de ia seconde paire sont particulièrement longs et

fins (Fig. 4).

Discussion

Par sa morphologie, cette espèce se place mani¬

festement dans les Akermaniinae, sous-famille

d'Armadillidae créée par Vandel (1973), pour

ranger des formes que Fauteur pensait dérivées de

lignées différentes, mais parvenues selon lui au

terme de l’évolution armadilliennc, essentielle¬

ment caractérisée par un phénomène de devolva-

tion. Non valable sur un plan phyletique, cette

subdivision rassemble toutefois des forma, qui se

caractérisent routes par une ornementation très

développée de tubercules, d'épines ou même

d aiguilles.

En ce qui concerne 1 affectation générique, nous

rattachons cette espèce au genre Pseudolaureola

lvvvon, Ferrara et Taiti, 1992, en privilégiant

comme caractère discriminant majeur, celui rete¬

nu par Vandel (1977), à savoir l'aspecr foliacé et

la terminaison en pointe des pleurépimères et des

néopleurons.

Ce rattachement n’est peut-être que provisoire,

mais dans l’attente d’une meilleure connaissance,

ZOOSYSTEMA ■ 1998 • 20(1)

Une nouvelle espèce de Pseudolaureola

Fig. 4. — Pseudolaureola deharvengi n.sp , 6 , exopodite (A) et

endopodite (B) de la seconde paire de pléopodes.

sinon d’une révision des Akermaniinae, et tout

spécialement des espèces regroupées dans les

genres Akewuiniu Côllinge, 1919, Laure ola

Barnard, I960 et Pseudolaureola Kwon, Fcrrara et

Taiti. 1992 cette solution nous a paru préférable

à celle de la création d’un nouveau genre. Le pro¬

blème majeur concernant le rattachement de la

nouvelle espèce au genre Pseudolaureola était rela¬

tif à l'existence d’exopodites 1 mâles, certes

réduits, mais présents chez P. deharvengi , alors

qu’ils étaient indiqués comme absents dans la

description originale de f espèce-type P. atlavtica.

Un examen minutieux d’exemplaires de cette

dernière espèce, présents dans la collection

Vandel, nous a, en fait, convaincu que chez cette

espèce également, existaient chez le mâle des exo-

podites 1 certes très réduits et difficilement

observables mais réellement présents, ce qui ren¬

dait notre affectation générique tout à fait plau¬

sible. N'ayant toutefois pu réexaminer l'ensemble

des espèces regroupées dans ce genre et pour ne

pas préjuger des observations faites par les

auteurs précédents, nous avons donc modifié

légèrement la diagnose du genre donnée par

Vandel (1977).

Le nom générique de Pseudolaureola lut utilisé

pour la première fois par Schmalfuss & fcrrara

(1983) pour regrouper trois espèces que Vandel

avait auparavant réunies dans le genre l.auréola .

mais ces auteurs ne désignent pas pour autant

l’espèce-type. C’est donc Kwon, ferra ta & Taiti

(1992) qui valident ce nom en désignant

Laureola atlantica Vandel, 1977 comme l’espèce-

type. Outre fespèce-type qui vit dans l ile de

Sainte-Hélène, ils incluent dans le genre

Pseudolaureola, P. wilsmorei (Nicholls et Bames,

1926) de l'ouesE de l’Australie et P. hyslrix

(Barnard, 1958) de Madagascar, espèces aux¬

quelles il convient donc d’ajouter P. deharvengi

n.sp., de Nouvelle-Calédonie.

Remerciements

Le matériel étudié a été récolté dans le cadre du

programme Divers!tas-Nouvelle-Calédonie

animé par le Muséum national d’Histoire natu¬

relle de Paris. Nous remercions M. Papineau, res¬

ponsable du Service de l’Environnement de la

Province Nord, pour l’aide qu’il a apportée aux

collecteurs.

ZOOSYSTEMA • 1998 • 20(1)

Dalens H.

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italiano (N. S.) suppl. 18 (3): 111-157.

Vandel A. 1973. — Les Isopodes terrestres de

l’Australie : étude systématique et biogéographique,

Mémoires du Muséum national d’Histoire naturelle

(N. S.) série A, Zoologie 82 : 1-171.

— 1977. — Isopodes terrestres, m La faune terrestre

de Elle de Sainte-Hélène, Annales du Musée royal de

VAfrique centrale, Tervuren, Belgique (220) :

385-426, pl. Il h. t.

Soumis le 1" août 1997 ;

accepté le 13 novembre 1997.

100

ZOOSYSTEMA • 1998 ■ 20(1)

A new species of hermit crab of the family

Parapaguridae (Decapoda, Anomura)

from French Polynesia

Rafael LEMAITRE

Smithsonian Institution, National Muséum of Natural History,

Department of Invertebrate Zoology,

Washington, D C. 20560 (United States)

Lemaitre R-1998, — A new species of hermit crab of the family Parapaguridae (Decapoda,

Anomura) from French Polynesia. Zoosystema 20 (1) : 101-107.

KEYWORDS

deep-water hermit crab,

new species,

Parapaguridae,

Oncopagurus,

French Polynesia,

Tuamotu Archipelago.

ABSTRACT

A new deep-water hermit crab species of the family Parapaguridae,

Oncopagurus oimos, is described from Moruroa (French Polynesia, Tuamotu

Archipelago). The new species is distinguished primarily by ils distinctive

colour pattern (still visible in alcohol-prescrvccl spécimens after one yenr),

and consisting of broad stripes on the left cheliped and ambulatory legs.

Other diagnostic ch ara a ers of the new species are: (1) males lacking fïrst

pleopods and having reduced unpaired left second pleopods: (2) slender left

cheliped; (3) rounded mesia! lace of the right chela. The new species is com-

pared to another congeneric species, O. tuamotu Lemaitre, 1994 that also

occurs in French Polynesia, as well as to other species of the genus in which

males lack first pleopods.

MOTS CLÉS

pagure d’eau profonde,

nouvelle espèce,

Parapaguridae,

Oncopagurus,

Polynésie française,

archipel des Tuamotu.

RÉSUMÉ

Une espèce nouvelle de pagure de la famille des Parapaguridae (Decapoda ,

Anomura) de Polynésie française. Une espèce nouvelle de pagure d'eau profonde,

appartenant à la famille des Parapaguridae. Oncopagurus oimos, est décrite de

Moruroa (Polynésie française, archipel des Tuamotu). Cette espèce nouvelle

se distingue en particulier par la couleur (encore visible après un an de

conservation en alcool) des bandes longitudinales sur le chélipëde gauche et

les pattes ambulatoires. D'autres caractères distinctifs sont : (I) la présence,

chez les mâles, d’un seul deuxième pléopode gauche ; (2) le chélipède gauche

grêle ; (3) la face mésiale de la pince droite arrondie. Cette nouvelle espèce

est comparée à une autre espèce du genre, O. tuamotu Lemaitre, 1994 qui se

trouve également en Polynésie française, ainsi qu’aux autres espèces du genre

dont les mâles n’ont pas de premiers pléopodes.

ZOOSYSTEMA ■ 1998 • 20(1)

101

Lemaitre R.

Study of parapagurid samples, obcaîned frorn

French Polynesia since Lemaitres (1994) report

on the species of this family frorn the area, has

revealed rhe existence of an undescribed species,

This new species belongs in (lie recentiy propo-

sed genus Oncopagurus Lemaitre, 1996, the spe¬

cies of which are charactemed primarily by:

(1) presence of upward curved epistomial spine;

(2) phyllobranch gills; (3) operculate right chela;

(4) males having poorly to moderately developed

first and second pleopods, with first sometimes

absent (Lemaitre 1996).

In the material cxamined sectiotr, the length of

the shield of the spécimens is indicated in paren-

thesis, measured (to the nearest 0.1 mm) frorn

the tip of the rostrmn to the midpoint of the

posterior région of the shield. The terni “serni-

chelate" is used lor rhe fourrh pereopod in accor¬

dance wirh the définition provided by

McLaughlin (1997). The spclling of the island

localiry where this new species was found is

according to Motteler (1986). A summary of

deep-water collecting activities in French

Polynesia, including a list of crustacean species

and station data, can be found in Poupin (1996).

Oncopagurus oimos n.sp.

(Figs 1-3)

Material EXAMINED. — French Polynesia.

Tuamotu Archipelago, Moruroa atoll, Marara ,

stn 499, 21°47,6’S - 138'’55,7’W, trapped, 200 m,

5.V.1996: holutype d (2.8 mm) MNHN-Pg 5505;

paratypes. 1 d (2.4 mm), 1 9 (2.7 mm) MNIIN-

Pg 5506: I d (2.9 mm) USNM 276085.

Etymology. — J’he spécifie name is boni tlie Greek

oimos, meaning St ripe, used as a noun in apposition,

and refers to the distinctive colour pattern of this spe¬

cies.

Distribution. — Known so far only from Moruroa

atoll, on the Tuamotu Archipelago, French Polynesia.

Depth: 200 ni.

Habitat. — Gastropod shells.

Description

Shield (Fig. IA) as broad as long; dorsal surface

evenly calcified, with scattered short setae; ante-

rior margins weakly concave; latéral projections

subtriangular, terminating acutely; antérolatéral

margins slopitig; posterior margin broadly roun-

ded. Rostrum broadlv rounded, weakly produ-

ced, and with short mid-dorsal ridge.

Ventrolateral margins of shield each with small

spine (noL visible in dorsal view, and often only

on one side). Anterodistal margin of branchioste-

gite rounded, unarmed, setose.

Ocular pedundes more than ha If length of

shield, with dorsal row of setae. Cornea weakly

dilared. Ocular acides subtriangular, terminating

in strong spine; separated basally by less than

basal width of one acide.

Antennular pedunclc long, slender, exceeding

distal margin of cornea by entire length of ulti-

mate segment; ventral flagellum with five to six

articles. Ulrimate segment twice as long as penul-

rimate segment, wirh scattered setae Basal seg¬

ment with strong ventromesial spine; latéral face

with distal subrecrangular lobe armed with one

or two small spines, and strong spine proximally.

Antennal peduncle (Fig. 1 B) exceeding distal

margin of cornea by abour one tliird length of

fifrh segment. Flagellum long, exceeding exten-

ded right clieliped and ambulatory legs, articles

with setae less than one to two flagellât artides in

length (Fig. IC). Fifth segment unarmed, but

with scattered setae. Fourth segment with strong

spine on dorsolateral distal angle. Third segment

with strong ventromesial distal spine. Second

segmenc with dorsolateral distal angle produced,

terminating in strong, simple spine; mesial mar¬

gin wirh spine on dorsolateral distal angle. First

segment with unarmed latéral face; ventromesial

angle produced, with three to four small spines

laterally. Anrennal acide slightly curved outward

(in. dorsal view), at most slightly exceeding distal

margin of cornea, terminaring in strong spine;

mesial margin with row of eight to tbirteen

spines, setose.

Mandible (Fig. 2A) with three-segmenred palp.

Maxillule (Fig. 2B) with externâl lobe of endo-

pod weakly developed, internai lobe wirh long

terminal seta. Maxilla (Fig. 2C) with endopod

exceeding distal margin of scaphognathite. First

maxilliped (Fig. 2D) with endopod exceeding

exopod in distal extension. Second maxilliped

(Fig. 2E) with exopod about seven rimes as long

as broad. Third maxilliped (Fig. 2F) with distal

two segments each twice as long as wide; crista

102

ZOOSYSTEMA • 1998 • 20(1)

New parapagurid from French Polynesia

dentata with about rwelve corneous-tipped teeth;

coxa and basis each .with one tooth mesially.

Sternite of third maxillipeds with small spine on

each side of midline.

Chelipeds markedly dissimilar. Right cheliped

(Fig. 1D-F) massive, with sparse setation. Fingers

curved ventromesially, each rerminating in cor-

neous claw; cutting cdges with irrcgularly-sized

calcareous teeth, Dactyl approximately as long as

mesial margin of palm, and set at strongly

Fig, 1. — Oncopagums oimos n.sp., Marara, stn 499, Moruroa atoll: A-H, holotype 6 (2.8 mm) (MNHN-Pg 5505); I, paratype S

(2.9 mm) (USNM 276085). A, shield and cephalic appendages: B, right antennal peduncle (latéral view); C, proximal portion of fla-

gellum of same D. right cheliped (dorsal view): E, right chela of same (ventral view): F, merus and carpus of same (mesial view). G,

left cheliped; H, left second pleopod (arrow) and portion of pleuron (latéral view); I, left second pleopod (latéral view). Stippled areas

on shield (A), merus and carpus of right cheliped (F), and left cheliped (G) indicate reddish colour pattern. Scales bars: A D-G

1 mm; B, C. H, I, 0.5 mm.

ZOOSYSTEMA • 1998 • 20(1)

103

Lemaitre R.

oblique angle to longitudinal axis of pal ni; mesial

margin broadly curved, well defined by row of

spines (corneous-tipped on distal half); dorsal

face wirh scattered small spines proximally; ven¬

tral face smooth; ventromesial lace concave.

Fixed finger broad at base; latéral margin well

defined by row of Spines (corneous-tipped cm

distal half); dorsal face wich scattered small

spines or tubcrcles; ventral face smooth. Palm

broader titan long; latéral margin broadly roun-

ded, well delimited by row of blunt to sharp

spines; dorsomesial margin marked by row of

small spines; mesial face rounded, with small

spines or tubcrcles; dorsal surlace smooth except

for scattered sntall tubcrcles; ventral surface

smooth. Carpus wich latéral margin well delimi-

ted by row of spines; dorsal face with numerous

small spines; dorsodistâl margin with row of

spines; ventromesial distal margin slightly expan-

ded mesially, with row of spines: ventral face

with scattered small tubercles. Merus with dorsal

row of small spines; ventromesial margin with

row of spines. Ischium with ventromesial row ol

small spines. Coxa with one or two small spines

on ventromesial and ventrolateral margins, and

ventrotnesial row of serae.

Left cheliped (Fig. IG) slcnder, wich chela as

long as carpus, well calcilted. Fingers terminating

in small corneous ctaws; dorsal and ventral sur¬

faces unarmed cxcept for scattered tufts of setae;

cutting edge of dactyl and fixed finger each with

row of minuté corneous tecth. Dactyl subequal

to palm in length. Palm unarmed or at mosr wirh

inconspicuous tuhercle cm dorsomesial angle,

and scattered setae; ventral face smooth except

for scattered setae. Carpus with strong dorsodis-

tal spine; dorsal margin with modérait secarion;

ventral face smooth Merus with setae on dorsal

margin; ventrolateral distal margin with row of

spines; ventral face smooth. Ischium unarmed,

with setose ventral face. Coxa with one spine on

ventromesial and ventrolateral distal margins,

and ventromesial row of serae.

Ambulatory legs (Fig. 3A-C) similar from right

to left, reaching. approximately to, or just

beyond, tip of extended right cheliped. Dactyl

slightly less than twicc as long-as propodus, ter¬

minating in sharp corneous claw; with dorsal and

dorsomesial rows of setae, and ventromesial row

of about ten to fourtecn corneous spines; latéral

and mesial faces with shallow, longitudinal sulcus

on proximal half (deeper on mesial face).

Propodus wirh row of setae on dorsal and ventral

margins. Carpus with small dorsodistâl spine and

setae dorsally and Ventrally. Merus unarmed,

with setae on dorsal and ventral margins.

Ischium and coxa unarmed. Anterior lobe of

srernite of third pereopods setose, subsemicircu-

lar, armed with simple terminal spine.

Fourrh pereopod (Fig. 3E) ÿemichclate. Dactyl

terminating in sharp corneous claw; with ventro¬

lateral row of small corneous spinules. Propodus

longer than dûrsoventral width, rasp fonned of

one tow of rounded scales. Carpus with long

setae on dorsal margin. Merus with rows of long

setae on dorsal and ventral margins.

Fifth pereopod (Fig. 3F) chelate. Propodal rasp

extending to mid-length of segment.

Umpods and telson (Fig. 3G-1) markedfy asym¬

metrical. Telson lacking transverse suture; dorsal

surface with scattered setae; terminal margin

with shallow médian cleft separating left and

right lobes, each wirh several marginal corneous

spines.

Male lacking first pleopods; with unpaired redu-

ced second left pleopod (Fig. 1H, I) consisting of

minute bud or very short segment with few ter¬

minal serae. Female lacking vestigial second right

pleopod,

Colour pdltem (Figs IA, F, G, 3A, B)

No observations of colour in tife were rnade of this

species. However, after approximately one year in

alcohol, a distinctive reddish colour pattern was

sfill visible in the specimens. The shîeld lias two

small reddish spocs (Fig. IA) on the anterior half,

one juSt behind each latéral projection. On both

chelipeds (Fig. 1F, G), the dorsodistâl portions of

the meri are reddish. The carpus of the right cheli¬

ped has a reddish atea on the proximal portion of

the latéral and mesial faces. The carpus of the left

cheliped has a distinctive broad reddish .stripe on

the dorsolateral and mesial laces; the dorsomesial

margin of the palm is light reddish; the dactyl is

reddish. The ambulatory legs (Fig, 3A, B) hâve a

reddish colour dorsodislally on the meri; die carpi

each hâve a reddish stripe on the latéral and mesial

faces, and the stripes of the carpi continue on the

104

ZOOSYSTEMA • 1998 • 20(1)

Lemaitre R.

meri but only on the latéral faces; the dactyls

hâve reddish dorsal faces.

Remarks

This new species is the fifth in the genus

Oncopagurus in which males lack first gonopods.

The other four are O. haigae de Saint Laurent,

1972 (Eastern Pacific); O. orientaUs de Saint

Laurent, 1972 (Indo-Pacific); O. tuamotu

Lemaitre, 1994 (known only front the Tuamotu

Archipelago); and O. cidaris Lemaitre, 1996

(known only from Australia). The males of these

Fig. 3 — OiKopagums olmos n.sp., Marara, stn 499. Moruroa atoll: holotype S (2.8 mm) (MNHN-Pg 5505). A, lett second pereopod

(latéral view): B, lett third pereopod (latéral view); C, dactyl of same (mesial view); D, sternite of third pereopods (ventral view): E, left

fourth pereopod (latéral view); F, propodus and dactyl of left fitth pereopod (latéral view); G, exopod of left uropod (dorsal view);

H, telson (dorsal view); I, exopod of right uropod (dorsal view). Stippled areas on second (A) and third (B) pereopods indicate reddish

colour pattern. Scale bars: A-D, 1 mm; E-l, 0.5 mm.

106

ZOOSYSTEMA • 1998 • 20(1)

New parapagurid from French Polynesia

four species hâve paired asymmetrical second

gonopods; liowever, Lemairre (1996, 1997) has

documenred incra-specific variation in males of

O. orientalh and O. cidurü , wItéré the second right

gonopod is occasionally lacking. l’hc dtree known

males of O. oimos n.sp. hâve only reduced, simple

unpaired left second plcopods. The segmentation

of the male second pleopods in thcse fivc species

differs. The second pleopods are unsegmemed in

O. oimos n.sp., O. bnigae and O. attenta Us-, they

are two-segraented in O. tuamotu ; and one- or

two-segmenred in O cidaris. Ofall known species

of Oncopagurus , O. oimos n.sp. is the only one in

which females hâve only unpaired left second

pleopods; fèraales of ail other species hâve vestigial

right second pleopods as vvell.

The distinctive colmu pattern of O. oimos n.sp.,

can be used to distinguish it from O. tuamotu,

the only other congeneric species known from

French Polynesia. The new species has broad

stripes on the left cheliped and ambulatory legs

(Figs IG, 3A, B), whereas O. tuamotu has red

bands on the cheliped and walking legs (see

Lemaitre 1994: 411, fig. 281). In addition to the

REFERENCES

Lemaitre R. 1994. — Crustacea Decapoda:

Deep-water hermir crabs (Parapaguridae) from

French Polynesia vvith descriptions of four new

species, in: Crosniet A. (ed.), Résultats des

Campagnes MUSORSTOM, Volume 12,

Mémoires dit Muséum national d'Histoire naturelle

161 : 375-419.

— 1996. — Hennit crabs of the farnily Parapaguridae

(Crustacea: Decapoda: Anomura) from Australia:

species ot Strobopagurits Lemaitre, 1989,

Sympagurus Smith. 188.3 and two new généra.

Records of the Australian Muséum 48: 163-221.

— 1997. — Crustacea Decapoda: Parapaguridae

from the Karubar Ctui.se in Indonesta. wirh des¬

criptions of two new species, in Crosniet A. &

previously meniioned différences in pleopod

conditions, other characters can also be used to

distinguish O. oimos n.sp. from O. tuamotu. The

mesial face of the right palrn is rounded in the

new species, with a wealc irregular dorsomesial

row of spines, whereas the mesial face is expan-

ded distally, and rhere is a distinct dorsomesial

and ventromesial tows of spines in O. tuamotu.

The left chela is noticcably more slender, as long

as the carpus, and the fingers arc set tiearly paral-

lel to the longitudinal axis of the palm in the

new species; the chela is shorter than the carpus,

and the fingers are set obliquely (pointing ven-

trolaterally) to die longitudinal axis of the palm

in O. tuamotu.

Acknowlc d gemen ts

The collecring efforts of Joseph Poupin in French

Polynesia hâve been invaluable in documenting

the crustacean fatma from this area. Flis efforts

are gratefulJy acknowledged. I thank also Alain

Crosnier for lostering this and other studies

based on the important material gathered during

many recent French expéditions.

Bouchet P. (eds), Résultats des Campagnes

MUS ORSTOM, Volume 16, Mémoires du

Muséum national d'Histoire naturelle 172 :

573-596,

Mcl .mghlin P, A. 1997. — Cmstacea Decapoda: her-

mit crabs of the family Paguridae from the Karubar

Expédition in Indonesi», fit Crosniet A. & Bouchet

P. (eds). Résultats des Campagnes MUSORS-

FOM, Volume 16 (5). Mémoires du Muséum natio¬

nal d'Histoire naturelle 172 : 433-572.

Motteler L. S. 1986. — Pacific isl-and names. A niap

and name guide to the New Pacific. Bishap

Musetjtn Miscellt-neous Publication 34: 1-91.

Poupin J. 1996. — Allas des crustacés marins profonds

de Polynésie française. Récoltes du navire Marara

(1986/1996). Service Mixte de Surveillance

Radiologique et Biologique de I homme et de

I cnvironnement, Montlhéry, 59 p.

Submitted on 30 May 1997;

accepted on 8 September 1997.

ZOOSYSTEMA • 1998 • 20(1)

107

Phyllotymolinidae, nouvelle famille

de Brachyoures Podotremata

(Crustacea, Decapoda)

Marcos TAVARES

Universidade Santa Ürsula, Instituto de Ciências Biolôgicas e Ambientais

22231-040 Rio de Janeiro (Brasil)

Tavares M. 1998. — Phyllotymolinidae, nouvelle famille de Brachyoures Podotremata

(Crustacea, Decapoda). Zoosystema 20 (1) : 109-122.

MOTS CLÉS.

Crustacea,

Decapoda,

Brachyura,

Podotremata,

nouvelle famille,

faune bathyale.

RÉSUMÉ

Une nouvelle famille de crabes, les Phyllotymolinidae, est décrite pour les

genres Phyllotymolinum Tavares, 1993, le genre-type, Genkaia Miyaké et

Takeda, 1970 et Lonchodactylus Tavares et Lemairre, 1996. Quatre espèces

sont incluses dans la nouvelle famille : Phyllotymolinum crosnieri Tavares,

1993 (Nouvelle-Calédonie er Vanuatu, 280-301 m) ; Genkaia gordonae

Miyaké et 1 akeda, 1970 (Japon, Tsushinta, nord-est de Kyushu, 68-100 m) ;

Genkaia keijii Tavares, 1993 (Nouvelle-Calédonie, 135-150 m) et

Lonchodactylus messinyi Tavares et Lemairre, 1996 (Bahamas, 262 m).

KEYWORDS

Crustacea,

Decapoda,

Brachyura,

Podotremata,

new family,

bathyal fauna.

ABSTRACT

Phyllotymolinidae, new family of Brachyura Podotremata (Crustacea ,

Decapoda). A new crab family, Phyllotymolinidae, is described to accommo-

date the généra Phyllotymolinum Tavares, 1993, the type genus, Genkaia

Miyaké et Takeda, 1970 a ntl Lonchodactylus Tavares et Lemairre. 1996. Four

spccies are included in the new family: Phyllotymolinum crosnieri Tavares,

1993 (New Caledonia and Vanuatu, 280-301 m); Genkaia gordonae Miyaké

et Takeda, 1970 (Japon, T sushinta, north-easrern Kyushu, 68-100 m);

Genkaia keijii Tavares, 1993 (New Caledonia, 1,35-150 m); and

Lonchodactylus messingi Tavares efLemaitre, 1996 (Bahamas, 262 m).

ZOOSYSTEMA • 1998 • 20(1)

109

Tavares M.

INTRODUCTION

Une nouvelle famille, celle des Phyllotymolinidae

fam. nov., est établie ici pour recevoir les genres

Genkaia Miyaké et Takeda, 1970, Phyllotymo-

linum Tavares, 1993 et Ltmehvdactylus Tavares et

Lemaitre, 1996. Des trois genres mentionnés,

seul Genkaia n’a pas été originalement décrit

dans les cyclodorippidés. Lors de sa description,

Miyaké &C Takeda (1970 : 27) puis Takeda

(1973 : 77) ont vu en Genkaia un proche parent

des genres dromiens Uypoconcba Guérin-

Méneville, 1854 et Conchoedromia Chopra,

1934, et l’ont provisoirement rattaché aux

Dromiidae. C’est Tavares (1993a : 281) qui a

montré qu’il n’existe pas de liens immédiats entre

Genkaia et les Dromiaeea et l’a transféré dans les

Cyclodori ppi dac.

Chez tous les Cyclodorippidae la disposition de

l’endostome et des appendices bucaux peut être

qualifiée d’ox)'stome, avec la fermeture du plan¬

cher buccal par les seuls troisièmes maxillipèdes

(Mxp3). On notera que le palpe articulé sur la

face interne du mérus de Mxp3 améliore la coap¬

tation entre celui-ci et 1’endosrome En revanche,

les modifications pour la respiration rencontrées

chez les Phyllotymolinidae concernent surtout

les antennes, dans leur uualitc. L’endostome esc

brachygnathe ; le bord latéral du cadre buccal er

les Mxp3 s’étendent vers l’avant, mais en demeu¬

rant rectangulaires. 11 n’y a aucune coaptation

entre les Mxp3 et l’endostome, L’appareil respira¬

toire des Phyllotymolinidae ne peut donc pas

être considéré comme une disposition oxyscome.

Les structures impliquées dans le système respira¬

toire des Cyclodorippidae et des Phyllotymolini-

dae ne sont pas homologues et ne peuvent pas

dériver l’un de l’autre. Ce sont principalement les

importantes novations uniques chez les

Brachyoures, fournies par la chambre prosro-

miale chez Phyllotymolinum, Genkaia et Loncho-

dactylus qui imposent la création de la nouvelle

famille des Phyllotymolinidae.

La terminologie utilisée pour désigner les diffé¬

rentes structures de la carapace est celle de

Tavares (1991b). Les abréviations suivantes ont

été employées pour certains appendices :

Mxpl, 2, 3

P2, P3, P4, P5

Pli, P12

MNHN

ZMA

maxillipèdes 1,2 et 3 ;

péréiopodes 2, 3, 4 et 5 ;

pléopodcs ! et 2 ;

Muséum national d’Histoire

naturelle, Paris ;

Zoôlogisch Muséum, Amsterdam.

PHYLLOTYMOLINIDAE fam. nov.

Dromiidae - Miyaké & Takeda 1970 : 26. — Takeda

1973a : 77 (p)‘0 parte) ; 1973b : 22 (pro parte) ; 1985:

97. - Sakai 1976 : 7 [proparte).

Cyclodorippidae — Tavares 1993a : 280, 285 ( pro

parte). - Tavares & Lemaitre 1996 : 464. — Tavares

1997 : 262 {pro parte).

GENRE-TYPE. — Phyllotymolinum Tavares, 1993.

Genre neutre. Espèce-type Phyllotymolinum cromieri

Tavares, 1993 par désignation originale et par mono-

typie.

GENRES INCLUS. — Phyllotymolinum Tavares, 1993 ;

Genkaia Miyaké et Takeda, 1970 ; l.nnchodactylus

Tavares et Lemaitre, 1996.

Espèces INCI.USIiS. — Phyllotymolinum cromieri

l avares, 1993 ; Genkaia gordonae Miyaké et Takeda,

1970 ; Genkaia keijii Tavares, 1993 ; Lonchudactylus

nu'ssrngi Tavares er Lemaitre, 1996.

Distribution. — Même si les connaissances actuelles

sur les Phyllotymolinidae sont fort limitées, il semble

que, à l'exemple des Cyclodorippidae (Tavares 1997),

l’Atlantique américain et l’océan Indo-Ouest-

Pacifique possèdent une faune phyllotymolinicnne qui

leur est propre. Des trois genres connus de

Phyllotymolinidae, deux sont exclusivement indo-

ouesr-pacifiques : Genkaia (G. gordonae. Japon,

Tsushima, nord-est de Kyttshu, 68-100 m ; G. keijii ,

Nouvelle-Calédonie, 135-150 ni) er Phyllotymolinum

(P. cmnieri , Nouvelle-Caledonie et Vanuatu, 280-

370 m). Lnnchodactylus est entièrement américain

(L. messmgi. Bahamas, 262 m).

Description

Carapace bien calcifiée, assez plane, peu sculptée.

Fronr subtriangulaire. Limites entre la face dor¬

sale de la carapace et les flancs assez nettes.

Largeur fronto-orhitaire supérieure à la moitié de

la largeur maximale de la carapace.

Cavité orbitaire communiquant librement avec la

cavité antennaire ; le lobe infra-orbitaire,

médiocre, ne sépare nullement l’une de l’autre les

cavités oculaire et antennaire. Bords supérieur et

110

ZOOSYSTEMA • 1998 • 20(1)

Phyllotymolinidae, fam.nov.

inférieur de l’orbite bien délimités. Dent exorbi¬

taire proéminente. Pédoncules oculaires mobiles,

courts, orientés transversalement par rapport à

Taxe de la carapace. Antennes remarquablement

robustes ; premier article antennaire mobile ;

articles 2 + 3 en forme d’un grand auvent.

Avancée de l’endostome très courte. Bords laté¬

raux du cadre buccal assez allongés, atteignant le

segment 2 + 3 de l'antenne.

Troisièmes maxillipcdcs se coaptant parfaitement

avec le bord latéral du cadre buccal. Exopodites

des trois paires de maxillipèdes pourvus d’un fla¬

gelle normal. Exopoditc de Mxp3 avec l’extré¬

mité effilée, très long, atteignant l'extrémité

distale du mérus ; palpe articulé h l'angle antéro-

externe du mérus, ses articles 2 et 3 comprimés

dorso-ventralement. Propode de l’endopodite de

Mxp2 sans avancée.

Chélipèdes égaux, P2 et P3 similaires, peu séti-

fères, faiblement ornés ; dactyle comprimé latéra¬

lement. P4 et P5 également similaires, mais

ischion de P3 deux fois plus long que celui de

P4 ; P4 et P5 réduits, sauf leur coxae, et subdor¬

saux ; dactyle préhensile, se fermant sur la face

latérale du propode.

Cavité sterno-abdominale peu délimitée et exca¬

vée. Abdomen femelle fbrGJé de sept segments,

tous assez élargis. Premier segment abdominal

couvrant tout l’espace entre les coxae des P5.

Chez les femelles de Genkdid er Lonchoditctylus,

pléopodes 1 (Pli) de la femelle vestigiaux, unira-

més, insérés sur la face ventrale du premier seg-

Fig. 1. — A. B, Phyllotymolinum crosnieri Tavares, 1993, S holotype. 5,7 x 6,9 mm (MNHN-B 24617) ; A, vue ventrale de la région

antérieure du cadre buccal ; B, face externe des segments abdominaux 1 à 6, et du telson ; C, Ketamia depressa (Ihle, 1916)

(Cyclodorippidae), ? paralectotype, 4 x 4,3 mm (ZMA-De 102973). id. à A.

ZOOSYSTEMA • 1998 • 20(1)

Tavares M.

Fig. 2. — A, B, Phyllotymolinum crosnieri Tavares. 1993, 4 X 4,5 mm (MNHN-B 24618) ; A, vue d'ensemble de la région frontale

de la carapace ; B, détail de la région antennaire : à noter les bords du cadre buccal allongés jusqu'à toucher le deuxième article

antennaire. Irès développé et en forme d'auvent ; C. D, Xeinostama eucheir Stebbîng, 1920 (Cyclodorippidae), 9, 5 x 5,5 mm

(MNHN-B 24717) : C, vue frontale de la carapace ; D. détail des régions antennaire el oculaire. À noter les différences entre l'endo-

stome de P. crosnlen et celui de X. eucheir.

ment abdominal. Pli absents chez la femelle de

Phyllotymolinum. Pléopodes 2 à 5 biramés, P15

plus court que les pléopodes 2 à 4, non ovifère.

Uropode absent chez la femelle et chez le mâle.

Abdomen mâle de sept segments (jusqu'à présent

le mâle de Lonehoductylus messingi n’est pas

connu).

Novations et affinités

DES PHYLLOTYMOLINLDAE

Il n’est pas question de mener dans cette note

une analyse approfondie des affinités entre les

Phyllotymolinidae fam. nov. et les- autres

Podotrentata. Une analyse cladistique des

familles de Podotremata est en cours de prépara¬

tion (Guinot & d'avares). Toutefois, de la

confrontation des connaissances accumulées au

cours des études sur les Cyclodorippidae (Tavares

1991a, b, 1992a, b, c, 1993a, b, 1994, 1996,

1997 ; Tavares & Lemaitre 1996), dont les

Phyllotymolinidae sont assez proches, nous avons

dégagé certaines idées que nous avons jugé utile

d’exposer dès à présent.

Chambre prostomiale

Les Phyllotymolinidae ne sont pas les seuls

Podotremata à posséder une chambre prosto-

miale. Une telle chambre existe, par exemple,

chez tous les Cyclodorippidae. Cependant, il faut

bien faire la distinction entre les structures et

appendices qui sont impliqués dans la formation

de la chambre prostomiale, d une part chez les

Cyclodorippidae et d’autre part chez les Phyllo¬

tymolinidae.

La chambre prostomiale des Cyclodorippidae est

constituée par le plancher de l’endostome recou¬

vert par les appendices suivants : mandibules,

maxillules, maxilles et maxillipèdes. L’endostome

112

ZOOSYSTEMA • 1998 • 20(1)

Phyllotymolinidae, fam.nov.

Fig. 3. — A. B. Phyllolymolinum crosnieri Tavares, 1993, v, 4 x 4.5 mm (MNHN-B 24618). vues dorsale (A) et ventrale (B) du

Mxp3 ; à noter l'exopodite muni de ilagelle et le palpe articulé sur le bord interne du mérus ; C, D, Clythrocerus nitidus (A. Milne

Edwards. 1880) (Cyclodorippidae), rf, 4 x 4,5 mm (MNHN-B 13489), vues dorsale (C) et ventrale (D) du Mxp3 ; à noter l'exopodite

dépourvu de flagelle et le palpe articulé sut la face interne du mérus.

joue, dans tous les genres de la famille, un rôle parfaite de chaque endopodite des Mxp3

primordial pour la formation de cetre chambre (Fig. 3D). L’exopodite, lui aussi effilé di.stalement,

prostomiale. Il est généralement très allongé et esr presque toujours dépourvu de flagelle

forme une gouttière, rétrécie vers l avant et qui (Fig. 3C, D), ce qui améliore la coaptation entre

peut même dépasser de beaucoup le bord frontal l’exopodite et les bords du cadre buccal. Lépipo-

de la carapace (Figs IC, 2C). Comme relevé par dite de Mxp3 est absent cher: tous les Cyclo-

Ortmann (1892 : 552) et par Ihle (1916b : 110, dorippïdae. L.e Mxp2 conserve une forme plus

fig. 62), le Mxp3 épouse la forme de la gouttière habituelle (Fig. 4B) et agit, probablement,

endostomienne : il esr exrrémement allongé et comme un filtre. Là aussi, l’exopodite est parfois

son mérus se termine par une extrémité distale dépourvu de flagelle. L’endopodire de Mxpl est

avancée et effilée (Fig. 3C) ; le palpe est articulé généralement plus allongé que l'exopodite et

non pas sur l’angle antéro-externe du mérus, mais s’enroule vers son extrémité distale, comme l’a

à sa face interne, permettant une coaptation plus bien vu Ihle (1916b : 109, fig. 57) ; souvent

ZOOSYSTEMA • 1998 - 20(1) 1 13 I

Tavares M.

114

ZOOSYSTEMA • 1998 • 20(1)

Phyllotymolinidae, fam.nov.

l’exopodite ne porte pas de flagelle. La maxille et

la maxillule ne montrent pas de transformations,

mais les mandibules sont modifiées en fonction

du type de respiration oxystome : elles se coaptent

entre elles, et leur forme très allongée (Fig. 4C,

D) aide à guider le courant d'eau vers I avant.

Chez les Phyllotymolinidae la chambre prosto-

miale est formée surtout du troisième maxilli-

pède, du bord latéral du cadre buccal, de la

mandibule et, cas unique chez les' Brachyoures,

de l’antenne. Les bords latéraux du cadre buccal

s allongent jusqu’à toucher les deuxième, troi¬

sième et quatrième articles atiten irai res (Figs IA,

2A, B). Ceux-ci sont assez développés et en

forme d’auvent (très caractéristique chez Phyllo-

tymolinum , Fig. 2B), constituant avec les bords

latéraux du cadre buccal une sorte de couloir,

rétréci vers l avant, remplaçant, du point de vue

fonctionnel, la gouttière endostomienne décrite

ci-dessus. On notera que les modifications du

deuxième article atrtennaire (Fig. 5A, B) offertes

par certains genres de Cyclodûrippidae

(Deilocertis y Corycodus, Neocorycodus et Clythro-

cerus) ne sont nullement de même uature que

celles trouvées chez le.s Phyllotymolinidae et,

d’ailleurs, elles n’ont aucune relation avec la for¬

mation d’une chambre prostomiale. Chez les

Phyllotymolinidae, les Mxp3 sont d'un type

ordinaire, si ce n’est qu ils se coaptent parfaite¬

ment entre eux et avec les bords du cadre buccal,

assurant ainsi la fermeture de la chambre prosto¬

miale (Fig. 3A, B). Ici, l'eudostotne ne joue pas

un rôle essentiel : il est court et n’est pas en

forme de gouttière (Figs IA, 2A). Chez Genkaia,

la mandibule ne participe pas à la formation de

la chambre prostomiale ; en revanche, chez

Phyllotymolirium, la mandibule est tour aussi

allongée que celle des Cyclodorippidae.

Fermeture du dactyle de P4 et de P5

Comme d ordinaire chez les Brachyoures (voir

aussi Guinot 1968), c’est toujours par deux

Fig. 4. — A. Phyllolymolinum crosmeri Tavares, 1993, ?, 4 •

4,5 mm (MNHN-B £4618). détail de l'endopodile de M*p2 : à

noter le propode sans avancée ; B-D, Glytdrocerus nilidus

(A. Milne Edwards, 1880) (Cyclodorippidae), i, 4 x 4,5 mm

(MNHN-B 13489) ; B, détail de tensembte propode-daotyle ; a

noter l'avancée du propode orné à peu près comme le dactyle ;

C, D, mandibules gauche et droite, vues dorsale (C) et ventrale

(D) : à noter la forme très allongée.

Fie. 5. — A. Xeinostoma eucheir Stebbing, 1920 (Cyclodorippi¬

dae).. V, 5 s 5.5 mm (MNHN-B 24717), détail des quatre pre¬

miers articles antermaïres ; B, Clythrocerus nilidus (A. Milne

Edwards, 1880) (Cyclodorippidae), ,-î, 4 x 4,5 mm (MNHN-

B 13489), vue d'ensemble de l'antenne

condyles que les articles des pattes thoraciques

s’articulent entre eux (ou au corps, dans le cas de

la coxa) Le plan d'articulation est alterné : la coxa

s’articule au corps par un condyie ventral et par

un condyie dorsal ; le basis s’articule à la coxa par

deux condyles latéraux ; l’articulation enrre

l’ischion et le basis se lait à nouveau par un

condyie ventral et par un condyie dorsal, et entre

l’ischion et le mérus par deux condyles latéraux,

et ainsi de suite. L’articulation à condyles latéraux

entre deux articles permet une mobiliré nette¬

ment plus importante que celle où la jointure des

articles est dorso-ventrale. Chez les Phyllotymo¬

linidae, P4 et P5 offrent, malgré leurs fortes

modifications, un plan d’articulation qui reste

rigoureusement le même. Cependant, c’est sur la

face latérale du propode que les dactyles de P4 et

ZOOSYSTEMA • 1998 • 20(1)

115

Tavares M.

Fig. 6. — A, B, Xeinosloma eucheir Stebbing, 1920 (Cyclodorippidae), 9, 5 x 5,5 mm (MNHN-B 24717) ; C, D, Phyllotymolinum

crosnieri Tavares, 1993, 9. 4 x4,5 mm (MNHN-B 24618) : péréiopode 5 ; à noter le propode articulé sur le carpe par deux condyles

dorso-ventraux et la torsion du propode et, en conséquence, le dactyle articulé sur le propode par deux condyles également en posi¬

tion dorso-ventrale.

de P5 se ferment. Le propode est parfois muni

d un petit tubercule ou de fortes soies» ce qui aug¬

mente la capacité préhensile de l’ensemble propo-

de-dactyle (Fig. 6D). Rappelons qüc l'articulation

entre le propode et le dactyle est assurée par deux

condyles latéraux, ce qui assure beaucoup de sou¬

plesse mais impose, normalement, une fermeture

du dactyle sur la face ventrale du propode et non

pas sur sa face latérale. Néanmoins, chez, tous les

Phyllotymolinidae (ainsi que chez certains

Podotremata), la moitié distale du propode de P-i

et de P5 est tordue : les deux condyles latéraux

assument de ce fait une position presque dorso-

ventrale (Fig. 6A-D), ce qui permet au dactyle de

se fermer sur la face latérale du propode, tout en

gardant une très forte mobilité. C’est ainsi que le

crabe acquiert la possibilité de tenir un objet au-

dessus de la carapace.

Appareil de maintien de l'abdomen

Efficace, l’appareil de maintien de l’abdomen des

Phyllotymolinidae est bien différent des systèmes

décrits par Guinot (1979) pour la plupart des

familles de Podotremata où la rétention de

l’abdomen est souvent assurée par la coxa des

divers appendices. Le fait que chez les

Phyllotymolinidae le système de maintien (sys¬

tème à butoir) de l’abdomen fait appel à des élé¬

ments anatomiques différents (novation sur le

sternitc 6) de ceux des Cyclodorippidae (système

à glissière, Figs 7A-C, 8), conduit à considérer

l’appareil de rétention des Phyllotymolinidae

116

ZOOSYSTEMA • 1998 • 20(1)

Phyllotymolinidae, fam.nov.

Fig. 7. — A D, Clythrocerus nitidus (A. Milne Edwards, 1880) (Cyclodorippidae), â, 4 x 4.5 mm (MNHN-B 13489) A, vue

d’ensemble du sternum thoracique ; B, détail de la cavité sterno-abdominale ; C, détail de l'un des bords latéraux de la cavité stemo-

abdominale ; D. vue d’ensemble du PII. Â noter, en A et B, que le sternum, assez large, n’est pas entièrement recouvert par l’abdo¬

men quand celui-ci est replié sous le céphalothorax.

ZOOSYSTEMA • 1998 • 20(1)

117

Tavares M.

Fig. 8. — Clythrocerus nitidus (A. MÎIne Edwards, 1880)

(Cyclodorippidae), 4 x 4,5 mm (MNHN-B 13489), les trois

derniers segments abdomianux. A noter que la cavité sterno-

abdominale fait saillie au niveau des P3 (Fig. 7B. C) et retient

l’abdomen en l'empêchant de se soulever. De plus, le moulage

entre le dernier segment abdominal et cette paille du sternlte 6

rend ce système plus efficace.

comme non homologue de celui des autres

Podotremata. Chez Phyllatymolinum le telson est

très élargi à la base, il dépasse largement de

chaque côte le bord antérieur du segment 6

(Fig. IB). Le sternite 6 possède de chaque côté,

au niveau de la coxa de P3, deux saiilies en tonne

d’éperon obliques, bordées par une gorge. Lors

du mouvement vers l avant de l'abdomen, l’épe¬

ron sert de butée à l’expansion postéro-latérale

du telson qui va pouvoir s'engager dans la gorge.

Une ébauche de moulage entre l’abdomen et le

sternum rend ce mécanisme plus parfait. Seul, un

mouvement vers Lavant, suivi d’un soulèvement

de l’abdomen, permet à celui-ci de se désengager.

Chez Genkaitl, le système de rétention est simi¬

laire, sauf que dtaque éperon sur le sternite 6 est

remplacé par une sorte de crête ornée de gra¬

nules. Le mâle de L. messingi étant inconnu à

l’heure actuelle, nous ne disposons d’aucun ren¬

seignement sur l’appareil de maintien de l abdo-

men chez le genre Lanchodactylus.

Chambre incubatrice

Les Phyllotymolinidae femelles possèdent des

segments abdominaux assez calcifiés, très bom¬

bés, avec des bords incurvés vers le bas. Ainsi la

ponte est entièrentenr enserrée dans la sorre de

boîte ainsi formée. Chez Pbyllotymolitmm , les

exopodites des pléopodes, assez larges, assurent la

fermeture latérale de la boîte (Fig. 9D). On

observe une tendance vers la réduction du

nombre de pléopodes (chez PhyUotymolinum, pas

de Pli) et de leur taille. Les Phyllotymolinidae

sont les seuls Podotremata à avoir, chez la femelle,

un P13 très réduit et non ovifère.

Cyclodorippoîdea

il n’est pas envisageable de considérer les Phyllo¬

tymolinidae comme une sous-famille de Cyclo-

dorippidae. Outre les différences étudiées

ci- dessus, il existe bien d’autres rraits distinctifs

concernant notament la formation de la cavité

stemo-abdominale, le nombre de segments abdo¬

minaux, les pléopodes chez la femelle. Nous ne

décelons aucune synapomorphie qui permettrait

de réunir les Phyllotymolinidae ec les Cyclo¬

dorippidae. D'ailleurs, dans l’état de nos connais¬

sances actuelles, la notion de Cyclodorippoîdea

(Cymonomidae +- Cyclodorippidae + Phyllo¬

tymolinidae) est à vérifier.

Clé pour les genres et espèces de Phyllotymolinidae fam. nov.

1. Ouvertures de la spermathèque presque contiguës et non indiquées par une bosse

bien distincte . Lonehodaaylns messingi

2. Ouvertures de la spermathèque très écartées, indiquées par une bosse bien distincte..

3. Pli absents chez la femelle. Exopodite de P12-5 foliacé. Sternites thoraciques 6-7

contigus . Phyllotymolinum crosnieri

118

ZOOSYSTEMA • 1998 ■ 20(1)

Fig. 9. — A-C, Xeinostoma eucheir Stsbbing, 1920 (Cyclodorippidae). 9, 5 x 5,5 mm (MNHN E 24717) ; D, Phyllotymohnum oros-

nieri Tavares, 1993, 9, 4 x 4,5 mm (MNHN B 24618) : abdomen chez la femelle. B, C, la flèche indique le haut de la figure ; les

pléopodes s'articulent à l'extrémité des segments abdominaux el non pas sur leur face ventrale : D, les exopooites des pléopodes,

assez larges, assurent la fermeture latérale de l'abdomen et la ponte est entièrement enserrée dans la sorte de boîte ainsi formée.

4. PII présents, uniramés et vestigiaux, Exopodite de P12-5 comme d’ordinaire.

Sternite 7 chevauchant une partie du sternite 6. Genkaia (5)

5. Distance entre la saillie latéro-brancbiale de la carapace et de la dent exorbitaire cor¬

respondante aussi grande que la moitié de la largeur fronto-orbitaire de la carapace.

Régions métabrancbiales presque lisses. Doigts des chélipèdes allongés (doigt fixe

orné de deux carènes longitudinales assez nettes sur chacune des faces interne et

externe : doigt mobile avec une seule carène). Bord préhensile du doigt fixe armé de

dents menues. Face ventrale du propode du chélipède avec une avancée bien déve¬

loppée et en forme d’aile ; tace ventrale du mérus munie d’une bosse assez grosse ....

...*.....s. G. keijii

—Distance entre la saillie latéro-branchiale de la carapace et de la dent exorbttaire cor¬

respondante nettement moins grande que la moitié de la largeur fronto-orbitaire de

la carapace. Régions métabrancbiales abondamment garnies de granules aplatis.

Doigts des chélipèdes plutôt courts (paire de carènes sur le doigt fixe et carène

impaire du doigt mobile très peu nettes). Bord préhensile du doigt fixe armé de

dents triangulaires fortes et assez développées. L’avancée aliforme de la face ventrale

ZOOSYSTEMA • 1998 • 20(1)

Tavares M.

Fig. 10. — Phyllotymollnum crosnieri Tavares, 1993, J, 4,2 x 4,2 mm (MNHN), vue dorsale. Échelle : 2 mm.

du propode des chélipcdes moyennement développée ; pas de bosse (ni d autre pro¬

tubérance accentuée, seulement des granules) sur la face ventrale du mérus .

... ..........................G. gordonae

Liste des espèces de Phyllotymolinidae et

de Cyclodorippidae illustrées dans cette

NOTE

Clythrocerus nitidus (A. Milne Edwards, 1880)

(Cyclodorippidae), Bâche, W Stimpson coll.,

Sandy Key , 135 ni : d, 4 X 4,5 mm (MNHN-

B 13489).

Ketamia depressa (Ihle, 1916)

(Cyclodorippidae), Indonésie, Siboga, stn 260,

îles Kei, 5 a 36,5’S - 132°55,2’E, 90 m : 9 para-

lectotype, 4 X 4,3 mm (ZMA-De 102973).

Phyllotymotinum crosnieri Tavares, 1993

Nouvelle-Calédonie, SMIB 6, stn DW 117,

18°59,4ü'S - 163°25,40’E, 280 m : 3 holotype,

5,7 x 6,9 mm (MNHN-B 24617).

Phyllotymolintim crosnieri Tavares, 1993

Nouvelle-Calédonie, SMIB 6, stn DW 117,

18°59,40’S - 163°25,40’E, 290 m : ?, 4 x

4,5 mm (MNHN-B 24618).

Phyllotymolinum crosnieri Tavares, 1993

Nouvelle-Calédonie, Lagon , stn 500, 19°04’S -

163°30’E, 4 ni. 1985, 225 m: 3,4,2 x 4,2 mm

(MNHN).

Xeinostomu euebeir Stebbing, 1920

(Cyclodorippidae), Vauban , 12°38,5’S -

48°16,5’E, 240 m : ?, 5 x 5,5 mm (MNHN-

B 24717).

Remerciements

Nous remercions vivement D. Guinot (Muséum

national d Histoire naturelle, Paris) et Colin

L. McLay (Canterbury University) qui ont relu

ce manuscrit et nous ont lait bénéficier de leurs

critiques constructives.

Une partie de ce travail a été menée au Labo-

120

ZOOSYSTEMA • 1998 • 20(1)

Phyllotymolinidae, fam.nov.

ratoire de Zoologie (Arthropodes) du Muséum

national d'Histoîre naturelle, Paris. Nous

sommes heureux d’exprimer notre gratitude à

son directeur Yves Coineau, qui nous a ouvert les

portes de son Laboratoire et des précieuses col¬

lections qui y sont déposées.

Nos plus vifs remerciements vont à Alain

Crosnier, directeur de recherche à l’Orstom, qui

nous a confié pour étude les espèces indo-ouest-

pacifiques mentionnées dans ce travail et au

Professeur Lipke B. Holthuis pour son aide

concernant les questions de nomenclature. Dirk

Platvoet (Zoologisch Muséum, Amsterdam) nous

a aimablement envoyé des spécimens de

Cyclodorippidae conservés dans son institution.

Nous sommes reconnaissant à Francine Meury

(Centre universitaire de microscopie électro¬

nique, Université Pierre et Marie Curie,

Paris VT), qui a apporté son concours lors des

séances de microscopie à balayage et de prépara¬

tions photomicrographiques.

Le CNPq (Conselho National de Desenvol-

vimento Cientffico e Tecnolôgico, proc.

n° 520254/95-3) et l'Université Santa Ursula,

témoignant du grand intérêt qu'ils ont toujours

porté à nos recherches sur la systématique, ont

financé une partie des recherches présentées ici.

RÉFÉRENCES

Guinot D. 1968. — Recherches préliminaires sur les

groupements naturels cher, les Crustacés Décapodes

Bracnyoures. V. Établissement d’un caractère évo¬

lutif : l'articulation ischio-mérale des chélipèdes.

Bulletin du Muséum national d'Histoire naturelle ,

Paris, série 2, 40 (1) : 149-166. figs 1 -19.

— 1979. — Données nouvelles sur la morphologie, la

phylogenèse et la taxonomie des Crustacés

Décapodes Brachvoures. Mémoires du Muséum

national d'Histoîre naturelle, série A, Zoologie 112 :

1-354, figs 1-70, pis 1-27, tabî. 1-5.

Guinot D. Ôt Richer de Forges B. 1995 — Cmstacea

Decapoda Brachyuia r révision de la famille des

Homolidae de Haan, 1839, in Crosnier A. (ed.),

Résultats des Campagnes MUSORSTOM, volume

13. Mémoires du Muséum national d'Histoîre natu¬

relle 163 : 283-517, figs 1-77.

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cies of the Dromiacea (Crustacea Decapoda) from

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Papers ofthe Zoologieal I.aboratoiy. Faculty of Agri¬

culture, Kvushit Uni vers! ty 3 (3): 19-28. figs I, 2.

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anglais]; 1-461 [en japonais]; 1-16, pis 1-251.

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Brachyura ol the Palau Islands. 1. Drorniidac,

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somatidae, Majidae and Parthenopidae. Bulletin of

the liberal Arts and Sciences- Course , Nihon

Univershy School nf Medtcinc 1: 75-126, figs 1-6,

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around the Isushima Islands Collccted by the

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dorippoidea Ortmann et remarques sur les genres

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(Crustacea, Decapoda, Brachyura). Bulletin du

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12 1990 (1991), (3-4) : 623-648, figs Ml.

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Stimpson, avec la description de Tymolus brucci

n.sp. d’Australie occidentale (Crustacea, Brachyura,

Cydodorippoidea), Bulletin du Muséum national

d'Histoîre naturelle , Paris, série 4. A 13 (3-4) :

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ZOOSYStEMA • 1998 • 20 (1)

121

Tavares M.

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509-514, figs 1,2.

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(Ortmann, 1892) (Crustacea, Decapoda,

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national d'Histoire naturelle, Paris, série 4, A 14

(1) : 201-207, figs 1-3.

— 1993a. — Crustacea Decapoda : les

Cyclodorippidae et Cymonomidae de l'Indo-

Ouest-Pacifique à l’exclusion du genre Cymonomus,

in Crosnier A. (ed.), Résultats des Campagnes

MUSORSTOM, volume 10, Mémoires du Muséum

national d'Histoire naturelle 156 : 253-313,

figs 1-20.

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veaux genres et trois nouvelles espèces de Cyclo-

dorippoidea américains (Crustacea, Decapoda,

Brachyura). Vie et Milieu A3 (2/3) : 137-143, fig. 1.

— 1994. — Xeinostoma inopinata sp. nov., a new crab

from Réunion Island, soutn Indian Océan

(Crustacea: Brachyura: Cyclodorippidae:

Xeinostomatinae). Memoirs of the Muséum of

Victoria, Melbourne 54: 121-123, fig. 1.

— 1996. — Révision systématique des

Cyclodorippidae américains (Crustacea, Decapoda,

Brachyura). Bulletin du Muséum national d’Histoire

naturelle, Paris, série 4, A 18 (1-2) : 233-295,

figs 1-24.

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MUSORSTOM, volume 18, Mémoires du Muséum

national d’Histoire naturelle 176 : 261-271.

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messingi, a new genus and species of Cyclodorippi¬

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Soumis le 19 septembre 1997 ;

accepté le 26 novembre 1997.

122

ZOOSYSTEMA • 1998 • 20(1)

Collemboles interstitiels de sables continentaux

et fluviatiles du Venezuela

Jean-Marc THIBAUD

Laboratoire d’Entomologie, Muséum national d’Histoire naturelle, EP 90 du CNRS,

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

Alba DIAZ

Departamento de Biologia, Facultad de Ciencias, Universidad de Los Andes,

Mérida5101 (Venezuela)

MOTS CLÉS

Collembola,

taxinomie,

nouvelle espèce,

sables continental et fluviatile,

Venezuela.

Thibaud J.-M. & Diaz A. 1998. — Collemboles interstitiels de sables continentaux et fluvia¬

tiles du Venezuela. Zoosystema 20 (1) : 123-127,

RÉSUMÉ

Dans ce travail nous citons huit espèces de Collemboles interstitiels des sables

continentaux et fluviatiles du Venezuela, dont une espèce nouvelle pour la

science : Xenyllogastrura venezueliensis n.sp. (Hypogastruridae).

KEYWORDS

Collembola,

taxinotny,

new species,

continental and fluvial sands,

Venezuela.

ABSTRACT

lnterstitial Collembola ofcontinental and fluvial sands from Venezuela. In this

workwe list eight interstitial species of Collembola from continental and flu¬

vial sands in Venezuela, one of these species being new for science :

Xenyllogastrura venezueliensis n.sp. (Hypogastruridae).

ZOOSYSTEMA • 1998 • 20(1)

123

Thibaud J.-M. & Diaz A.

INTRODUCTION

Les Collemboles ont été récoltés, par le premier

auteur, lors d'une mission effectuée au Venezuela

en décembre 1996. Ils onr été extraits, par la

méthode du lavage, du sablé de dunes continen¬

tales proches de la ville de Coro dans l’État du

Falcon, du sable lluviatile du Rio Chanta près

d’El Vigia et du sable d'un torrent glaciaire du

Piramo de Mttcubajl dans l’Etat de Mérida.

Huit espèces ont été identifiées, dont trois nou¬

velles pour le Venezuela, et une nouvelle pour la

science : X 'enyllagastrUra venezueliensis n.sp.

L’holotype er des paratypes sont déposés au

Laboratoire d’ Entomologie du Muséum national

d’Histoire naturelle à Paris (MNHN), d’autres

paratypes sont déposés au Département de

Biologie de l’université des Andes à Mérida.

Famille HypogasTRURIDÆ Borner, 1913

Xenyllogastrura venezueliensis n.sp.

(Figs 1 -3)

Matériel-type. — Holotype 9 et 9 paratypes 6 et

? (MNHN) et 4 paratypes <3 et 9 (Mérida,

Universidad de los Andes),

LOCALITÉ-TYPE. Venezuela, sable dunaire conti¬

nental à Coro (État du Falcon), 6.XII.1996,

J.-M. Thibaud rcc.

Description

Longueur de l’holotype femelle : 400 pm ; des

paratypes : 380-400 pm. Couleur blanche ;

seules les cornéules résiduelles sont légèrement

pigmentées en gris. Grains tégumentaires moyens

(1 pm). Soies du même type, soies sensorielles un

peu plus épaisses.

Antennes petites (57 pm), 0,8 fois plus courtes

que la diagonale céphalique. Rapports articles

antennaires 1 : II : il I : IV = 1 : 1,2 : 1,36 : 1,6.

Articles antennaire 1 et II avec, respectivement,

sept et douze soies. Articles antennaires III et IV

sans séparation nette. Organe sensoriel de

l’article antennaire 111 classique du genre, avec

deux microsensilles globuleuses, cachées partielle¬

ment par un repli tégumentaire et encadrées par

deux sensilles de garde plus longues er cylin¬

Fig. 1. — Xenyllogastrura venezueliensis n.sp., articles anten¬

naires III et IV.

driques, ne dépassant pas la limite supérieure de

ce repli tégumentaire, et par trois soies de garde ;

présence d’une quinzaine d’autres soies et de la

microsensille ventro-externe. Article antennaire

IV avec sept ou huit sensilles cylindriques ; pré¬

sence de la microsensille, ici assez longue, de

l’organite subapical et d'une grosse vésicule api¬

cale (Fig. 1).

Organe postantennaire avec 4 + 4 tubercules péri¬

phériques , 33,3 % des individus présentent une

asymétrie avec 4 + 3 tubercules. Le diamètre de

cet organe postantennaire est égal à deux ou

trois fois celui d’une cornéule (Fig. 2). 4 + 4 cor¬

néules résiduelles d’un diamètre réduit (2 à

3 pm). Les pièces buccales sont classiques du

genre

Tibiorarses I, Il et 111 avec, respectivement, dix-

sept, dix-sept et seize soies, sans ergot capité ;

fémurs avec chacun dix soic-s et trochanters avec

chacun cinq soies. Griffes trapues, sans dent,

avec un cm podium assez court.

Iube ventral avec 6 + 6 soies. Rétinacle avec

2 + 2 dents. Fürca courte (28 pm). Rapport

manubrium : mucro-dens = 1,3 : l. Dens avec

deux soies dorsales. Présence de deux épines

anales très petites (2 à 3 pm).

Chétotaxie dorsale (Fig. 2). Signalons, sur les

124

ZOOSYSTEMA • 1998 • 20(1)

Collemboles interstitiels du Venezuela

thorax II et III, la présence des m2, la position en

p3 des soies sensorielles, l'absence des microsen-

silles latérales et, sur l’abdomen V, la présence des

soies m2.

Chétotaxic ventrale des abdomens II à IV (Fig. 3).

Signalons, sur la tête, la présence de 2 r 2 soies le

long de la ligne ventrale, sur le thorax l’absence de

soie, sur l'abdomen IV la présence, à la base du

manubrium, d'une paire de macrochères à large

embase, caractéristiques du genre.

Ces chétotaxies présentent parfois certaines varia¬

bilités. Par exemple, sur le sternite abdominal IV,

la rangée postérieure porte 1 + 1 ou 3 + 3 soies.

Discussion

Cette nouvelle espèce est très proche de

Xenyllogastrura oaocvlata (Steiner, 1955)

d’Europe et de Méditerranée, espèce redécrite par

Deharvcng & Gers (1979), et proche de X. are-

naria Fjellberg, 1991 des îles Canaries.

Elle se différencie de la première par la longueur

des deux sensilles de garde sur l’article anten-

naire III, ainsi que par celle de la microsensille

sur l’article antennaire IV et par quelques détails

de sa chétotaxie.

Elle se sépare de la seconde par la chétotaxie des

tibiotarses, par la Variabilité oculaire, par la lon¬

gueur de l empodium, par la taille du mucro-

dens, par la longueur des deux sensilles de garde

sur l'article antennaire III, et par quelques détails

de sa chétotaxie.

Ajoutons enfin que cette citation est la première

du genre Xenyllogastrura , genre méditerranéen-

européen-macaronésien, dans la région néotropi¬

cale.

Famille BraCHYSTOMELUDAE Stach, 1949

Rapoportbella yolandae

(Rapoport et Mano, 1969)

Matériel examiné. — 1 2, sable fluviatile du Rio

Chama à El Vigia, 2.XJI.1996.

Remarque

Espèce édaphique-psammophile, néotropicale,

décrite d’un sol de savane du Venezuela et retrou¬

vée dans plusieurs îles des Petites Antilles.

t P t S\ -•

i! .

'! ' -, i \ t '

-'1 ( i s “ ■

, I » * > î

-I V l ^ i ! 1

“ , /

I ‘'-‘'■'I'

î k» ?

'N 'I i

-H-

; ' 7 i

us /»

vv ; > -7

l. * /

, ...

i r \

A i |

r 1 ^ /

/ v

j \ i

n A4

r <• G A

.LL.L.I.j

j i * * \

g i

f i / f 'S

^ 1 cv f"

■s

Fig. 2. — Xenyllogastrura venezuellensis n.sp., chétotaxie dor¬

sale.

Famille ONYCHIURIDAE Borner, 1913

Mesaphorura macrochaeta Rusek, 1976

Matériel EXAMINÉ. — 6 9 et 14 ex. en alcool, sable

d’un torrent glaciaire dans la vallée de Mucabajf

(Pâramo de Mucubajf, État de Mérida), 3.XH.1996.

Remarque

Espèce édaphique-psammophile, holarctique,

retrouvée ici dans un milieu montagnard.

Protapborura encarpata (Denis, 1931)

Matériel examiné. — 12, sable d’un torrent gla-

ZOOSYSTEMA • 1998 • 20(1)

125

Thibaud J.-M. & Diaz A.

Fig. 3. — Xenyllogastrura venezueliensis n.sp., chétotaxie ven¬

trale des abdomens II à IV.

ciaire dans la vallée de Mucabajl (Pâramo de

Mucubajt État de Mérida), 3.XII.1996.

Remarque

Espèce édaphique, néotropicale, déjà signalée par

Diaz & Najt (1995) du même pâramo.

Famille ISOTOMIDAE Borner, 1913

Folsomides purvulus Stach, 1922

Matériel EXAMINÉ. — 1 9 et 1 juv., sable fluviatile

du Rio Chaîna à El Vigia, 2.XIL1996. 1 9, sable

d’un torrent glaciaire dans la vallée de Mucabajl

(Pâramo de Mucubajt, État de Mérida), 3.X1I.1996.

Remarque

Espèce édaphique-psammophile, cosmopolite,

déjà signalée au Venezuela.

Folsomina onychiurina Denis, 1931

Matériel EXAMINÉ. — 8 9, sable fluviatile du Rio

Chama à El Vigia, 2.X1I.1996.

Remarque

Espèce édaphique-psammophile, cosmopolite,

déjà signalée au Venezuela.

Cryptopygus thermophilus (Axelson, 1900)

Matériel examiné. — 1 9 et 1 â , sable fluviatile du

Rio Chama à El Vigia, 2.X11.1996.

Remarque

Espèce édaphique-psammophile, cosmopolite,

déjà signalée au Venezuela.

Isotomieüa et. symetrimucronata

Najt et Thibaud, 1987

MATÉRIEL EXAMINÉ. — 1 9, sable d’un torrent gla¬

ciaire dans la vallée de Mucabajl (Paramo de

Mucubajt, État de Mérida), 3.X11.1996.

Remarque

Espèce édaphique, pantropicale, décrite de

l'Équateur, déjà retrouvée en Amazonie brési¬

lienne, dans les îles Seychelles, dans l’atoll de

Fangataufa (Polynésie) et dans trois îles des

Galapagos (Thibaud, Najt & Jaquemart 1994).

Cet unique exemplaire est, malheureusement,

dépourvu de dens et de mucron.

Remerciements

Ce travail a pu être réalisé grâce à une conven¬

tion internationale C.N.R.S./C.O.N.I.C.I.T.

(Venezuela).

RÉFÉRENCES

Deharvcng L. & Gers G. 1979. — Sur les genres

Xenyllogastrura Denis, 1932 et Orogastrura n.g.

Travaux du Laboratoire d’Écobiologie des

Arthropodes édaphiques, Toulouse I : 1-10.

Diaz A. & Najt J. 1995, — Cnllemholes (Insecra) des

Andes vénézuéliennes. Bulletin du Muséum national

d'Histoire naturelle. Paris, série 4, A 1994 (1995),

16(2-4) : 417-435.

Fjellberg F. 1992. — Collembola ol the Canary

Islands. I. Introduction and survey of the family

Hypogastruridae. Entomologia scandinavica 22:

437-456.

126

ZOOSYSTEMA ■ 1998 • 20(1)

Collemboles interstitiels du Venezuela

Rapoport E. H. & Manos S. 1969. — Colembolos de

Venezuela. I. Acta Biologia Venezuolana 6:

117-128.

Thibaud J.-M., Najt J. & Jaquemart S. 1994. — Les

Collemboles (Insecta) de l’Archipel des

Galapagos. II. Isotomidae. Bulletin de l’Institut

royal des Sciences naturelles de Belgique ,

Entomologie 64 : 199-204.

Soumis Le 7 mars 1997;

accepté le 2 septembre 1997-

ZOOSYSTEMA • 1998 • 20(1)

127

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Références bibliographiques

Hoeg J. T. &C Liitzen J. 1985. — Comparative

morphology and phylogeny of the family

Thotnpsoniidae (Cirripcdia: Rhizocephala:

Akemrogonida) with description of ihree nevv

généra and seven new species. Zoologica Scrîpta

22: 363-386.

Rôckel D. t Xorn W. &c Kohn À. J. 1995. —

Mannal of the living Conidae , volume 1: Indo-

Pacific région. Christa Llemmen, 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. & Lhmann H. (eds), Hiology of

Australasian Progs and Reptiles. Surrey Beatty and

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Manuscripts, without limitation of the nurnber of

pages, must conform strictly with the instructions

to authors and will be sent to the Editor:

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130

ZOOSYSÎEMA • 1998 • 20(1)

Instructions aux auteurs

sented as follows: Smith (2001), Smith (2001,

2002), (Smith 2001), (Smith 2001; Cary 2002),

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- references to illustrations and tables should be

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References

Ffoeg J. T. & Lützen J. 1985. — Comparative

morphology and phylogeny of the family

Thompsoniidae (Cirripedia: Rhizocephala:

Akentrogonîda) 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-

Pacific région. Christa Hemmen, Wiesbaden,

517 p.

Schwaner T. D. 1985. •— Population structure of

black tîger snakes, Notechis ater niger, on offsho¬

re islands of South Australia: 35-46, in Grigg G.,

Shine R. &C Ehmann H. (eds), Biology of

Australasian Frogs and Reptiles. Surrey Beatty and

Sons, Sydney.

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ZOOSYSTEMA • 1998 • 20(1)

131

Mise en page

Noémie de la Selle

Packaging Éditorial

Achevé d’imprimer

sur les presses de l’Imprimerie Durand

28600 Luisant (France)

Mars 1998

Dépôt légal n° 9975

Printed on acid-free paper

Imprimé sur papier non acide

Date de distribution le 30 mars 1998

Couverture: Branchinecta minuta Smirnov, 1948 (Crustacea, Anostraca),

détail de l’enveloppe de l’œuf (MEB).

Photographié A. Thiéry (Université d’Avignon).

zoosystema

1998 • 20 (1)

Vente en France

Muséum national d'Histoire naturelle

Diffusion Delphine Henry

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Conception Graphique : Isabel Gautray

ISSN : 1280-9551

101

109

123

Vacelet j. 6t Perez T.

• Two new généra and species of sponges (Porifera, Demospongiae) without skeleton from a

Mediterranean cave ^ jk

m w

Diouf M., Bâ C. T. & Durette-Desset M.-C.

• Deux nouveaux Neoheligmoneila (Nematoda, Trichostrongylina, Nippostrongylinae) parasites de

Mastomys erythroleucus (Muridae) au Sénégal jjflHkc. sjjpp

Young P. S.

• Cirripedia (Crustacea) from the "Campagne Biaçores" in the Azores région, including a generic révi¬

sion of Verrucidae ■BaMjBjay an—

Dalens H.

• Isopodes terrestres de Nouvelle-Calédonie II. Sur une nouvelle espèce du genre Pseudolaureola

(Crustacea, Oniscidea)

Lemaitre R.

• A new species of hermit crab of the family Parapaguridae (Decapoda, Anomura) from French

Polynesia JB Æ

Tavares M.

• Phyllotymolinidae, nouvelle famille de Brachyoures Podotremata (Crustacea, Decapoda)

Thibaud J.-M. & Diaz A.

• Collemboles interstitiels de sables continentaux et fluviatiles du Venezuela