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Micromorphology and microstructure

as expressions of scleractinian skeletogenesis

in Favia fragum (Esper, 1795) (Faviidae,

Scleractinia)

Jean-Pierre CUIF

Département des Sciences de la Terre, Université Paris-Sud,

Bâtiment 504, F-91405 Orsay cedex (France)

cuif@geophy.geol.u-psud.fr

Christine PERRIN

Laboratoire de Paléontologie, Muséum national d’Histoire naturelle,

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

perrin@mnhn.fr

Cuit J.-P. & Perrin C. 1999. — Micromorphology and microstructure as expressions of scle¬

ractinian skeletogenesis in Favia fragum (Esper, 1795) (Faviidae, Scleractinia).

Zoosystema 21 (2) : 137-156

ABSTRACT

Microstructural différences observable within septa are currently recognized

as major phylogencric critcria among Scleractinia. If the emergence of niole-

cular techniques provides a good mean ofassessing thèse criteria, sonie recent

results using molfccuîar approach seem to give only low support for niicro-

structural-based phylogcuetic rclationships of Scleractinia. This, however,

may resuit from the numerous uncertainties concerning rhe microstructural

pattern of species upon which major taxa arc based. Three-diniensional cha-

racters of septal growth units and rheir varïabiliry hâve here been investigared

using the specimens of Favia fragum included within the Milne Edwards

Collection, these being the oldest référencé ntaterial still available today.

Integrating these data with results front the most recenr work on this species

KEYWORDS leads to the reconstruction of the three-dimensional pattern of septal archi-

^Sderacnnia' tecture. The phylogenetic significance of this requires assessment with alter-

skeletogenesis. native approaches.

ZOOSYSTEMA • 1999 • 21 (2)

137

CuifJ.-P. & Perrin C.

MOTS CLÉS

Favia fragum ,

Scleractinia,

squelettogenèse.

RÉSUMÉ

Micromorphologie et microstnuiure chez Favia fragum (Esper, 1795) (Faviidae ,

Scleractinia) en uni qu 'expressions de h squeletrngenese des scléractiniaires.

Chez les scléractiniaires, ce sont actuellement les différences microstructu¬

rales observables dans les septes qui sont reconnues comme critères phylogé¬

nétiques majeurs. Si l'émergence des méthodes moléculaires fournit des

moyens d’évaluer ces critères, des résultats récents utilisant l'approche molé¬

culaire semblent donner seulement une faible valeur à la phylogénie des sclé¬

ractiniaires fondée sur les microsrructures. Ceci, toutefois, peut résulter des

nombreuses incertitudes concernant les microstruclures des espèces sur les¬

quelles les taxons majeurs snnr basés. Les caractéristiques rridimenrionnelles

des unités de croissance septales et leur variabilité ont été ici étudiées sur les

spécimens de Favia fragum de la collection Milne Ldwards, qui sont le plus

ancien materiel de référence actuellement disponible. La prise en compte de

l’ensemble des données résultant des recherches plus récentes concernant

cette espèce permet la reconstitution tridimentionnelle de l'architecture sep¬

tale. La signification phylogénétique de ce type d’analyse devra etre évaluée

par rapport aux résultats obtenus par les approches concurrentes.

INTRODUCTION

As pointed out by Veron et al. (1996) the well

known Wells’ evolutionary tree for Scleractinia

“has long been in need of révision”. This is well

shown by important diserepancies existing bet-

ween the highest taxonomie subdivisions of the

alternatively proposée! evolutionary schemes ot

Scleractinia, even when considering only the

most receni hypothèses of Veron himself (1995),

Roniewicz &: Morycuwa (1993) and Chevalier &

Beauvais (1987).

Pioneered by Pratz (1882), Ogilvie (1896) and

nutnerous others, description of fibre arrange¬

ments in corallian structures soon became reco-

gnized as a prominent rool in cotai taxonomy

and later, was formally regarded of fundamental

importance by Wells (1956: fig. F237, F340). By

contrast, rccent DNA sequencing studies led

Veron et al. to the conclusion thaf "morphologi-

cal characters of extanr corals dLstinguish fanulies

more rcliahlv than do internai microskeletal cha¬

racters frequently used in coral paleonrology”

(1996: 1). Il truc, this conclusion should Icad to

a basic mcthodological change in the taxonomie

approach to the scleractinian skeleton.

Numetous obscure points still persist in the

microstructural knowledge of the “key species”,

i.e. the type species lor the généra on which

lamilies are based. Therelore, microstructural

study of rhese "key species” is a prerequisite for

crirical reexamination ol related supra-specifîc

taxa.

Among these "key species”, Madrepora fragum

Esper, 1795 the type species of the genus Favia,

is one ol the most important, as il is the type

genus for one of the major sclcractinian taxa: the

family Faviidae, and the suborder Faviina

(Triassic to Recent). In addition, this species also

illustrâtes the pour State ol basic microstructural

data. In most cases, précisé descriptions of septal

growth including recent improvement in under-

standing ol biomineralization processes are com-

pletely lacking. This cannot constitute a reliable

consistent basis for taxonomie considérations.

The aim of this paper is theiefore to présent an

accurare description of the type species

Madrepora fragum in the ligln ot recent know¬

ledge ol sderactinian skelerogenesis. Fmphasis is

espeeially placed on integraung morphological,

micromorphological and microstructural data in

order to provide a better understanding ol the

thrcc-dimcnsional skclcial growth ptoccss. These

analyses hâve been carried out on the Favia fra-

138

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia fragum

gum colonies of rhe Milne Edwards Collection

houscd at thc Muséum national tf Histoire natu¬

relle of Paris. Because the type specimen of Favia

fragum is now lost, the marerial of Milne

Edwards represents today die most relevant spé¬

cimens from an historical point of view.

SUMMARY OF PREVIOUS STUDIES

Nomenclature

Originally created by Olten (1815), rhe genus

Favia was inciuded in the rejection by the ICZN

of ail taxa proposée! in the Oken's book (decision

417). Howcvcr, thc genus group naine Favia was

made availahlc- by thc work of Milne Edwards

(1857) vvho Uscd it for a group of species inclu-

ding Madrepora fragum Esper, 1795. Verrîll

(1902) designated Madrepora fragum as type spe¬

cies' of the genus Favia. From a forma! point of

view, the genus ninte Favia shotlld bc consideted

as Favia Milne-Edwards, 1857 [not Oken,

1815).

Concerning the choice of Madrepora fragum

Esper, 1795 as type spccies ot Favia by Verrill

(1902), Alloiteau (1957) and later Zlatarski &

Martine/. Estalella (1982) followcd Verrills pro¬

posai although Madrepora fragum Esper, 1795 is

considered as a posterior synonym of Madrepora

ananas Pallas, 1766. Verrill (1902: 90) emphasi-

zed that thc use pf ananas could introduce confu¬

sion sincc the name Madrepora ananas was

already used by Linnacus (1758) for a Palaeozoic

coral froni Gotland now belonging to the genus

Acervularia Schweigger, 1819 and also applied to

extant spccies by earlier authors (not mentioned

in the Verrills paper).

Since Milne Edwards (1857), the genus Favia

gained in taxonomie importance as it was desi¬

gnated as type genus of the Faviidae by Gregory,

1900 and thc family Faviidae considered as rhe

basis for the suborder Faviina Vaughan & Wells,

1943.

In the first publication of Favia fragum (Esper,

1795), Esper although not using conventional

terms for describing coral specimens, referred to

the geometry of the calices, the wall structure

and even the distal margin of the septa with sur-

prising précision. This type specimen which was

sampled from Récent reefs of rhe Caribbean, rs

now lost (Zlatarski & Martinez Estalella 1982).

Milne Edwards tk Hainte (1849), describing a

Recent specimen from Haiti, provided more

detailed information about the geometry and

structure ot thc calices. These especially concern

intracalicinal budding, septal arrangement in

four cycles and the distal margin of major septa

showing a W'ell-dcvelopcd tooth at the axial mar¬

gin. Fhe specimen was referred to Pamstrea fra¬

gum by the authors (Milne Edwards tk I lamie

1849) but later transterred to the genus Favia by

Milne Edwards (1857). Subséquent murpholo-

gical descriptions ot this spccies are numerous,

inclutling observations on the living tissues (c.g.

Duerden 1902; Matthai 1919).

THE FIRST MICROSTRUCTURAL ANALYSIS OF F AV/A

FRAGUM BY Al I OITEAU (1957)

Alloiteau was the first to analyse the microstruc¬

ture of Favia fragum from thin sections.

According to this author, septal and costal micro-

structures slightly differ. The costal part is for-

med by very large and simple rrabecnlae

(Alloiteau 1957: 138, “poutrelles"), clearly sepa-

rat-ed from each other by a thin demiculate

suturai line. Each trabecula seen in transverse

thin section contains a large calcification centre

consist-ing of several dark dots from which ara-

gonitic fibres radiatc. The septal part shows

snialler trabeculac with less well-dehned borders

and the calcification centres are someiimes so

close that they furm a thick subcontinuous

médian dark line. Additioiul calcification centres

also occur on both sides of this médian line.

Aragonitic libres radiate from each centre and are

perpendicular to the latéral faces of septa.

The study of Favia fragum by Zlatarski tk

Mari inkx Estait! la ( 1982)

Based on detailed observations of an extensive

collection from Cuba, the work of Zlatarski &

Martine/, Estalella (1982) provides one of the

most compréhensive studies of Favia fragum,

including description of the successive stages of

the budding process and assessment of intra-

specific variability. In particular, calicinal arran¬

gement, usually regarded as a generic character of

Favia [plocoid, Wells (1956); subplocoid,

ZOOSYSTEMA • 1999 • 21 12}

139

CuifJ.-P. & Perrin C.

Alloiteau (1957)]. appears to be highly variable.

The calicinal surface may be plocoid, cerioid or

meandroid and oftêli shows interniediace stages

between these various arrangements. Other

variable characters observed by rhese authors,

include calicinal si/.e, ornamentation of perithe-

ca, thickness ofsepta, abondance of septal denta-

tion and colunrella development. Observations

on the skeletal microstructore are in good accor¬

dance with the previous description of Alloiteau

(1957). While in the costal part, the trabeculac

within thin sections are well difïerentiated with

irregular quadrangular periphery and large calci-

fication centres, they gradually incline towards

the septal axial margin torming a large dark

médian line in the central part ol septa.

APPROACHRS AND TECHNIQUES

Specimens of Favia fragum were successively

observed on different scales and treated in several

ways.

Morphological data were obtained from observa¬

tions of each colony with a stereomicroscope.

Detailed qualitative observations were related to

measuremems of geometrical characters usually

regarded as standard criteria for scleractinian

taxonomy.

Micromorphology and its variability were stu-

died by scanning-electron microscopy (SEM).

According to the morphological data, représenta¬

tive calices were selected within each colony and

sampled ttsing a dental saw. These samples were

then coated with gold-palladium for 3D-observa-

tions with SEM.

Microstructure was studied in optical micro¬

scopy from the thin sections of the Alloiteau

Collection housed at the Laboratoire de

Paléontologie, MNHN, Paris.

Ultrastructural data were obtained by 2D-obser-

vation of transverse sections with SEM. Selected

calices within each colony were cross-sectioncd

and polished with diamond paste. Several

etching préparations ot these surfaces were tried,

in parricular for précisé observation of calcifica¬

tion centres and their internai strucrure. Some

polished sections were treated with an enzymanc

solution of alcalase, pH 7.5 for three hours at

35 °C with stirring ol ihc préparation. A second

set of samples was also prepared by etching with

a solution of formic acid 0.1% and glutaralde-

hyde 3% for 40 seconds. Ecched préparations

were then coated with gold-palladium loi SEM

observation of ukrastructurc, The latter was also

compared with the microstructural data.

Ail SEM préparations are included in rhe MiLne

Edwards Collection housed at the Laboratoire de

Biologie des Invertébrés marins et Malacologie,

Muséum national d’Histoire naturelle, Paris.

DESCRIPTION OF FAVIA FRAGUM FROM

THE Mil .NE EDWARDS COLLECTION AT

THE MNHN

The Mu ni Edwards Collection

AT TH K MNHN

This collection housed at the Laboratoire de

Biologie des Invertébrés Marins et Malacologie

(BIMM), Muséum national d’Histoire naturelle,

Paris, comprises five specimens or groups of spe¬

cimens sampled from diverse localities of the

Caribbean. These are regisrered as follows:

FAV559, Favia fragum (Esper), Antilles (bought

at Marseille), one small colony (Fig. IA, B);

FAV560, Favia fragum (Esper), Haïti

(Mr. RiconJ, 1831 ), three small colonies and rwo

fragments (Fig. 1C-H, K, L);

61 fi, Favia fiagum (Esper), Antilles (bought at

Marseille, Mr. I.. Rousseau, 1841), one small

colony (Fig. 1M-O);

631, Favia fragum (F.sper), Antilles (Mr. Roissy,

1837), one small colony (Fig. 11, J);

1260, Favia fragum (Esper), Saint-Thomas

(Mr. Duchassaing. 1870), one colony (Fig. 2A, B).

ExternaL .shape of colony

The Milne Edwards specimens are small low

colonies (about 4-10 cm in diameter and a few

ccntimeters high) with globular. commonly

hemispherical shape. The ovoid external mor-

phology of rwo colonies (FAV560-1. 2) appears

ro be relared to the elongated shape of rheir sub¬

strates, both consisting of a mytîlid shcll. The

smallest specimens (FAV560-3, 4, 5) are low

encrusting growth forms.

140

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia jragum

Fig. 1. — Favia fragum (Esper. 1795), specimens Irom the Mllne Edwards Collection at the MNHN; A, B, specimen FAV559; C. spé¬

cimen FAV560-1: D, E. specimen FAV560-2; F, specimen FAV560-1; G, H, specimen FAV560-3: I. J, specimen 631; K, L, specimen

FAV560-4. MO, specimen 616. Scale bar: 1 cm.

ZOOSYSTEMA • 1999 • 21 (2)

tr k ‘ V - '

9 HP

'Jjf

r mm

Fig. 2. — Favia fragum (Esper, 1795), specimens from the Mllne Edwards Collection al the MNHN. calicinal relationships: A D. spe

cimen 1260: C, subplocoid part of coiony; D, subcerioid part of the colony; EF, specimen 616, E, plocoid calices; F, part of the colo

ny showing elongated meandroid calices. Scale bar: 1 cm.

ZOOSYSTEMA • 1999

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Skclctogenesis in Favia fragum

Fig. 3. — Favia fragum (Esper, 1795), budding, specimen 631; A, distomodeal budding; B. tristomodeal budding within the same

colony. Scale bar: 1 cm.

CAUCINAl RH.AI lONSHII’S

Colonies .ire usually plocoid (Fig. 2E), vvalls oF

individual calices bcing separated by dissepi-

mental coenosteum, Distances between w.i11s of

neighbouring calices are however highly variable

leading co a subplocoid or even a subcerioid

(1260, Fig. 2C, D) appearance of colonies. Short

sc-ries of calices are unconimon (FAV560-I, 2)

although one specimen comprises some verv

elongated calices which tend to form short

meanders at the pcriphery of the colony (616,

Fig. 2F). Intracolonial variability is high, even

within the small colonies of the Milne Edwards

Collection and calicinal relationship seems to be

partly dépendent on the position of calices

within the colony.

Budding

Colony growth occurs through întracalicina!

usually distomodeal budding which results front

subdivision of the mother calice into two

approximately equal parts (Fig. 3A) or sometimes

corresponding respcctively to one third and two

thirds of the initial calice. Division occurs

through the axial development of two opposed

major septa (commonly SI or sometimes S2),

axial margms of which gradually fuse unnl they

form a distinct and compact séparation. On each

side of tliis new wall, ncw septa arc then devclop-

ed resuhing (rom twisting and growth of distal

teeth of the initial septa. fn some cases, both the

opposite septa are rwisted before becoming fused

(631). Tristomodeal budding is rare (Fig. 3B)

and results from the same process of subdivision

into three equal parts. Polystomodeal budding

may be observed on specimen 616 (Fig. 2F), In

this case, centres are differentiated before the

séparation of voung calices and sériés of up to

five to six centres are formed. F h esc parts of the

colony tend to become meandroid.

Camus

The most cominon calices are shallow with

roun ded or elongated shapes often distorted by

intracalicinal budding. Shapes of calices and dis¬

tances between centres are, however, highly

variable between colonies but also within the

same colony. Calicinal shape varies from angular

to suhangular (FAV560), subpolygonal (631). up

to very elongated meandroid calices (616).

Occurrence of irregular calices usually incrcases

towards the periphery of colonies. Diameter of

adult monostomodeal calices averages 5-6 mm

ranging from 4.5 to 7,5 mm in most colonies

wllile their depth is around 1.5-2.5 mm.

However, potential variability of calicinal size

ZOOSYSTEMA • 1999 • 21 (2)

143

Cuif J.-P. & Perrin C.

and depth appcars co bc relativcly high. Size of

clongatcd meandering pôlystomodeal calices

occurring within specimen 616 is about lü-15 X

2-3 mm. In addition, colony 631 shows much

larger calices, diamctei ofwhich ranges from 10-

14 X 7-9 mm (average 11.5 X 8 mm) with calici-

nal depth varying between 5 and 8 mm.

Intracai.icinai. structures

Septa

Cycles, numbtr of septa. Number of septa

ranges front 32 to 54 within adult monostomo-

deal calices (Fig. 4A) and is about 40-41 on ave¬

rage. Septa are organized in four cycles with

sometimes a few additional abortive septa contri-

buting ro the formation of thc septothecal wtdl,

The number of septal cycles appears ver) - consis¬

tent within ail studied colonies. Septa ofthe first

two cycles (SI and S2), et|ual orsubequal, reach

the columella. Septa ofthe third cycle are usually

thinner and their development slightly varies bet¬

ween colonies. AU or only some S3 may teach

the calicinal centre: the other third-cyclc septa

extend to half to three quarters of the calicinal

radius and may sometimes fuse with a neigh-

bouring S2 at their internai border. The fourth

cycle is incomplète, most of its septa being wcak-

ly developed and never exceeding onc quarter of

the calicinal radius. Septa correspond to costo-

septa and are usually straigln or slightly curved.

They are often gradually thinning from the calice

peripheiy to the centre. Septa are compact

althougb somc porcs may occur at their axial

margin or even sometimes in their médian part,

those of the younger cycles being in general more

porous than septa of the two First cycles. The

overall septal porosity varies between colonies

but seems to bc similar within thc same colony.

Process of pore formation is clearly inferred at

the axial margin in some calices. Pores are form-

ed from the horizontal development of the

uppermost internai tooth joining and further

fusing widt the vcrticallv-extended axial tooth of

the same septum, this Icaving an empty rounded

gap below the junction of both teetli.

Latéral faces of septa show conical-to-sharp

points forming more or less regular Unes parallel

to the septal growth direction. Vertical near the

calicinal wall, these lines are gradually inclined

towards the axial septal margin. Close to the dis¬

tal margin, each line ol points corresponds to the

position of a distal tooth (Fig. 4B. C). Points on

the latéral septal faces therefore resuit from the

graduai burial of previous successive teetli within

the incrémental layers o( tibrous aragonite

contiibuting to the septal rhickening.

Septal dentation. Teeth of the distal margin

consist of groups of snrall points with a conical

or curved, sharp ot smooth shape locatcd at the

extremity of the tooth axis (Fig. 4D-F). The

height of teetli regularly incrcases from thc wall

to thc axial margin, the higher teeth devcloping

at the uppermost internai and axial septal margin

(Fig. 4B, C). lncrcasc of tooth height oecurs

through thc élongation of thc dental main axis

and not by thc increasing size ol lIic points.

Teeth close to the calicinal periphery are sub-

parallel to the wall while their inclination is gra-

dtially increasing towards the internai margin

(Fig. 40. In some calices, those of thc upper¬

most internai margin are nearly horizontal. A

strong axial tooth may also lie developed vertical-

ly on those septa reaching the columella (i.e.

mainly SI and S2), these whole axial teeth form¬

ing within somc calices a paliform crown around

the columella. In .sortie rare cases (631), rhe axial

tooth may also be formed ol its ovvn small fan-

system (Fig. 5F). Septa of the First two cycles

bear stronger and better-developed dentation

than those of thc later cycles. 1 fêtai lcd observa¬

tions of dental extremities reveal highly variable

shape and organisation within a given grottp of

points (Fig- 5A-E), The ends of axial teeth and

especially palilorm teeth show a higher number

of points with no preferential orientation while

organisation of the dental structure is gradually

increasing towards the peripheral calicinal tnar-

gin where points are progressive!}' arranged in

small paddle-like structures or fans ortenred nor¬

mal to rhe septal plan. The degree of organisa¬

tion is usually higher at the distal external

margin of septa SI and ,S2 but lurgely varies bet¬

ween calices and colonies. Some specimens never

clearly show the paddle structure, even at the dis¬

tal margin of major septa. By contrasr, paddles

are particularly commun within specimen 631

(Fig. 6).

Microstructure. SEM observations of transverse

144

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia fragum

Fig. 4. — Favia fragum (Esper, 1795), SEM, distal margins of septa; A. specimen 1260. calice showing septal arrangement, columel-

la and dentation ot septal margins; B, C, specimen 631, B, Internai part of a calice, shape and arrangement of distal teeth and for¬

mation of columella; C. part of a calice showing the gradational change of the shape and orientation of teeth at the distal margins of

septa; D-F. specimen FAV560-1 : D, middle part of a distal septal margin; E, more peripheral part of the distal margin; F, detail of D.

Scale bars: A, C, 1 mm; B. 2 mm; D-F, 100 pm.

ZOOSYSTEMA • 1999 • 21 (2)

145

CuifJ.-P. & Perrin C.

Fig. 5. — Favia fragum (Esper, 1795), SEM, variability of axial teeth; A, specimen FAV560-2; B, detail ot A: C specimen 631;

D, specimen FAV560-4; E, specimen FAV560-5; F, specimen 631. Scale bars: A, D, 200 pm; B, 100 pm; C, E, 500 pm; D, 200 pm;

F, 1 mm.

146

ZOOSYSTEMA • 1999 • 21 (2)

Skelctogenesis in Favia fragum

Fig. 6 . — Favia Iragum (Esper, 1795), SEM relatlonship between ultrastructure and micromorphology. specimen 631: A. B. paddle-

like tooth at the dislal margin of septum A, ornamentation of latéral face of septum is the resuit of gradational burial of latéral ends of

teeth by successive incrémental layers ot aragonite: C. D, sets of aragoniflc fibres seen between two neighbouring teeth at the distal

margin of septum Scale bars. A, 200 pm: B-D, 100 ym,

sections of septa takcn ai sttcccssively greater the sanie set hâve the sarne orientation (Fig. 8).

depth within the calice allow the internai struc- Each layer records the successive incrémental

titre and, ihcretore, the successive stages of septal growth of cotai skelcron. Fibre orientation

growth to bc understood (Fig. 7). Calcification through successive layers appears ro be broadly

centres appear as smooth round dots, about consistent although growth direction may slight-

25 pnt in diametet and devold of fibres, but ly vary between successive layers. Sets of fibres

higher magnification shows that they consist of are also clçarly distinguished on well-preserved

very small equant crystals. They are sutrounded distal surface of septa and obviously show a fre-

by concentric layers consisting o( sets of fibres, qtient oblique direction of fibres witlt respect to

Fibres are arranged approximately perpendicular the growth surface (Fig. CiC, D).

to the layer boundaries. Orientation of fibres Distribution of calcification centres within the

within the saine layer slightly varies between septa clearly Controls the external septal micro¬

adjacent sets but fibres of.successive layers within morphology and especially the formation of

ZOOSYSTEMA • 1999 • 21 (2) \/f] I

Cuifj.-P. & Perrin C.

Fig. 7. — Favia fragum (Espef, 1795), SEM, ullfastruclure of the

distal margln, specimen 631 ; A. etched section of a tooth on the

peripheral distal margin showing the line ol calcification centres

from which aragonitic fibres radiale; direction of calicinal margin

is to the bottant of photograph; B, etched section of internat dis¬

tal margin ol a septum; difterenttal development ot aragonitic

fibres is especially well shgwn withirt lhe younger incrémental

layers; direction of calicinal axis to the top of photograph. Scale

bars: 200 pm

rceth. Transversc sections through teeth show

several calcification centres alignée) roughly nor¬

mal to the septal plan. Formation ot teeth occurs

through the development ol aragonitic layers

around thèse centres. Differenrial growth of ara¬

gonite fibres from each centre gradually fills up

the space between rwo ncighbouring teeth and

junction of fibres showing opposite growth from

successive teeth is usttally well marked by an irre-

gular line observed on SEM préparations and in

thin sections (Fig. 8).

Columella

Columella is spongy and results from latéral

development, twisting, Ittsion and tangling of

axial margins and often axial teeth (Fig. 9A-C).

Upper surface of columella may produce small

spines similar to rhe denral points (Fig, 9C).

When axial teeth arc vertically oriented, they do

not contribute to columella structure but tend to

form a paliform crown. Diameter of columella

varies between 1.3 to 2.5 mm, size and the densi-

ry of structure both depending on the develop¬

ment of axial septal margins.

Endotheca

Endotheca consists of frequent thin interseptal

dissepiments wich a slighr upward convexity.

They are developed from latéral faces of neigh-

bouring septa and do not occur ar the sanie level

across different interseptal spaccs. Central junc-

tion sensu Sorauf (1970) wtthin the interseptal

space is marked by a Utile downward fold.

Gcntly inclirted close to tbc calicinal peripliery,

dissepiments rend to becomc subhorizontal to

the centre.

Wau.

Wall is continuons and mainly septothecal, form-

ed by the thickening of septa in their external

part but also by the adjunction of rudimentary

radial éléments developed from new calcification

centres (Fig. 9D-F).

Etched transverse sections of wal! under SEM

show successive stages of wall formation.

Development of wall begins by latéral growth of

bars from opposing latéral faces of rwo neigh-

bouring septa. This occurs through the forma¬

tion of calcification centres at both latéral septal

laces and incrémental growth ot fibres around

each centre. Thc.se centres are roughly alignée!

normal to both septal planes and new centres are

gradually added at each extremity of both

convergent bars leading to their fusion

(Fig. 1 OA). Thickening of this new wall results

from the development of aragonite clusters

around each centre but also, witliin the central

part, from the adjunction of new calcification

centres arranged perpendictilar ro the wall

(Fig. 10B, C) and finally forming a rudimentary

radial element.

Ext 11 IIUCA

Costae and costal dentation

Costae arc usually well-developed, thicker than

septa and non-confluent or subconfluent in rnost

colonies, although specimen 631 shows frequent

cases ol costal confluence. Non-confluent costae

of neighbouriug calices are often separated by a

narrow groove, previouly mentioned by Zlatarski

& Martinez Estalella (1982) but also by Esper

(1795). When costae are snbconflticnt, this groove

is m<ire irrcgular and disconrinuous. Costae cor-

responding to SI, S2 and S3 are longer than

148

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia fragum

Fig. 8. — Favia fragum (Esper, 1795), ultrastructure of a septum; A, SEM, photomosaic of specimen 631 ; B, sketch traced from A;

c.c., calcification centres: i.L lîmit of incrémental layer: j.f.. suture of fibres from neighbouring growth units; s.f.. sets of aragonitic

fibres. Scale bars: 200 pm.

those of the last cycle which are rhick and short.

Costal dentatîon (Fig. 11) comprises simple

conical thick teeth close to rhe wall (Fig. 11 B),

while furthcr from the calicinal margin, they are

gradually inclined ourwards and may develop a

few small points at their distal end, Costal denta-

tion, however, ts never as well-developed as the

septal teeth and the dental ends of cosrae often

show random orientation of points (Fig. 1 IA).

In particular. the paddle structure, which appears

to represent the highest degree of dental orgatti-

zation, remains rare on costal teeth and was only

observed in colony 631.

Coenosteum

Coenosteum consists of exothecal dissepiments

similar in thickness and shape to their endothecal

counterparts.

Epjtheca

Epitheca is telativcly thin and continuously sur¬

rounds the cornllum external surface. Its orna¬

mentation consists of numerous thin para IJ el

transverse striations, sometimes slightly undulat-

ing. Morphological variabilit)' of epitheca afleets

its thickness and continuity, and the intensité of

its ornamentation.

THREE-DIMENSIONAL SKELETAL

DEVELOPMENT OF FAVIA FRAGUM

Calcification centres

Calcification centres hâve been defined as points

from which aragonitic fibres radiate. Alrhough

their nature and composition as well as the exact

process of calcification are still not fully under-

ZOOSYSTEMA • 1999 • 21 (2)

149

CuifJ.-P. & Perrin C.

Fig. 9, — Favia iragum (Esper, 17951, SEM, columella and wall, A, specimen FAV560-4. view of a calice wîlh spongy columella

developed from fusion of axial septal margins; B. specimen 631. central part of calice with spongy columella resultlng front Iwisting

and tangling of axial teeth; CF, specimen 1260: C, axial margins of septa and their ralafioriship wilfi columella; D, early stage of sep-

tothecal wall formation, latéral expansion of fhe neighbouring septa Ihrough adjunction of calcification centres, the large areas to left

and right are septa: E, calcification centre and surrounding aragomtic fibres wlthin thp wall. delail of F, F. laler stage of wall formation

showing the development of a new radial element; structures at top and bottom are septa. Scale bars: A, B. 1 mm; C, 200 pm; D-F,

100 pm.

150

ZOOSYSTEMA ■ 1999 ■ 21 (2)

Skeletogenesis in Favia fragum

Fig. 10. — Favia tragum (Esper, 17951. successive stages of wau formation, traced irom SEM photographs ol specimen 616. axial

part of calice is beyond (lie top ol A, B and C: A. early stage ol watt formation |us| atter junctlon ot both latéral expansions (rom

neighbouring septa, note trie calcification centres broadly arranged In line parallel to the wall and tbe small notçft on the extemal

margin of wall ïndicafing the point of junction; B. ttiickening of wall by adjunction of calcification centres on its internai side, see also

Fig. 9D: C, lalei siage ol wall formation showing the earliest calcification centies allgned parallel to the wall and the later centres

arranged perpendicular to fhe wall and contributing to the formation of a new septum, see also Fig. 9F. Scale bars: 100 pm.

stood, thcy obviously correspond to starting

points of aragonite fibres, Thereforc, their distri¬

bution in space and cime strongly Controls mor-

phological and structural development of coral

skelcton. StM observations of slightly etched

transverse sections ol septa and walls ol Favia

fragum show calcification centres as non-fibrous

crystallized striictures contrasting with the sur-

rounding incrémental layers of aragonite

(Fig. 12A, B), fi should be tiuted, however, thar

the appearance ol iliese centres is highlv variable

according to sarnple préparation and is particu-

larly sensitive to etching conditions (Fig. 12).

Analysis ol Favia fragum skeleton shows that sep¬

tal and costal micromorphologies, as well as the

columella and scptothecal wall are structures

dirccdy depending on the threc-dimensional loca¬

tion of calcification centres. Centres described

from rhin sections in previous Works (Alloitcau

1957: Zlatarski &C Martinez Hstalella 1982) appcar

as dark large dots (Fig. 13C) or as séries of centres

forming a dark line when septal teeth are indincd

to the axial margin. SF.M observations, however,

show the.se previously described large dots corres¬

pond in fret to groups of centres approxiioately

aligned normal to the septal plan. Th U arrange¬

ment occurs both in septa and costae but is also

involved in formation of the columella and the

septothccal wall. This thcrelore seems to reflect

the skeletal growth process of Favia fragum.

ZOOSYSTEMA • 1999 • 21 (2)

151

Cuif J.-P. & Perrin C.

Fig. 11. — Fa via Iragum (Esper, 1795). SEM. costal (tentation; A, specimen 616. weil-developed costae with costal teeth bearing

group of points; B, specimen FAV560-2, costal dentation showing simple conical teeth on the ieft. Scale bars; A, 200 pm; B, 100 gm.

Fig. 12. — Favla Iragum (Esper, 1795), SEM, calciticatlon centres; A. specimen 61S-2, transverse section slightly elched for

40 seconds with a solution ol lormic acid i% and glutaraldehyde 3%: B„ detail ol A, same préparation, calciticalion cenlre clearly

appears as non-tibrous crystalline structure; C. specimen 631. transverse section prepared with an enzymatic solution ot alcalase

pH 7,5 for 3 hours 35 minutes, calciticalion centres and then immédiate surroundlng incrémental layers are strongly etched; D, detail

of C, same préparation, internai structure of calcification centre cannot be seen from this type of préparation. Scale bars; A, C.

100 pm; B, 10 pm; D. 50 pm.

152

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia fragum

Fig. 13. — Favia fragum (Esper, 1795), thin sections ffom lhe Alloiteau Collection, MNHN Paléontologie; A, 512451a, transi/erse

section in the wall area; B. SI 2451 b, transverse section showing the wall and the peripheral part of calice; C. detail ot B. Scaie Dars:

A. 1 mm; B, 500 pm; C 200 pm,

SKELETAJ. GROWTH

Development of coral skeleton results from

incrémental growth shovvn by concentric lavers

ot fibrous aragonite around calcification centres.

This forms groups of points occuiring at tooth

ends of distal margin ot septa and costae, each

point resulring from growth around a calcifica¬

tion centre. Paddlc structures are therelorc form-

ed from regttlarly aligned calcification centres,

the arrangement ot points at distal margin retlect-

ing the arrangement of rliese centres wirhin the

septal microstructure. Sériés ol centres are paral-

lel to growth direction and bran,ched axis of cal¬

cification centres was never observed.

Length of aragonitic fibres, and therelorc thick-

ness of incrémental growth layers, vary according

to growth direction. This differential develop¬

ment of aragonite crystals Icads ro the burial of

previously formed points on the latéral septal

faces and also to the graduai infilling of space

between neighbouring reeth along the distal mar¬

gin of septa (Fig. 8).

SEM observations of the distal septal margin

show that aragonitic crystals within sets ot fibres

are clearly oblique to the external surface of sep¬

tum (Fig. 6) contrary to the widely accepted view

that fibres are perpendicular to the basal ecto-

dermal layer.

DISCUSSION

To date, the Milne Edwards specimens of Favia

fragum may lie regarded as the oldest available

material relevant to the historical development of

its species définition. Moreover, the first descrip¬

tions ol septal and thécal microstructures of

Favia fragum (Alloiteau 1957; / ! a t a rs k i &

Martinez Estalella 1982) both included morpho-

logical observations of the Milne Edwards speci¬

mens.

From the generic characterization by Milne

Edwards (1857: vol. 2, 426) to the latest treat-

ment by Véron (1986: 450), morphological rcla-

tionships berween adjacent corallites hâve been

used to discriminate between Favia and related

généra. For both authors, colonies belonging to

Favia are plocoid while Favites Finit, 1807, the

elosest genus, is charactcrized by cerioid colonies.

Variability of this geometrical arrangement leads

ZOOSYSTEMA • 1999 • 21 (2)

153

Cuif J.-P. & Perrin C.

Fie. 14. — Favia fragurn (Esper, 1795), original figures from

Alloiteau 1957: A, septal ornamentation (Alloiteau 1957: fig. 98);

B, transverse section of septothecal wall at the junciion of four

calices (Alloiteau 1957: fig. 96). Scale bar. A. no scale: B. 1 mm.

Veron (1986; 451) [o consider, howcvcr, that the

generic distinction based upon plocoid versus

cerioid arrangement may be somewhat “arbitra-

ry”. This is well illustrated by the variability in

calicinal pattern in Milne Edwards' spécimens.

The budding process givlng risc- to “equal cotai

lires” in Favia and to “corallites of different si/.c

in Favites is ajso regarded by Veron (1986) as a

subsidiary distinctive character.

This establishcd use of purcly morphological

characters for defining supra spécifie taxa lias to

be related to diverse commènes on the unsatisfact-

ory State of classification of Favia and related

généra (e.g. Vàughatl 1918) and to the paradoxi-

cal remark of Veron (1986. 445), çonsidering the

family haviidae, thaï sometimes fspecies may be

more readily recognized than genus”.

The first attempt to introducc additional criteria

at generic level was made by Alloiteau (1957:

138), who accurately described tlie micro-

morphology ot septal growing edges, the orna¬

mentation of septal sides of Favia fragurn

(Alloiteau 1957: fig. 98) (Fig. 14A) and also

made the firsi observations concerning septal

microstructure. Curiously, little was donc in this

study for coordinating ail morphological and

microstructural data which are presented separa-

tely. Because oi recent improvements in under-

standing the mechanisms of invertebrate

biomineralization and irnproved imaging which

facilitâtes comparison of similar-scaled picrures

of micromorphology and microstructure, it is

now possible to integrate ail morphological and

microstructural data on Favia fragurn within a

single description of its skeletal growth.

J he basic unit controlling septal organization

within the colonies of the Milne Edwards

Collection is the denriculate group of calcifica¬

tion centres whose typical arrangement is best

seen on the middlc part of distal septal margins.

As internai arrangement of chcsc basic growth

unies dues not correspond to any previously-

described type of uabeculae, we prefer to describe

die micnosiructure for itself wirhour referring to

usual terniinology. Each growth unit comprises

up to six or seven centres arranged in a transverse

alignmenr normal to rhe septal plan. The most

latéral of ihese centres are not completelv buried

by the lurther growth of fibres during the later

stages of septal development and therefore

remain visible on the latéral faces ofsepta. This is

the origin ot the sériés of granules illustrated by

Alloiteau (1957: fig. 98) (fig. I4A) and regarded

by him and latc-r by Zlatarski & Martinez

Esralella (1982) as latéral ornamentation.

However, tliese sériés of granules, the strength

and conrinuity ot which dépend on the regulari-

ty ot the growth process, hâve to be considered as

part of the basic septal plan and not as ont,inten¬

tai {= supetftcial) features. At this point, atten¬

tion must be Jrawn to the possible misinter-

pretation that may resuit from. observation of the

septal faces alctnc. In particular, lincar séries of

granules may be produced by a number of differ¬

ent septal m icrosttuctures (tf. description of sep¬

tal structure in ManFivaltia Eamouroux, 1821

(Cill & Eafuste 1971). Only relevant comparison

between septal face morphology, septal growing

margins (where first stages of calcification are

seen) and septal microstructures allow septal

organization to be corrcetly understood.

As alrcady mentioned in the above descriptive

part, rhe basic growth unit of septa in Favia fra-

gum may be composcd ot a variable number of

calcification centres, which gives risc to a first

level of variability between septa ol successive

cycles within the sanie calice and between septa

of different corallites. From this point of view,

154

ZOOSYSTEMA • 1999 • 21 (2)

Skeletogenesis in Favia fragum

specimen 631 appears remarkable for its régula ri-

tv and clac well-expressed organization of its sep¬

tal margins. In mhcr spécimens, despite a higli

morphological variability in size and shape of

calices, tlie sa me basic structure of septal growth

units occur: The variability in number and deve¬

lopment of calcification centres in each growth

unit cannor be attributed to subspecies distinc¬

tions or to controlling ecological parameters

since thèse arc unknown for the Milne Edwards

specimens.

The range of microstructural variability may be

clearly assessed by taking into account changes of

mîcrostructural features at the distal ends of sep¬

tal growrh units front the axial septal margin to

the costal margin. Ar rhe axial septal margin, rite

basal parts of septal growth units beeome very

elongated, Icading to the formation of digitiform

septal extensions (Fig. 5A-C . called “axial teeth

by previons authors (Alloiteau 1957; Martinez

Esralella 1982). As mentioned by these aurhors,

the spongy columella is exclusively built by these

septal expansions, and this agrées wcll with our

own observations (Fig. 9AHowever, these

“axial réélit" sltould not be regarded as distinct

features of septal architecture. Round calcifica¬

tion centres aie clearly scett on their tips and

continuous growth axes occur within their elong-

ated parts. This indicatcs that their internai

structure and therefore their growth pattern, are

basically the sanie as for typical septal growth

units of the rruddle part ol septal margins.

Clearly, “internai teeth' of other authors resuit

from only slight modifications in growth pattern

of the Liasic septal units.

Different patterns of growth variation can also be

nbserved on rhe cn.xtal margins and are mainly

due to variations in the degree of development ol

costal teeth and to a reduced number ol points at

their distal ends (Kg, 11). In ail cases, however,

rhe santé basic schente of the septal growth unit

can be recognized.

In Fiis introductory cbapter to the Family

Faviidae, Véron (1986: 443) qualiFied the septal

structure of these cotais as “simple", in the

Treatise of Pjjlcontoiogy, Wells (1956: F400)

regarded the simple trabcculae as a diagnostic cri-

teria for the suhlamily Faviinae. Analysis ol

Milne Edw'ards specimens of Favia firagnm sug-

gests that this canttoi be applicd to the septal

structure of this “key species”. First, accurate

observation of septal arrangement shows that, in

spire ol the morphological différences and varied

origips of the specimens, the basic septal archi¬

tecture remains cite sanie. Secondly, rite arrange¬

ment of calcification centres within individual

growth units always shows a polycentric pattern,

excluding a simple trabecul.tr structure. As the

alternative terni of conipound trabeculae may

include very different microstructural arrange¬

ments, these W'ould hâve to be analyscd befote

reassessing tlie significance of polycentric pat¬

terns. Finally, this iltree-dimcnsional arrange-

menr resulting front précisé posicioning of

calcification centres and permanent comrol of

libre growth direction, is strictly maintained

during successive growth stages, whatever growth

unit is considered.

CONCLUSION

By usingand comparing both morphological and

structural data on different scalcs within and bet-

vveen the studied colonies of Favia fragum, a pre-

liminary assessment ol potentially diagnostic

criteriacan be made.

Chnractcrs with a relatively high potential for

skclet.il plasticity are tltose related to tlie external

geometry of calices including their si/.e and

shape, t 11 ci r relat lonships with surmunding

calices, the length of Costa? and abondance of

coeitOSteum, To a lesser degree, some intracali-

cinal éléments may also vary in size and dotai lcd

shape, i.e. columella and distal dentation of septa.

Variation of wall aspect appears to he related to

the successive stages ol wall formation and there¬

fore is direct ly dépendent on septal development.

Constant charactcrs tltat may he used as discrimi-

nating criteria for taxonomy include number of

cycles and budding process, but also distribution

and arrangement of calcification centres which

cotttrol three-dimensional growth process and

therefore, micromorphnlogy, These resu Ils

emphasize the nced for taxonomie studies eom-

bining the 2D-analysis of seleractinian micro-

and ultrastructure with 3D-observations of

micromorphology and morphology. This inte-

ZOOSYSTEMA ■ 1999 • 21 (2)

155

Cuif].-!>.& Perrin C.

gratetl approaeh can be used to obtain a better

understanding of die growth process and its con-

trol. It aiso appears more promising than basing

scleractiman taxonorny only on analogies and on

différences of shape and macromorphology.

Acknowledgements

The authors are especially grateful to Professor

D. Doumenc (Laboratoire de Biologie des

Invertébrés marins et Malacologie, Muséum

national d’Histoirc naturelle) for loan of spéci¬

mens and lor permission of preparing them for

SEM observations. Wc also vvisli to thank for

their help the technical staff of the Laboratoire

de Paléontologie, Muséum national d’Histoire

naturelle and of the Département des Sciences de

la Terre, Université Paris-Sud: C. Chancogne

(MNHN) for technical assistance in operatmg

the SEM, D Serrette (MNHN) for macro-

phorographs, A. Denis and J. Didclot (UPS) for

preparing micro- and SEM-photographs. We

also thank Professor Eva Roniewicz (University

of Warsaw) and Dr Brian R. Rosen (The Natural

History Muséum, London) for their critical

review of the manuscript.

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Sciences 1 1: 63-168.

Wells J. W. 1956. — Selcractinia: part F, 32.8-444, in

Moore R. C. (ed.), Treatisc an invenebratc palennto-

logy. Ceological Sodcry ol America; University of

Kansas Press, Lawrence, Kansas.

Zlatarski V. N. tk Martine/. Esialdla N. 1982. — Les

sclêractiniaires de Cuba avec des données sur les orga¬

nismes associés. Académie bulgare des Sciences,

Sofia, 472 p.

Submitted un 23 fanuary 1998;

accepted on 3 July 1998.

156

ZOOSYSTEMA • 1999 • 21 (2)

Trois nouveaux Heterocotyle (Monogenea,

Monocotylidae) parasites branchiaux de

Taeniura grabata (Euselachii, Dasyatidae)

en Tunisie

Lassâd NEIFAR

Laboratoire de Biologie et Parasitologie marines, Faculté des Sciences,

Université Tunis II, 1060 Tunis (Tunisie)

Louis EUZET

Station méditerranéenne de l’Environnement littoral, Université Montpellier II,

1 Quai de la Daurade. F-34200 Sète (France)

euzet@univ-montp2.fr

Oum Kalthoum BEN HASSINE

Laboratoire de Biologie et Parasitologie marines, Faculté des Sciences,

Université Tunis II, 1060 Tunis (Tunisie)

MOTS CLÉS

Monogenea,

Monocotylidae,

Heterocorylinae,

Heterocotyle,

parasite,

élasmobranch»,

Dasyatidae,

Taeniura grabata,

branchies,

Tunisie.

Neifar L., Euzet L. & Ben Hassine O. K. 1999. — Trois nouveaux Heterocotyle

(Monogenea, Monocotylidae) parasites branchiaux de Taeniura grabata (Euselachii,

Dasyatidae) en Tunisie. Zoosystema 21 (2) : 157-170.

RÉSUMÉ

Trois nouveaux Heterocorylinae parasites branchiaux de Taeniura grabata

péchés en divers points de la côte tunisienne, sont décrits : Heterocotyle

mokhtarae n. sp., H. striata n. sp. et H, forcifera n. sp. Heterocotyle mokhtarae

et H. striata ont un hapteur avec plusieurs crêtes sinueuses disposées selon la

formule I - I -2-3-3. Chez H, forcifera il n’y a qu'une seule crête sinueuse. H.

mokhtarae a un pénis tubulaire et un bulbe éjaculateur trois fois plus long

que le pénis. H. striata a un pénis conique orné d’une formation ovoïde en

lame à bords finement striés. H. forcifera a une pièce accessoire à deux

branches denticulées soudée au pénis tubulaire. La limite des critères définis¬

sant le genre Heterocotyle et l'importance systématique des caractères invo¬

qués, nombre de crêtes sinueuses sur les septa du hapteur et présence d'une

pièce accessoire dans l’appareil copulateur, sont discutés. On met en évidence,

entre les trois espèces, l’existence des différences constantes dans le nombre

de sinuosités formées par les crêtes des septa. Une clé du genre Heterocotyle

est proposée.

ZOOSYSTEMA • 1999 • 21 (2)

157

Neifar L., Euzet L. & Ben Hassine O. K.

KEYWORDS

Monogoncrt,

Monocotylidiie.

Heterocotyliniic,

Hetetùcotylt ?,

parashe,

elasmobranch,

Dasyaticlar,

Taeniura grabata ,

giüs,

Tunisia.

ABSTRACT

Three new Heterocotyle (Monngcnea, Manocotylidae) gillparasites of Taeniura

grabata (Èuselachii, Düsydtidae) jrom Tunisia.

Description are given for three Heterocotylinae. gills parasites of Taeniura

grabata collected front various Iocalities oi tire Tunisian coast: Heterocotyle

mokbtarae n. sp., //, striata n. sp., and H. forcifera n. sp. The haptor of

H. mokbtarae and H. striata has numerous «muons septal ridges and

1-1-2-3-3 configuration. The haptor of II. forcifera has only onc sinuous sep¬

tal ridge. H. mobtarac has a tuluilar pénis, and an ejucul.uory bulb three

tintes as long as the pénis. H. striata has a conit penî.s wllh an ovoid slide-

shaped piece with a slightly striated edgv, H. forcifera has an accessory piece

with two jagged branches fused to the tubtilar penis. The limit.s of the criteria

defining the genus Heterocotyle and the systcmatic importance of the charac-

ters used, the number of sinuous ridges on the septa of the haptor and the

presence of an accessory piece associated with the copulatoty apparatus, are

discussed. Constant différences are evidenced for the number of sinuosides

on che septal ridges between these three species. A key of the genus

Heterocotyle is proposed.

INTRODUCTION

L'originalité de la faune des élasmobranches de

Tunisie résulte en grande partie de la présence,

en particulier dans la région méridionale (golfe

de Gabès), de plusieurs espèces à affinités tropi¬

cales. Taeniura grabata (E. Geoffroy Saint-

Hilaire, 1817) est une des ces espèces Signalée

en Méditerranée de la Tunisie à l’Égypte, cetre

pastenague n’est pas citée sur les côtes de

l’Algérie et du Maroc (McEachran & Capapé

1984). Sur la côte atlantique de PAlrique, clic ne

paraît exister qu'au sud du Cap Vert.

Nous avons récolté sur les branchies de Taeniura

grabata de nouveaux Monogenea dont trois sont

à classer parmi les Heterocotylinae Chisholm,

Wheeler & Bevériey-Burton, 1995. La morpho¬

logie du hapteur avec quatre formations striées

dorsales, la présence de septa radiaires et circu¬

laires marqués veimalemenc par des crêtes

sinueuses situent ces trois espèces dans le genre

Heterocotyle Scott, 1904. Nous les considérons

comme nouvelles er leur description constitue

l’essentiel de ce travail.

MATÉRIEL ET MÉTHODES

Les sept Taeniura grabata examinées proviennent

de la pêche côtière, cinq au nord dans le secteur

de Bizerte (27.11.1996 ; 18.V.1996 ; 07.VI.1996 ;

25.III. 1997 et 25.111.1997) et deux au sud dans

le secteur de Xar/.is (10.VI.1996 ; 1 1, VI. 1996).

Les poissons sont nécropsiés le plus rapidement

possible après leur capture. Les arcs branchiaux,

séparés par incision dorsale et ventrale, sont pla¬

cés dans des boîtes de Pétri emplies d’eau de mer

filtrée. Les monogènes repérés au stéréomicrosco-

pe entre les lamelles branchiales sont récoltés

vivants et étudiés soir directement au microsco¬

pe, soit fixés légèrement aplatis entre lame et

lamelle. l'alcool à 70°, le formaldéhyde à 5 %, le

mélange de Bouin-Hollande sont les fixateurs

utilisés. Les individus fixés sont lavés par plu¬

sieurs bains d'eau distillée puis colorés avec le

carmin au borax alcoolique de Grenadier ou le

carmin acétique de Sémiçhon, Après déshydrata¬

tion er passage dans l'essence de girofle, ils sont

montés entre lame et lamelle au baume du

Canada. Plusieurs individus sont fixés colorés et

158

ZOOSYSTEMA • 1999 • 21 (2)

Heterocotyle nouveaux de Taeniura grabatc

montés directement entre lame et lamelle avec le

mélange picrate d’ammonium-glycérinc selon

MaLmberg (1957). Après quelques heures, néces¬

saires à la diffusion du milieu de montage, ces

préparations sont lutées avec du Glyccel

(GURR®).

Immédiatement après leur dissection, quelques

arcs branchiaux sont fixés in toto dans une solu¬

tion de formaldéhyde à 5 % ou dans de l’alcool

éthylique à 75°. Les parasites récoltés entre les

filaments de ces arcs sont comme précédemment

lavés, colorés et montés. Afin de préciser la mor¬

phologie des pièces sclérifiées du hapreur et du

système génital, quelques individus ont été éclair¬

cis et montés dans la gomme au ehloral de

Berlèse. Ces préparations sont lutées avec du

Glyceel.

Nous n'avons pas noté la position exacte des dif¬

férentes espèces sur les ares branchiaux. Un petit

nombre d individus ont été observés entre les

lamelles de la hranchlc hyoïdienne.

Les dessins sont réalisés avec tin tube à dessin.

Toutes les mesures minimum-maximum

(moyenne) sont données en micromètres. Pour

unifier la nomenclature nous utiliserons « unci-

nulus » pour les petits crochets larvaires (croche¬

tons ou « hookk-ts •■) (Pariselle &T Fuzet 1995)

qui persistent dans la membrane marginale du

hapreur. Nous n utilisons pas « gripus ■ pour les

grands crochets car nous n’avons pas de données

sur leur ontogenèse. Pour désigner les divers

septa er loculi, nous suivons la nomenclature

proposée par Neffar et al ( 1998) dans leur travail

sur les Heterocotyle parasites de Dasyath pastinaca

(Linné, 1758).

Dans le genre Heterocotyle Scott, 1904, la surface

des septa est marquée, selon l’espèce, par un

nombre variable (un, deux ou trois) de crêtes

sinueuses. Dans le cas où il existe plusieurs

crêtes, nous nommons « crête médiane » celle qui

est sur le plan médian du septum et « crêtes (ocu¬

laires » celles qui sont sur les flancs du septum.

Nous précisons la disposition des uncinuli chez

Heterocotyle , car nous pensons que, chez les

Heterocotylinae, elle peur aider à comprendre les

rapports existant entre les différentes morpholo¬

gies du hapreur. Les 14 uncinuli sont situés dans

la membrane marginale avec une disposition

symétrique. De chaque côté, les uncinuli I et

II sont au niveau du loculus postérieur, I plus

près du plan de symétrie ; les uncinuli III et

IV sont au niveau du loculus postéro-latéral,

III plus près des crochers ; l’undnulus V esr au

niveau de l’extrémité du septum latéral, l'uncinu-

lus VI au milieu du loculus antëro-latéral et le

Vil au milieu du loculus antérieur (Fig. 1 B).

Dans l’appareil copulateur mile nous distin¬

guons deux parties, le pénis creux par où passe le

sperme et les sécrétions du bulbe éjaculateur et la

pièce accessoire soudée sur la base du pénis.

Cetre pièce n’existe pas chez toutes les espèces.

Chez les trois espèces décrites, le pénis étant peu

courbé la mesure donnée est celle de la droite joi¬

gnant les extrémités proximale et distale. La

mesure des branches arquées de la pièce accessoire

de H. forcifera n. sp. représente la corde de l'arc.

Dans ce travail nous enrendons les Herero-

cotylinae selon la définition donnée par

Chisholm et al. (1995) cr le genre Heterocotyle

avec la diagnose revue par Chisholm &C.

Whittington (1996).

Le materiel original (holotype et paratypes) est

déposé au Muséum national d’Histoire naturelle

(MNHN) et au Natural History Muséum

(BMNH), London.

SYSTÉMATIQUE

Heterocotyle mokhtarae n. sp.

(Figs 1 ; 2)

Holotype. — Déposé au MNHN numéro 593 HP

Tk 131. Paratypes (10) déposés au MNHN numéros

593 HF Tkl32 à 593 HP Tk 135. Paratypes (10)

déposés au BMNH numéros 1998.4,3.6. à

1998.4.3.12

HÔTE-TYPE. — Taeniura grabata (E. Geoffroy Saint-

Hilaire, 1817).

LOCALISATION. — Branchies, entre les lamelles bran¬

chiales.

I .ÔCALITÉS. — Bîzerte, Zar/is (Tunisie).

MATÉRIEL ÉTUDIÉ. — 50 individus, mensurations sur

25, Poissons examinés : 7. Prévalence 100 %.

ÉTYMOLOGIE. — Nommé en hommage à Madame le

ZOOSYSTEMA • 1999 • 21 (2)

159

Neifar L., Euzet L. & Ben Hassine O. K.

Professeur F. Mokhtar-Maamouri de l’Université

Tunis II.

Description

Heterocotylinae. Individus adultes mesurant,

sans le hapteur, I 200-1 650 pin (I 400 pm) de

longueur et 360-560 pm (470 pm) de largeur au

niveau de l’ovaire. Hapteur étiré transversale¬

ment, un peu plus large que le corps, 380-

600 pm (515 pm) (Fig. 1 A). Surface ventrale du

hapteur subdivisée en un luculus' central et huit

loculi périphériques' (deux loculi postérieurs,

deux postéro-latéraux, deux antéro-latéraux et

deux antérieurs). Huit septa radiaircs (un médian

postérieur, deux postéro-latéraux, deux latéraux,

deux antéro-latéraux, un médian antérieur) entre

un septum circulaire intérieur entourant le locu-

lus central et un septum circulaire extérieur mar¬

quant la base d’une membrane marginale. Septa

surmontés ventralement par des crêtes sinueuses,

une sur le septum médian postérieur et les septa

postéro-latéraux, deux sur les septa latéraux, trois

sur les septa antéro-latéraux et le médian anté¬

rieur. Septum circulaire intérieur avec deux crêtes

sur sa moitié antérieure et une crcte sur sa moitié

postérieure. Septum circulaire extérieur avec une

seule crête (Fig. IB). Un crochet, long de 40-

45 pm (44 pm) du bout du manche à la courbure

de la lame, situé à l’extrémité postérieure de

chaque septum postéro-latéral. Quatorze uncinuli

longs de 9-10 pm situés dans la membrane mar¬

ginale avec la disposition symétrique décrite plus

haut. Quatre structures arquées, à aspect strié,

composées d’une palissade très serrée de minus¬

cules lamelles triangulaires sclérifiées. Structures

saillantes à la lace dorsale du hapteur, au niveau

des loculi postérieurs et postéro-latéraux

(Fig. iQ.

Bouche ouverte au fond d’une large dépression

ventrale antérieure entourée par des fibres mus¬

culaires laissant libre une lèvre ventrale. Glandes

médio-antcricurcs et latéro-antérieures débou¬

chant sur le bord de cette lèvre. Pharynx en

barillet mesurant 155-210 pm (180 pm) de lon¬

gueur et 95-170 pm (118 pm) de diamètre, avec

une petite partie buccale entourée par un faible

sphincter et une grande partie postérieure mar¬

quée par sept couches épaisses de fibres radiaires.

Fibres musculaires latérales fixées au pharynx à la

limite entre les deux parties et disposées en éven¬

tail dans le corps. Glandes œsophagiennes,

situées de chaque côté du pharynx et débouchant

à sa base. Deux branches intestinales simples ter¬

minées en cul-de-sac dans la région postérieure

du corps. Un petit nombre de granules pigmen¬

taires dorsaux de chaque côté du pharynx

(Fig. 1 A). Glandes postérieures non observées.

Vessies du système osmo-régulateur dilatées et

visibles, chez quelques individus dont l’holotype,

une de chaque côté immédiatement en arrière du

pharynx.

Testicule médian postérieur entre les branches

intestinales. Canal déférent visible, chez les indi¬

vidus vivants, sur le côté gauche du corps entre le

vagin et la branche intestinale, difficilement

observable chez les individus fixés et colorés.

Canal déférent passant sur le côté droit au niveau

du bulbe éjaculateur puis dilaté en une vésicule

séminale aboutissant, avec de nombreuses

glandes latérales, à la base d’un long bulbe éjacu¬

lateur de 300-400 pm (345 pm) de longueur et

45-70 pm (60 um) de largeur. Bulbe contenant

des files de granules représentant les sécrétions

des glandes latérales et, dans sa partie basale,

deux cavités en U, plus ou moins développées

(Fig. 2A). Sécrétions et sperme passant entre ces

cavités. Pénis tubulaire simple mesurant 145-

165 pm (154 pm) de longueur et 10-12 pm

(1 i pm) de diamètre. Base élargie en entonnoir

coiffant l’extrémité antérieure du bulbe éjacula¬

teur. Parrie distale évasée marquée par une côte

longitudinale et une ouverture à bord spiuulé

(Fig. 2B). Paroi de la gaine du pénis légèrement

épaissie dans sa partie terminale près de l'ouver¬

ture génitale. Ouverture vaginale sub-latérale

gauche. Vagin, 90-150 pm (121 pm) de lon¬

gueur er 45-90 pm (60 pm) de largeur, entouré

par une couche de cellules glandulaires. Chambre

vaginale, à paroi plissée légèrement épaissie,

mesurant 180-300 pm (223 pm) de longueur sur

40-130 pm (87 pm) de largeur. État de vacuité

de cette chambre lié à la présence (17 individus

sur 50) d’une masse très polymorphe assimilée à

un spemvacophore. Chambre vaginale terminée

par un étroit canal se jetant dans un réceptacle

séminal peu développé. Ootype médian, 4 paroi

épaisse, avec une pairie diSLalc élargie ouverte au

pore génital ventral et une partie étroite marquée

160

ZOOSYSTEMA • 1999 • 21 (2)

Fig. 1 . — Heterocolyle mokhtarae n. sp. A. animal in toto (hololype), vue ventrale ; B, hapteur, vue ventrale ; C, hapteur, vue dorsale.

I à VII uncinuli. Échelles : 250 pm.

à la base par le débouché des glandes de Mehlis.

Dans l’ootype, œuf tétraédrique, mesurant 85-

100 pm (92 pm) de côté, avec un filament opposé

à l’opercule.

Remarques

Le hapteur avec huit loculi périphériques autour

d'un loculus central, des septa à crête sinueuse et

surtout quatre structures dorsales striées placent

ZOOSYSTEMA • 1999 • 21 (2)

Neifar L., Euzet L. & Ben Hassine O. K.

Fig. 2. — Heterocotyle mokhtarae n. sp. A, bulbe éjaculateur

avec cavités pcstérieures ((lèches) ; B, pénis. Échelles : 50 pm.

ce Monocotylidae dans le genre Heterocotyle. La

longueur du pénis, supérieure à 150 pm, permet

de séparer cette espèce de tous les Heterocotyle

connus, sauf de H. confina Timofeeva, 1983.

H. mokhtarae n. sp. se distingue de H. confina

par le bulbe éjaculateur beaucoup plus long que

le pénis et par la morphologie du pénis avec une

côte dans sa portion distale. Nous considérons ce

parasite de T. grabata comme une espèce nouvelle.

Heterocotyle striata n. sp.

(Fig. 3)

I lOLOTYl 1 ! . — Déposé au MNHN numéro 591 HF

Tkl20 Paratypes (10) déposés au MNHN numéros

591 HF Tk 121 à 591 HF Tk!25. Paratypes (10)

déposés au BMNH numéros 1998.4.3.1 à 1998.4.3.5.

HÔTL-Tv l’F. — Taeniuru grabata (E. Geoffroy Saint-

Hilaire, 1817).

LOCAI IsATION. — Branchies, entre les lamelles bran¬

chiales.

Localité. — Bizerte, Zarzis (Tunisie).

Matériel étudie. — 50 individus, mensurations sur

25. Poissons examinés : 7. Prévalence 100 %.

ÉTYMOLOGIE. — Nommé à cause de l’aspect strié du

pénis.

Description

Adultes mesurant, sans le hapteur, 1 100-

1 500 pm (I 260 pm) de longueur et 310-

550 pm (430 pm) de largeur au niveau de l’ovai¬

re (Fig. 3A). Hapteur postérieur étiré

transversalement, plus large que le corps, et

mesurant 400-600 pm (498 pm). Septum

médian postérieur et septa postéro-latéraux avec

une crête, sepra latéraux avec deux crctes, septa

antéro-Iatéraux et septum médian antérieur avec

trois crêtes. Septum circulaire intérieur avec deux

crêtes sur la moitié antérieure, une sur la moitié

postérieure. Septum circulaire extérieur avec une

seule crcte (Fig. 3B). A l’extrémité postérieure de

chaque septum postéro-latéral un crochet de 50-

60 pm (55 pm) avec la pointe sensiblement égalé

au manche et à la garde. 14 uiuinuli (9-10) avec

la disposition décrite ci-dessus. Quatre structures

striées dorsales présentes. Pharynx 140-200 pm

(173 pm) de longueur et 90-140 pm (111 pm)

de diamètre en deux parties. Glandes oesopha¬

giennes peu développées. Un petit nombre de

granules pigmentaires latéro-dorsaux de chaque

côté du pharynx. Sur le bord apical de la lèvre

ventrale deux paires d'amas représentant le

débouché des glandes antérieures. Glandes posté¬

rieures non observées. Une ouverture medio-

ventrale entre l’ovaire et l'appareil copulateur.

Testicule médian postérieur, plus ou moins lobe,

paraissant parfois subdivisé en trois parties.

162

ZOOSYSTEMA • 1999 • 21 (2)

Heterocotyle nouveaux de Taeniura grabatt

Bulbe éjaculareur mesurant 180-250 fim

(217 pm) de longueur sur 60-11 1 pm (79 pm)

de diamètre avec deux cavités parfois visibles

dans sa partie basale. Glandes latérales avec un

groupe gauche développé de part et d’autre du

pharynx et un groupe droit moins important

entre le bulbe éjaculateur, l’ootype et la branche

intestinale. Glandes aboutissant, autour du canal

déférent, à la base du bulbe. Bénis en tronc de

cône, 130-155 pm (144 pm) de longueur et 28-

32 pm (29 pm) de largeur en son milieu, à base

élargie coiffant l'extrémité antérieure du bulbe.

Surface du pénis marqué dans le tiers distal par

deux rangées de Filaments régulièrement disposés

en peigne. Rangées contiguës à leur origine et à

leur extrémité. Filaments diminuant de taille

jusqu’à l'ouverture du pénis dont le bord parait

spinulé (Fig. 3C). Vitelloducres transverses for¬

mant toujours un U en avant de l’ovaire. Cham¬

bre vaginale à paroi plissée et légèrement

sclérifiée. Ootvpe très long formant un .S entre

les deux branches de I ovaire puis une anse anté¬

rieure avant l'élargissement terminal. Glandes de

Mehlis situées à gauche au niveau de la masse

initiale de l’ovaire. Dans l’ootype, œuf tétra¬

édrique de 7ü- 7 5 pm (72 pm) de côté, avec un

très long filament opposé à l’opercule.

Remarques

Nous plaçons ce Monocotylidae dans le genre

Heterocotyle en nous fondant sur la présence de

quatre structures striées faisant saillie sur la face

dorsale du hapteur, La morphologie du pénis-

place cette espèce à côté de H. pasrinàcae Scott,

1907 et H. çhinensis Timofeeva, 1983. Elle s’en

distingue par la présence de deux rangées de fila¬

ments qui donnent au pénis une allure striée

Dans le hapteur de Heterocotyle striata, si la dis¬

position générale des crêtes sinueuses est du type

1-1-2-3-3, il existe, avec les espèces du genre

ayant cette disposition, de légères différences,

Ainsi, les crêrcs Incnlaires qui doublent la crête

médiane des septa latéraux, anréro-latéraux et du

septum médian antérieur ne s’attachent pas exac¬

tement de la même manière sur la crête du sep¬

tum circulaire extérieur. Cher H. pastimeae et

chez H. mokhtarae , les crêtes loculaires se termi¬

nent au niveau de sinuosités contiguës à celle sur

laquelle s’attache la crête médiane. Chez H. striata ,

il y a régulièrement deux sinuosités entre

l’attache de la crête médiane et celle des crêtes

loculaires (Fig. 3B).

Nous avons observé une masse ovoïde, que nous

nommons spermatophore, plantée au niveau de

l’ouverture vaginale. Les individus avec la

chambre vaginale emplie par une masse compa¬

rable sont rares (7/50).

La présence et le rôle de l’ouverture médio-ven-

tralc reste un problème que nous espérons

résoudre rapidement.

Heterocotyle forcifera n. sp.

(Fig. 4)

Hotui vi'h. — Déposé au MNFIN numéro 592 HF

Tkl26. Paratypes (10) déposés au MNHN numéros

592 H F Tk 127 à 592 HF Tkl30. Paratypes (10)

déposés à BMNH numéros 1998.3.4.13 à

1998.4.3.16.

HOtf.-Type. — Taeniura grabata (E. Geoffroy Saint-

Hilaire, 1817).

HABITAT. — Branchies, entre les lamelles branchiales.

Localité. — Bizerte, Zarzis (Tunisie).

Matériel étudié. - 50 individus, mensurations sur

25. Poissons examinés : sept. Prévalence 100 %.

ÉTYMOLOGIE. — Nommé pour rappeler la morpholo¬

gie en forceps de la pièce accessoire de l’appareil copu-

lateur.

Description

Adultes mesurant, sans le hapteur, 610-1 150 pm

(837 pm) de longueur et 260-460 pm (334 pm)

de largeur au niveau de l’ovaire (Fig. 4A).

Hapteur postérieur en cupule musculaire presque

circulaire plus étroite que le corps, 250-390 pm

(289 pm) de diamètre Tous les septa surmontés

par une seule crête médiane régulièrement

sinueuse. Membrane marginale large. Un crochet

de 45-55 pm (49 pm) de longueur à l’extrémité

de chaque septum latcro-postcrieur, Débouché

des glandes antérieures parfois visibles comme

deux paires d'amas de part et d’autre de l'extré¬

mité apicale. Glandes postérieures non observées.

Pharynx mesurant 122-177 pm (146 pm) de

longueur et 90-125 pm (106 pm) de diamètre.

Glandes œsophagiennes développées. Quelques

ZOOSYSTEMA • 1999 • 21 (2)

163

Neifar L, Euzet L. & Ben Hassine O. K.

Fig. 3. — Heterocalyle striata n. sp A., animal in toto (holotype), vue ventrale ; B, hapteur, vue ventrale détaillant les rapports entre

les crêtes septales médianes et loculaires et les sinuosités de la crête du septum circulaire extérieur (flèches) ; C, pénis, vue

dorsale ; D, pénis, vue ventrale. Échelles : A, 250 pm ; B, 100 pm ; C et D, 50 pm.

ZOOSYSTEMA • 1999 • 21 (2)

Heterocotyle nouveaux de Taeniura grabatc

rares granules pigmentaires (souvent totalement

absents) dans les régions latéro-dorsales du pha¬

rynx.

Testicule médian postérieur. Petit bulbe éjacula-

teur globuleux, 65-110 pm (93 pm) de longueur

et 45-80 pm (61 pm) de diamètre. Glandes laté¬

rales difficilement visibles. Pénis tubulaire légère¬

ment arqué, 33-55 pm (49 pm) de longueur,

présentant en son milieu quelques minuscules

épines superficielles. Pièce accessoire, soudée sur

la base du pénis, composée de deux bras en

demi-cercle portant sur leur face interne une

série de denticüles irréguliers (Eig, 4B, C). Vagin

tubulaire long de 40-50 pm (44 pm). Chambre

vaginale à paroi légèrement sclérifiée en deux

parties. l a première mesurant 45-55 pm (5U pm)

de longueur er 20 pm à 45 pm de largeur La

seconde plus régulière (30 x 20 pm) en tronc de

cône à paroi cannelée, reliée au réceptacle sémi¬

nal par un long canalicule. Petit réceptacle sémi¬

nal globuleux médian en avant de l'ovaire.

Ootype très long formant, sur le côté gauche du

corps, une boucle aplatie entre Povaite et le vagin

puis dessinant sur le côté droit une anse antérieure

avant l’élargissement terminal. Glandes de

Mehlis non observées. Œuf 50-95 pm (“2 pm)

de côté, avec un très long filament parfois pelo¬

tonné à l’intérieur de Pootype.

Remarques

Comme les précédentes, nous plaçons cette espèce

dans le genre Heterocotyle en nous fondant sut la

présence de crêtes sinueuses sur les septa et de

quatre structures striées saillantes à la surface

dorsale du hapteur. L’espèce que nous venons de

décrire se rapproche de H. armâtes Timofeeva,

1983 et de If. timcrkana llargis, 1957 par la

structure de l’appareil copulateur qui possède

une pièce accessoire soudée au pénis et pat la

morphologie du hapteur oit tous les septa ont

une seule crête sinueuse. Elle s'en distingue par la

pièce accessoire composée de deux branches au

lieu d’une seule chez H. armata et H. arfiericana.

Chez H. fortifiera nous n avons pas vu de grosse

masse dilatant la chambre vaginale. Chez un

individu, nous avons observé une petite structure

(spermatophore ?) plantée au niveau du pore

vaginal et chez un autre une masse de taille com¬

parable dans le vagin. En outre, dans la chambre

vaginale de plusieurs individus, il existe des amas

de spermatozoïdes.

DISCUSSION

Pour les trois espèces décrites les glandes anté¬

rieures et leurs conduits ne sont pas ou sont diffi¬

cilement visibles chez les individus fixés et

colorés. On peut seulement observer deux paires

d’amas de part et d’autre de l’extrétnitc apicale de

la lèvre ventrale. Sur le vivant, et chez quelque

individus éclaircis au Berlèse, la disposition des

glandes, de leurs conduits et de leurs débouchés

est identique à celle décrire par Chisholm Se

Whittingron (1996). Nous avons (Neilar et ai.

1998) retrouvé cette disposition chez les

Heterocotyle de D. pastinaca. Par contre nous

n'avons pas observé de glandes antérieures à

bâtonnets. Nous n’avons pas décelé la présence

de glandes postérieures.

Dans l’appareil copulateur des Heterocotyle ,

Chisholm &: Whittingron (1996) nomment

pièce accessoire deux formations qui nous parais¬

sent différentes. L'une est la pièce sclérifiée, à

morphologie variable, que l’on trouve soudée au

pénis chez H. umericana Elargis, 1955 et

H. ivrtnata Timofeeva, 1983 et H. fortifiera n, sp,

Le cas de H. dasyatis (Yamaguti, 1965) est parti¬

culier, le faisceau de baguettes ne paraissant pas

soudé au pénis (Chisholm 1995). L'autre forma*

tion est située près de l'extrémité distale du

pétris, Elle nous paraît représenter la partie épais¬

sie (sclérifiée ?) de la paroi de la gaine dans

laquelle coulisse l’appareil copulateur mâle pour

faire saillie à l’extérieur au moment de la copula¬

tion. Cette gaine est parfois bien visible autour

de l’extrémité distale du pénis de H. mokhuirae

(Figs 1A ; 2A).

Dans les Dàsyhatütrematinae, autre sous-famille

des Monocotylidae, Chisholm et al. (1995) ont

proposé de séparer, dans le nouveau genre

Timofeevia, l’espèce T. rajae (Timofeeva, 1983)

un parasite branchial de Raja porosa Günther,

18“4, décrit comme Dasybatolrêma rajae. Le

genre TiinofeeVut est supporté par une seule apo-

morphie, la présence d'une pièce accessoire bifide

fixée à la base d'un pénis tubulaire. En outre,

l’illustration originale de cette espèce montre que

ZOOSYSTEMA • 1999 • 21 (2)

165

Neifar L., Euzet L. & Ben Hassine O. K.

Fig. 4. — Helerocolyle fordtera n. sp. A, animal in toto (holotype), vue ventrale : B, C. appareil copulateur mâle : pénis (p) et pièce

accessoire (ap). Échelles . A, 250 pni ; B, C, 25 pm.

la morphologie générale du hapteur et celle des

crochets, sans long manche, ne correspondent

pas exactement à celle des Dasybcltotrema Price,

1938.

Chez H. armata et H. forcijèra , l’appareil copula¬

teur ressemble à celui de Timofeevia avec une

pièce accessoire soudée sur la base du pénis. Chez

H. amerieana Hargis, 1953, nous avons pu nous

assurer que la pièce accessoire est effectivement

soudée à la base du pénis. Cette pièce en crochet,

très légèrement spinulée dans sa concavité, pré¬

sente, de façon constante, une bulle ovalaire sur

sa convexité (Fig. 5). Notons que cette morpho¬

logie de l’appareil copulateur, avec un pénis et

une pièce accessoire soudée sur sa base, est carac-

téristique de très nombreux genres de

Dactylogyridea.

Par contre, chez ces espèces, la morphologie du

166

ZOOSYSTEMA • 1999 • 21 (2)

Heterocolyle nouveaux de Taeniura grabat.

Fis. 5. — Heterocotyle americana Hargis, 1955. Appareil copu-

iateur mâle : pénis (p) et pièce accessoire (ap) avec bulle ova¬

laire (flèche). Échelle : 50 gm.

hapteur est différente puisque chez H. ctrmata et

H.forcifera tous les septa sont surmontés par une

seule crête sinueuse et par une, deux ou trois

crêtes chez //. americana.

Parmi les Heterocotyle , si La présence d’une pièce

accessoire dans l'appareil copulateur représente un

lion critère spécifique, nous ne pensons pas qu il

puisse, comme d’ailleurs le nombre de crêtes sur

les septa, justifier actuellement la création de nou¬

veaux genres. Nous conservons donc Heterocotyle

avec la seule apomorphie : présence de quatre

structures striées à la surface dorsale du hapteur.

Chez certains individus colorés et montés, les

crêtes sinueuses apparaissent comme une succes¬

sion de points alternativement clairs et sombres.

Ceci représente peut-être un artefact résultant de¬

là technique de coloration au carmin acétique de

Sémichon. Mais cet aspect peut aussi traduire des

différences dans la structure même de la crête.

Sur les coupes, les crêtes apparaissent comme un

simple épaississement du syncytium légumentaire.

Une étude en microscopie électronique à trans¬

mission esr donc nécessaire pour connaître leur

structure exacte.

Comme Neifar et al. (1998) font signalé chez les

Heterocolyle parasites de Dasyatis pastinaca

(L., 1758) le nombre de sinuosités des crêtes

médianes et loculaires des différents septa est

fixe, ou varie dans de très faibles proportions. Il

en est de même chez les trois Heterocotyle para¬

sites de Taeniuragrabata. Ainsi, chez cinq indivi¬

dus de H. mokhtarae , nous avons compté 86-88

sinuosités sur le septum circulaire extérieur, 32-

34 sinuosités sur le septum circulaire intérieur,

sept pour le septum médian antérieur, huit ou

neuf pour les sepra antéro-latéraux, neuf pour les

septa latéraux, neuf pour les septa postéro-laté¬

raux et neuf pour Je septum médian postérieur.

Nous avons compté 14-16 sinuosités pour la

crête loeulaire des loculi antérieurs et antéro-laté-

raux. Nous pouvons résumer cette répartition par

86-88, 32-34, 7, 8-9, 9. 9 et 14-16. Chez six

individus de H. striata, le nombre de sinuosités

est comparable. En utilisant la même loimule

nous avons compte 86-88, 30, 7, 8, 8, 7-9 et 14-

15 sinuosités. Chez H. forci fera, outre l'absence

des crêtes loculaires, le nombre de sinuosités sur

les differents septa est plus faible. Pour le hapteur

de quatre individus, nous avons compté 72, 25,

7, 6, 7, 5, 5 sinuosités. On peut penser que,

entre cette espèce et les précédentes, cette diffé¬

rence entre le nombre de sinuosités résulte de la

taille du hapteur. fin effet, son diamètre,

(290 uni) chez H. forcifcra, esr nettement infé¬

rieur -à celui du hapteur de H, mokhtarae

(515 pm) et de H. striata (498 pm). Mais le

nombre de sinuosités paraît lié à l’espèce. Ainsi

chez H arma ta, dont le hapteur a la même mor¬

phologie et la même taille que celui de H. fbreife-

ru, ou peut compter sur la figure de Tîmofeeva

(1983) un nombre légèrement différent de sinuo¬

sités, soit : 79, 25, 6, 6, 7, 9, 10.

Les deux cavités observées dans le bulbe éjacula-

tcur de H. mokhtarae er H striata peuvent être

rapprochées îles deux chambres internes du

bulbe éjaculateur des Calieotylinae, c’est-à-dire

une des apomorphie.s .soutenant le caractère

mo-nophylétique de cette sous-famille. Il n’est pas

actuellement possible de proposer une homologie

entre ces formations, homologie qui modifierait

profondément la systématique proposée par

ZOOSYSTEMA • 1999 • 21 (2)

167

Neifar L., Euzet L. & Ben Hassine O. K.

Chisholm et al. (1995) pour les Monocotylidae.

Avec la description de H. caprieornensis ,

Chisholm & Whittington (1996) font une révi¬

sion du genre Heterocotyle. Ils reconnaissent 10

espèces. Deux autres, H. ellîptica Pillai & Pillai,

1976 de Pastinacbus sepheu (Porsskâl, 1775) et

H. robuste! (Johnston & Tiegs, 1922) de

Üroloplnts testaceus (Trygonopiem tcstacea Millier

8sc Henle, 1841), sont considérées comme sp.

inquirenda leur description étant insuffisante.

Neifar cl al. (1998) onr récemment décrit Hete-

rocotyle scoiti et H. similis , deux nouvelles espèces

parasites de D. pastinaca. Avec les trois parasites

de Taeniura grabata , le genre compte actuelle¬

ment 15 espèces, toutes parasites de Dasyatidae.

Ce sont : Heterocotyle americana Hargis, 1955 de

Dasyatis americana Hildebrand & Schroeder,

1928 ; H. arrnata Timofeeva, 1983 de Hitnati-

tura uârtuik (Forsskâl, 1775) ; H. caprieornensis

Chisholm & Whittington, 1996 de Himantura

fai Jordan & Seule, 1906 ; H. chinensis Timofeeva,

1983 de D. akajei (Müller &C Henle, 1841) ;

H. confusa Timofeeva, 1983 de Himantura uar-

nak ; H. dasyatis (Yamaguti, 1965) de Dasyatis sp.;

H. forciferd n. sp. de Taeniura grabata (Geoffroy

Saint Hilaire, 1817) ; H. granulalae Young, 1967

de Himantura granulata (Maclcay, 1 883) ;

H. rninirna (Mac Callum, 1916) de D. ccntrourd ?

(Mitchill, 1815) ; H. mokhtdrde n. sp. de T. graba¬

ta-, H. pastmacae Scott, 1904 de D. pastinaca ;

H. pseudominima Hargis, 1955 de IX americana

ou D. say (Lesueur, I81 T ) ; H. scotti Neilar, Euzet

& Ben Hassine, 1998 de D. pastinaca ; H. similis

Neifar, Euzet &C Ben Hassine, 1998 de D. pastina¬

ca ; H. striata n. sp. de T. grabata.

Déjà signalé par Lawler (1981). et discuté par

Chisholm &C Whittington (1996), l’hôte de

H. minima reste incertain. Après Price (1938)

nous citons D. centroura , mais avec doute, car les

monogènes de cette pastenague de la côte atlan¬

tique américaine n'ont pas été étudiés, Il existe

aussi une incertitude pour l'hôte de H. pseudomi¬

nima Hargis, 1955, Hargis donnant D. america¬

na ou D. say. Notons que nous n’avons pas

observé d’individus correspondant à H. pseudo-

mtntma parmi les parasites branchiaux d'un spé¬

cimen de D. amencana où nous avons par contre

trouvé plusieurs individus de H. americana.

Clé de détermination du genre Hetekocotyle

1. Appareil copulateur mâle avec une pièce accessoire .2

— Appareil copulateur mâle sans pièce accessoire. 5

2. Pièce accessoire simple .3

— Pièce accessoire complexe ..4

3. Pièce accessoire large à extrémité denticulée . H. armata

— Pièce accessoire en griffe avec un bouton sur la convexité . H. americana

4. Pièce accessoire à deux branches. H. forcifera

— Pièce accessoire formée par un faisceau de baguettes . H. dasyatis

5. Pénis large conique .. 6

— Pénis étroit tubulaire...7

6. Pénis à trois pointes distales. Paroi vaginale épineuse . H. pastinacae

— Pénis avec deux rangées de filaments. Vagin inerme .. H. striata

7. Pénis rectiligne...... 8

— Pénis fortement arqué . 1 1

168

ZOOSYSTEMA • 1999 • 21 (2)

Heterocotyle nouveaux de Taeniura grabatc

8. Longueur du pénis supérieure à 130 pm ...*...9

— Longueur du pénis inférieure à 130 pm. 10

9. Bulbe éjaculateur aussi long que le pénis . .. H. confusa

— Bulbe éjaculateur deux fois pius long que le pénis ... H. mokhtarae

10. Longueur du pénis voisinant 100 pm . H. capricornensis

— Longueur du pénis inférieure à 100 pm . H. chinensis

11. Pénis arqué, taille inférieure à 100 pm. H. pseudominima

— Pénis arqué, caille supérieure ou égale à 100 pm . 12

12. Taille de pénis supérieure à 100 pm... H. scotti

— Taille du pénis environ 100 pm... 13

13. Membrane marginale large . H. granulatae

— Membrane marginale étroite . 14

14. Pénis à extrémité distale évasée. H. minima

— Pénis à extrémité distale repliée. H. similis

Remerciements

Nous remercions R. Overstreet et S. Curran qui

ont récolté et envoyé les branchies de Dasyatis

americaria parasitées par Heterocotyle americaria.

Ce matériel nous a permis de préciser la morpho¬

logie de l'appareil copulateur de ce Monocoty-

lidae. Nous remercions Mr. O. Zoghlami

Directeur général des Écoles de pèche de Tunisie

et Mrs. Y. Omrani et W. Boughdir, respective¬

ment Directeurs de l'Lcole de Pêche de Bizerte et

de Zarzis, pour nous avoir très aimablement

accueillis dans leur établissement et facilité nos

récoltes.

RÉFÉRENCES

Chisholm L. A. 1995. — Taxonomy of Heterocotyle

dasyatis n. comb. (syn. Diploheterocotylit dasyatis)

(Monogenea: Monocorylidae). Journal of Parasito-

logy 81: 439-441.

Chisholm I.. A.. WheelerT. A. & Bevcrlcy Burron M.

1995. — A phylogent - tic analysis and revised classi¬

fication of the Monocorylidae Taschenberg, 1879

(Monogenea), System,)tir Paras itôa/gy 32; 159-191,

Chisholm L. A. & Whittington 1. D. 1996. — A révi¬

sion of Heterocotyle (Monogenea: Monocorylidae) with

a description of Heterocotyle capricornensis n. sp. front

Himantura fai (Dasyatididae) front Héron Island,

Créât Barricr Réel , Âustralia, International Journal

for Parashology 26: 1169-1190,

Lawler A. R. 1981. — Zoogeogrnphy and Host-specifi -

city of the Superjamily Capsalnidea Price, 1936

(Monogenea: Mounpisthocotylea). An Evaluation of

the Host-Parusite Records of the Superfomily Capsu¬

lai dect Price. 1936. and their (Jtility in Détermi¬

nations of Hosi-specifiaty and Zoogeogrnphy. Spécial

Papers in Marine Science. Numbcr 6. Virginia

Instkure of Marine Science and School of Marine

Science, College of William and Mare, Virginia,

650 p.

Malmberg G, 1957. — [On the occurrence ol

Gyroddctylus on Swedisb ftshes]. Skrijierutgivna av

Sodra Soeriges Fiskeriforentng ( 1956): 19-~6 (in

Svvedish, with description of species and a summary

in Hnglish).

McEàchran J. D. & Capapé C. 1984. — Dusyand.te:

19 T -202, in Whirehead P. J. P. étal, (eds), Poissons

de l'Atlantique du Nord-Est et de la Méditerranée ,

volume L UNESCO, Paris.

Neifar L.. Emet L. & Ben Hassitte O. K. 1998.

Nouveaux Monocorylidae (Monogenea), parasites

branchiaux de Dasyatis pastinaeü (Linné)

(Euselachii. Dasyatidae). Compléments à la des¬

cription de Heterocotyle pauinacac Scott, 1904-

Systematic Parasitology 41: 197-208.

ZOOSYSTEMA • 1999 • 21 (2)

169

Neifar L., Euzet L. & Ben Hassine O. K.

Pariselle A. & Euzet L. 1995. — Gill parasites of the

genus Cichlidogyrus Paperna, 1960 (Monogenea,

Ancyrocephalidae) from Tilapia guineensis (Bleeker,

1862), with description of six new species.

Systematic Parasitology 30: 187-198.

Timofeeva T. A. 1983. — [New représentatives of

monocotylids (Monogenea, Monocotylidae) from

cartilaginous fishes of the South-China and Yellow

Seas]. Proceedings of the Zoological Institute 121: 35-

47 (en russe).

Soumis le 17 avril 1998 ;

accepté le 25 septembre 1998.

170

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of the Cochlespirinae

(Gastropoda, Conoidea, Turridae)

Alexandra I. MEDINSKAYA

A. N. Severtzov Institute of Problems of Evolution,

Leninsky Prospect 33, Moscow 117071 (Russia)

Medinskaya A- I- 1999. — Forégul anatomy of the Cochlespirinae (Gastropoda, Conoidea,

Turridae). Zoosystema21 (2) : 171-198.

KEYWORDS

Cochlespirinae,

Conoidea,

anatomy,

foregut,

histology.

ABSTRACT

The foregut anatomy of 20 species, belonging to eight généra, of the sub-

fâmily Cochlespirinae is described. A cladistic analysis based on several most

important characters (morphology of probcwcis, position of buccal sphinc¬

ters, histology of venom gland, position o( the venom gland opcning, struc¬

ture of muscular bulb, and morphology' of radular tecth) revealed three more

or less well-defmed groupe within the subfamily. The main fearure characte-

rizing the subfamily as a wholt and separating groups within it, appeared to

be the structure of venom gland and ifs muscular bulb. The stlbgenus

Sibogasyrinx of the genus Lettcosyrinx was shown to deserve a genus status.

Some généra appeared to be intermediate between Cochlespirinae and

Crassispitinae in some anatomical characters, and their taxonomie position

remains not completely clear.

MOTS CLÉS

Cochlespirinae,

Conoidea,

anatomie,

système digestif,

histologie.

RÉSUMÉ

L'anatomie du système digestif des Cochlespirinae (Gastropoda, Conoidea ,

Turridae).

1. anatomie du système digestif de 20 espèces, appartenant à huit genres de la

sous-famille Cochlespirinae, est étudiée. Une analyse cladisrique, fondée sur

les plus importants caractères de ce groupe (la morphologie de la trompe, la

disposkion des sphincters, l'histologie de la glande à venin, la disposition de

l’ouverture de la glande à venin, la structure de la poire musculaire et la mor¬

phologie des dents de la radula) a permis de distinguer trois groupes plus ou

moins homogènes. Le caractère principal, qui permet de caractériser la sous-

famille et de la diviser en différents groupes morphologiques, est la structure

de la glande à venin et de sa poire musculaire. Le sous-genre Sibogasyrinx du

genre Lettcosyrinx mérite le statut de genre. Quelques genres semblent occu¬

per une position intermédiaire entre les deux sous-familles Crassispirinae et

Cochlespirinae, et leur position taxonomique n'est pas encore définitivement

établie.

ZOOSYSTEMA • 1999 • 21 (2)

171

Medinskaya A. I.

INTRODUCTION

The superfamily Conoidea is well-defined mor-

phologically due to che presence of venom gland

and highly modifted radular teeth, At the same

dme the relationships vvithin the superfamily are

still far frnm being clear, and this is one ol the

main problems of the current taxonomy of the

group.

Investigations of phylogenetic relations within

Conoidea and the classification of the super¬

family were mainly based on characters of shell

and radula (e.g., Powcll 1966; McLean 1971),

and only recent studies hâve paid much attention

to the anatomy of digestive System, which is the

most variable and, therefore, the most informa¬

tive division.

The first compréhensive analysis of conoidean

anatomy together with radula and shell morpho-

logy was published in 1993 (Taylor et al. 1993).

This demonstrated the importance of anatomical

characters for taxonomie and phylogenetic analy¬

sis of the group. Although this study was based

on the anatomy of 72 species, this is still only a

small portion of living species of the superfamily

(apparently no less titan 5000 species).

In anatomical studies of Conoidea, the very

diverse fantily Turridae is of a spécial interest.

The fantily consiste of tour currently recognized

subfamilies, viz i’urrinac, Clavatulinae, Crassis-

pirinae, and Cochlespirinae. and is characccrized

by a radula with a membrane and marginal teeth

usually of wishltone type (Taylor et al. 1993).

The taxonomie positron of nvany représentatives

of this fantily and rheir placement in the sub-

families are still eonrroversial.

Anatomical study of tlie subfamily Crassispirinae

(Kantor et al. 1997) lias sltown many inreresting

features and lias allowed a clearer définition of

the boundaries of the subfamily. I bis Work lias

made évident the necessity ofsimil.tr studies

dévoted to other subfamilies, for lire complété

analysis of relationships among Turridae and

more reliable séparation of Subfamilies.

Therefore, the aim of the présent study was to

analyzc the anatomy of species of the subfantily

Cochlespirinae and to teveal the rporphological

diversity of its représentatives with the objective

of establishing relationships of this subfamily

with other taxa of Turridae.

MATER1AL AND METHODS

The subfamily Cochlespirinae is considered to

include 22 Recent généra (Taylor et al. 1993).

The présent study covers eight of them, as well as

two généra {Kurilobadaha Sysoev & Kantor,

1986 and t-'Ucisyrinx Sysoev & Kantor, 1986),

which were prcviously assigned to Crassispirinae

(Taylor et al. 1993). Elle two lauer généra can-

not bc actually ine 1 udcd in Crassispirinae

(Kantor et al. 1997), and therefore thev are pre-

sently considered in Cochlespirinae, within

which they were originally descrihed (Sysoev &

Kantor 1986).

The study of anatomy of Cochlespirinae was

based on 20 species from tlie Pacific and the

Atlantic, collected from depths of 100 to 7000 m

(Table 1).

Plie anatomy was studied from longitudinal

serial sections of the foregut, eut at 10 ( 1 m and

stained in Mason’s triple stain. The radula was

examined by SF.M. lu some cases, the results of

liand dissections were also used.

The character analysis was performed using

PAUP version 3.1.1 (Swofford 1993).

ABBREV1ATIONS USKll IN THE FIGURES

bc buccal cavity

bip buccal lips

bm buccal mass

lit buccal tube

btsa amerior sphincter of buccal tube

btss shifting sphincter of buccal tube

cm colummellar muscle

con circumoesophageal nerve ring

ctl connective fissile layer of muscular bulb

ebt sac-like en large me ru of buccal tube

ech rhynchodcum epithelium change

eoe oesophagus cnlargcmcnt

ep epitnelial pad

gre glandular part of rhynchodeum

mb muscular bulb

oe oesophagus

ph prohoscis

pw proboscis wall

rhe rhynchocoel

rhs rhynchostomal sphincter

rhw rhynchodeum wall

172

ZOOSYSTEMA • 1999 • 21 (2)

Table 1. — List of cochlespirine species. which hâve been seclioned. with details of lheir collection location.

The additional material included sections of Megasuraila carpenteriana (Gabb, 1865), Aforla Inoperculata Sysoev & Kantor, 1988, A circinata (Dali, 1873), A. lepta (Watson, 1881),

A. moskaievi Sysoev & Kantor, 1987, A aulaca alaskana Sysoev & Kantor, 1987, A crebristriata (Dali, 1908). A, abyssalis Sysoev & Kantor, 1987, A. kupriyanovi Sysoev & Kantor,

1987, kindly provided to the authorby John Taylor (the first species) and Yuri Kantor (ail other species).

Species

Campaign

Station

Depth

Location

Coordinates

Antiplanes sanctiioannis

(Smith, 1875).

R/V Gidronavt

R/V Gidrobiolog

dredge No. 41

stn 124

110 m

550 m

Paramushir ld.,

Pacific side

50°15’N, 156°29'E

48°15'N, 140°41'E

Carinolurrispolycaste (Dali, 1919)

R/V Vityaz

stn 4179

1258 m

North California

42°40' N, 125°00' W

Cochlespira pulchella

(Schepman, 1913)

R/V Baruna Jaya, KARUBAR

stn CP 59

405-399 m

Indonesia, Tanimbar Is.

08°20'S, 132°11'E

Cochlespira radiata (Dali, 1889).

R/V Marion Dufresne, MD55

stn CP 11

248-262 m

SE Brazil

23°35 S, 40°06'W

Comitas murrawolga

(Garrard, 1961)

R/V Vauban,

MUSORSTOM 4

stn DW 229

445-460 m

Southern New Caledonia

22°52'S, 167°13’E

Co mitas onokeana vive ns

Dell, 1956

R/V Dmitry Mendeleev

stn 1269,

Sigsbee trawl

720 m

14°26'S, 174°29’W

Kurilohadalia elongata

Sysoev & Kantor. 1986.

Zoogical Muséum of Moscow

State University Lc-22398, paratype

R/V Vityaz

stn 2208

7210-7230 m

Kurile-Kamchatka

trench, east of North

Kurile Is.

49°29‘N, 158°41’E

Leucosyrinx tSibogasyrinx)

pyramidalis (Schepman, 1913)

R/V Baruna Jaya, KARUBAR

stn CP 72

699-676 m

Indonesia, Tanimbar Is.

08°36’S, 131°33’E

Leucosyrinx verrilll (Dali, 1881)

R/V Akademik Kurchatov,

14th cruise

stn 1209

1060 m

NE Atlantic

13°04'N, 63°06'W

Marshallena plvlipplnarum

(Watson, 1882)

R/V Coriotis. BIOGEOCAL

stn CP 260

1820-1980 m

Loyalty Basin

21°00'S, 166°58'E

Plicisyrinx decapitata

Sysoev & Kantor, 1986.

Zoogical Muséum of Moscow

State University Lc-22388, paratype

R/V Vityaz

stn 3166

5557 m

E South Kurile Is.

44°42,09'N, 153°49’E

Foregur anatomy of Cochlespirii

Medinskaya A. I.

rndular sac

rso

radular sac opening

rst

rhynchostome

s g

salivary gland

sgd

sallvary gland duct

proboscis scalk

v g

venom gland

vgd

venom gland duct

HISTORICAL REVIEW OF

COCHLESP1R1NAE AND THE PRESENT

CONCEPT Oh TH E SUBFAMILY

The subfamily Cochlespirinae with the type

genus Cothlespim Conrad, 1865 was established

by Powell (1942) for a group ol fbssîl and Recenr

généra diaractemed hy a “chtn tusiform shell,

with sliarp, sornetimes frilled, penpheral keel"

(Powell 1942: 31), Subsequently Powell (1966)

has synonymized Cochlespirinae with Turrieuli-

nae Powell, 1942 becati.se ofsLigbt différences

between the subfamilies. According to Powell’s

(1966, 1969) concept of Turriculinae, the sub-

family was rather heterogeneous and included

généra with different styles of radula, but united

by moderato to large shell size, more or less nar-

rowly fusiform shape, the sinus position on the

shoulder, a rail spire, and a long canal.

McLean (1971) generally employée! Powell's

concept of Turriculinae, but restricted the sub-

family to généra with a uniform type of radula,

consisring ol wishhone marginal teeih and a

variouslv devçloped (to absent) central toorh.

In the révision of Conoidea based on anatontical

characiers (Taylor et al. 1993), the authors found

that the type genus of the subfamily, Turricula

Schumacher, 1817. is vety dose anatomically to

représentatives of the subfamily Clavatulinac

Gray, 1853. Thercfore, Turricula was transferred

to Clavatulinae and Turriculinae became a syno-

nym of the latter. This made the name

Cochlespirinae valid for the group consisting of

the resl of Turriculinae sensu Powell. The dia-

gnosis ol Cocblespirinae in Taylor et al. mainly

followed that of earlier authors and was mainly

based on conchological characters, because ana-

tomical data did not givc suffîcient grounds for

the clear séparation of Cochlespirinae from, for

example, Crassispirinae McLean, 1971. There-

fore, 22 Recent généra (and 6 subgenera) recog-

nized by Taylor et al (1993) in Cochlespirinae,

Were characrerized by the following set of

conchological fearures: shell of medium to large

size, fusiform or pagodiform, with usually ntode-

rately elongate canal, generally deep anal sinus

on the shoulder, and usually multispiral and

smooth (at least initially) proroconch.

ANATOMY

I his section describes anatomy of the anterior

part of the digestive System and radula of each

species. Unfortunately, many species were repre-

sented by a single specimen, therefore the radula

descriptions are given only For some species.

Besides the original material, we usecl the data on

the anatomy of Aforia spp, (Sysoev & Kantor

1987, 1988), with some additions and correc¬

tions based on the study of original préparations.

Detailed descriptions of these species are given in

the respective papers, and some corrections are

listed in Table 3.

Cochlespira pu le bel la (Schepman, 1913)

(Figs I ; 16A, B)

As the anatomy of C. radiata (Dali, 1889)

(Fig. 16C) differs only slightly from that of

C pulchelta . the description mostly concerns

both species, with the différences being .specially

indicated.

T he rhynchodeum is narrow and long. The

rhvnchodeal sphincter is large, in posterîor posi¬

tion. Lhe change of glandtilar epithelium oçcurs

in the hrst third of the rhynchodeum length in

inverted position. The entire part of rhyncho¬

deum covered by epithelium similar to that of

proboscis, is evcriible.

l he proboscis is long, with a rather large basal

diameter and nartowing apically. At the apical

end of proboscis there are rwo anterior sphincters

with a sac-like buccal enlargement between them

and a medium-sized épithélial pad. The buccal

tube is almost straight in the apical part, and

with regular small folds in the basal part. The

lumen of the buccal tube is rather narrow and

174

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

Fig. 1. — Cochlespira pulchella , reconstruction based on longitudinal sections through the foregut.

expands greatly in front of the buccal mass.

Proboscis and buccal tube walls are rather mus-

cular, at the base the proboscis vvall thickness is

about 19% of the total proboscis diameter, the

buccal tube wall thickness is about 6%.

The buccal mass is large, muscular, and situated

posterior to the proboscis. Buccal lips are rather

long, muscular, and invertcd, The buccal mass

forms a caecum-like structure with the opening

of radular sac. In C. radiata it is better develop-

ed. The radular sac has a wide opening, and the

buccal sac is absent.

Ducts of salivary glands are long and open on

both sides of radular sac. Salivary glands are very

large and acinous. The vcnom gland opens under

the nerve ring and behind the buccal mass. The

histology of the venom gland does not change

posterior to the rterve ring. The muscular bulb is

very large (srnall in C. r/tdiata), comprised by

two oppo.sitely oriented layers of muscles, and

without a connective ti.ssue laycr. The lumen of

the muscular bulb is rather narrow, The ocsopha-

gcal loop is very long. The buccal mass is situa-

ted antcriorly, far front the nerve ring. The

oesophagus diameter does not change markedly

behind the nerve ring.

The anatomy of Cochlespira radiata and C. pul-

chella differs only in srnall details. The radula of

both species has a well-formed central tooth with

a strong central cusp. Marginal teeth of C. pid-

cbelltt (Fig. 16A, B) are of the wishbone type

with two limbs. Teeth of G radiata (Fig. 16C)

appear to he longitudinally folded and somewhat

twisted, with a central channel.

Leucosyrinx verrilli (Dali, 1881)

(Fig. 2)

The large rhynchodeum sphincter lies in an ante-

rior position and practieally encircies the rhyn-

chostome. The epithelium of the rhynchndeal

wall is glandular and changes histology in the

posterior quarter of the rhynchodeum length.

The posterior part of the rhynchodeum is lined

by an epithelium similar to that of proboscis, It

Ls evertible.

The proboscis is of medium length, with a rather

large diameter, widi the apical end loosely coiled

inside the rhynchodeum. At the tip of proboscis,

there arc two anterior sphincters and a sac-like

buccal enlargement between them. A well-deve-

loped épithélial pad is situated in front of the

ZOOSYSTEMA • 1999 • 21 (2)

175

Medinskaya A. I.

Fig. 2. — Leucosyrinx verrilli, reconstruction based on longitudinal sections through the foregut.

first sphincter. The buccal tube is straight.

Proboscis walls are not very muscular and uni¬

fia rmly thin along their entire length. The wall

thickness is about 7% of the total proboscis dia-

nteter and the buccal tube wall thickness is about

6 %.

The buccal mass is medium-sized, muscular and

situated postcrior to the proboscis. Buccal lips

are absent. The radular sac opening is rather nar-

row and the buccal sac is absent. Ducts of the

salivary glands are rather short, weakly coiled and

open on both sides of radular sac. Salivary glands

are very sinall and acinous. The venom gland

opens at the boundary ol buccal mass by a smaJI

duct. Venom gland histology is unchanged post-

eriorly. The muscular bulb is coniposed of rvvo

layers of longitudinal muscles separated by a

layer of unstructured connective tissue, with an

innerniost layer of circulât fibres. The muscular

bulb lumen is rather narrow, The oesophagus is

greatly looped. The oesophagus diameter does

not markedly increase behind the nerve ring.

Data on radula are absent.

Leucosyrinx (Sibogasyrinx) pyramidalis

(Schepinan, 1913)

(Figs 3; 16D, E)

The large rhynchodeal sphincter is anterior and

almost encircles rhe rliynchosrome. The epithe¬

lium of the rhynchodeal wall is glandular,

changes histology in the rear quarter of the rhyn-

chodcum Icngrh. The postcrior part of the rhyn-

chodeum is lined by an epithelium similar to

that of proboscis. It is cverliblc.

The proboscis is long, with rather a large basal

diameter; the apical end is very narrow, rolled up

into a bail, At the proboscis tip there are two

anterior sphincters and a sac-like buccal enlarge-

ment berween them. A distinct épithélial pad is

absent. The buccal tube is straight apically and

lolded basally, with a narrow lumen. It greatly

expands in front of the buccal mass. Proboscis

and buccal mass walls are weakly muscular and

thin. In the apical part the wall thickness is

slightly more than 10% of the total proboscis

diameter, whereas the buccal tube wall thickness

176

ZOOSYSTEMA • 1999 ■ 21 (2)

Foregut anatomy of Cochlespirinae

ebt

Fig. 3. — Leucosyrinx ( Sibogasyrinx ) pyramidalis, reconstruction based on longitudinal sections through the foregut.

is only about 4%. The proboscis diameter greatly

increases basally, with a corresponding increase in

the wall tbickness, whereas the buccal tube walls

remain thin-walled, and rheir thickness becomes

only about 2-3% of the total proboscis diameter.

The buccal mass is large, wcaldy muscular, cur-

ved and situated behîng the proboscis. Buccal

mass walls are thin, with hardly differentiated

end of buccal mass, because the hisrology of the

oesophageal wall does not differ front that of

buccal mass. The différence consists only in the

relative thickness of muscle layers and walls

themselves. Buccal lips are long, inverted and

rather muscular.

The radular sac has a wide opening and lacks a

buccal sac. Duces of salivary glands are long and

open on both sides of radular sac. Salivary glands

are medium-sizcd and acinous. The venom gland

opens into the oesophagus beyond the buccal

mass, behtnd the nerve ring (therefore, it does

not pass through the nerve ring). The ducr is

absent. Venom gland histology does not change.

The muscular bulb is composée! of two opposing

layers of muscles, without a connective tissue

layer. The lumen of muscular bulb is rather nar-

row.Thc oesophagus is moderately elongated.

The buccal mass is situated not far anteriorly of

the nerve ring. The oesophagus diameter greatly

increases behind the nerve ring.

The radula (Fig. 16D, E) is very similar ro that

of Cochlespira pulcbetla , having wishbone margin¬

al teeth and a well-deftned central tooth with a

strong cusp.

Comitas onokeana vivens Dell, 1956

(Figs 4; 5A, B; 17A)

The rhynchodeal sphincter is small, anterior and

situated slîghtly behind the rhynchostome. The

rhynchodeum wall epithelium is glandular, dif-

fering front that of proboscis wall. The rhyncho¬

deum is not evertible.

The proboscis is long, wide, straight and covered

by regular small folds along the enrire length.

The structure of the proboscis tip is very pecti-

liar. The muscular wall of proboscis becontes

ahrupdy veiy thin and simulraneously the extern-

al epithelium changes from high columnar ro

very low. No distinct change of epithelium

occurs at the apical end of proboscis. The exter-

nal epithelium is the saine as the internai buccal

tube epithelium.

In the two sectioned specimens (Fig. 5A, B), the

extent of eversion is different. In the place where

ZOOSYSTEMA • 1999 • 21 (2)

177

Medinskaya A. I.

Fig. 4. — Comitas onokeana vivens, reconstruction based on longitudinal sections through the foregut.

the buccal tube becomes more muscular, there

are two small sphincters on both sides of buccal

sac-like enlargement. There are two glandular

pads in the enlargement.

Morphologically, the structure of this portion

corresponds to that ot apical end of the proboscis

with two anterior sphincters, le is seen (Fig. 5B)

that the buccal tube may evert to such extent

that this portion appears practically at the very

end of the proboscis.

The buccal tube has small dcnscly arranged folds

along its enttre length. Proboscis walls are not

very muscular, with the same thickness along

their entire length. 1 he wall thickness is about 6-

7% of the total proboscis diameter, the buccal

tube wall thickness about 10%. The buccal tube

is lined with rather low columnar epithelium,

and becomes larger after beginning of buccal

mass. The oesophagus is lined with columnar

epithelium. The buccal mass is rather large, mus¬

cular and situated behind the proboscis. Buccal

lips are absent. The oesophageal wall torms not

muscular, rather long but thin lips in front of the

buccal mass.

The radular sac has a wide opening, whilst the

buccal sac is absent. Ducts of the salivary glands

are rather long and coiled, and open on both

sides of radular sac. Salivary glands are small and

acinous. The venorn gland opens ncar the radu¬

lar sac, ar rhe posterior part of the huccal mass.

The duel is absent. Venom gland histology does

nul change. The muscular bulb is composed of

two lâyers of longitudinal muscles separated by a

layer of unscructured connective tissue. 1 he

outer layer thickness is three to four limes more

than that of inner layer. The lumen of the mus¬

cular bulb is rather large. The oesophagus is

moderacely elongated. The oesophagus diameter

does not increase behind the nerve ring.

The radula (Fig. 17A) consisrs of fiat marginal

178

ZOOSYSTEMA • 1999 ■ 21 (2)

Foregut anatomy of Cochlespirinae

btss

Fig. 5. — A, B, Comitas onokeana vivens, reconstruction based on longitudinal sections through the apical part of the proboscis, dif¬

ferent extent of the eversion.

teeth thickened on the outer sides, with two flat C. murriuvolga has also a distinct central tooth

plates in the central part. instead of two fiat plates.

Comitas murrnwolga (Garrard, 1961) Antiplanes sanctiioannis (Smith, 1875)

(Fig. 17B.C) (Fig. 6A, B)

The anatomy of Comitas murrdwolga and C. ono- The rhynchodeal sphincter is anterior. The epi-

keana vivons differs onlv in minor details, thelium of the rhynchodeal wall is glandular and

C. murrnwolga has very small buccal Jips inverted differs front that of proboscis wall. The proboscis

into the buccal cavity. Marginal teeth are very is long, wide, straight and covered by regular

similar to those of C. onokeana vivens , but small folds along its entire length. The structure

ZOOSYSTEMA • 1999 • 21 (2)

Medinskaya A. I.

Fig. 6. — Antiplanes sanctiioanms. A reconstruction based on longitudinal sections through the foregut; B, apical part of proboscis.

of the proboscis tip is similar to that of Comitas. buccal en largement with a long and tall épithélial

The rather muscular wall of the proboscis pad. The situation is thus similar to that in

becomes abruptly very thin, but, in contrast to Comitas , where the structure of fhis part corres-

Camitas, without a change in outer epithelium, ponds morphologically to that of the proboscis

The anterior end of the proboscis is truncated tip, whereas this part is intermediate in its posi-

apically, with a rather long and narrow stalk near tion. The buccal tube has small densely arranged

the centtr of llatiencd area. No distinct change folds along its entire length.

of epithelium occurs at the tip of the proboscis. The proboscis waJis are rather muscular, with the

In the two secrioned specimens, the extent of same thickness alung their entire length. The wall

eversion is different. In the second specimen (not thickness is about 10-12% of the total proboscis

figured) the thin-walled portion is considerably diaraeter, whilsr the buccal tube wall thickness is

shorter, and the buccal mass is situated behind about 7-8%. The buccal tube is lincd with a

the proboscis. rather low columnar epithelium, bccoming taller

At the beginning of the wide portion of rhe pro- after rhe beginning of buccal mass. The oesopha-

boscis there are a small sphincter and a sac-like gus is lined with columnar epithelium.

180

ZOOSYSTEMA • 1999 - 21 (2)

Foregut anatomy of Cochlespirinae

Fig. 7. — Marshallena pnilippinarum , reconstruction based on longitudinal sections through the foregut.

The buccal mass is large, muscular, folded and

situated at the proboscis base. Buccal lips are

long, thin. rather muscular and nor everted. The

radular sac has a wide opening, wTiilst the buccal

sac is absent. Salivary ducts arc long, coiled, open

on both sidc.s of radular sac. Salivary glands are

large and acinous. 1 he venom gland opens near

the radular sac, at the rear boundary of buccal

mass. A duct is absent. Venom gland histology

does not change. The muscular bulb is compo-

sed of two layers of longitudinal muscles separa-

ted by a laver ol unstructured connective tissue.

The thickness of the outer laver is slightly less

than that of the inner. The lumen of the muscu-

lar bulb is small. The oesophageal loop is short.

Oesophagus diameter greatly increascs behind

the nerve ring.

The radula consists of two marginal teeth, com-

posed of two Hat plates.

Marshallena philippinarum (Watson, 1882)

(Figs 7; 17D, E)

The rhynchodeal sphincter is located in an ante-

rior position. The epithelium of the rhynchodeal

wall is glandular, but is developed only in the

anterior part of rhynchodeum. The rhyncho-

deum walls are very thin in the basal part, and

the glandular epithelium is represented only by

scattered cells. The rhynchodeum is not ever-

tible.

The proboscis is long, straight, except the apical

part, and of a small diameter which does not

change along the proboscis length. The structure

of the proboscis tip is similar to that of

AmipLmes. The muscular wall of proboscis gia-

dually becomes very thin withour a change in the

external epithelium, lire proboscis walls are verv

thin in the apical part. At the beginning of the

wide portion of the proboscis there is a large

sphincter, whcreas a sac-like buccal enlargement

and épithélial pad arc absent. The buccal tube is

rather srrongly coiled inside chc proboscis.

Proboscis walls are muscular in the basal pair.

The wall thickness is about 19% of the total pro¬

boscis diamc-ter, with rhe buccal tube wall thick¬

ness about 3-4%.

The buccal mass is vety large and muscular. le is

situated almost perpendicular to the proboscis

and benr so that its end lies almost at the probos-

ZOOSYSTEMA • 1999 • 21 (2)

181

MedinskayaA. I.

Fig. 8. — Kurilohadalia elongata, reconstruction based on longitudinal sections through the foregut.

cis base. In Figure 7 the buccal mass is shown at

a slightly reduced scale. The nerve ring lies

almost immediâtely behind the buccal mass, near

the proboscis base. Buccal lips are rather short

but muscular, and not everted. The radular sac is

very short, with a narrow opening, the buccal sac

is absent.

The salivary ducts are long, slightly coiled and

open on hoth sides of radular sac. The salivary

glands are large and acinous. (The glands and

ducts are not shown in Figure 7). The venom

gland opeits iar behind the radular sac, at the

rear boundary of the buccal mass. The duct is

absent. The histology of the venom gland is very

different Iront usual. A hollow, uncoiled tube

arises from the buccal mass, goes strietly along

the oesophagus and passes through the nerve

ring. It is lined with a glanduiar epithelium with

very large tells, the venom granules arc absent.

The histology ol the venom gland changes just in

front of the muscular bulb. l’he epithelium

abruptly disappears, and the tube ht contes ftlled

by venom granules. The granules are présent also

in the lumen of the muscular hulb. The muscu-

lar bulb is contposed of two layers ol longitudi¬

nal muscles separnted by a layer of unstructured

connective tisxue. The thickness ot the outer

layer is slightly less than titat ol the inner. The

lumen of muscular bulb is sntall. The oesopha¬

gus is slightly clongated. Oesophagus dianieter

does not increase markedly behind the nerve

ring, but rite oesophagus for ms an appendix,

extending berween the rltynchodcum wall and

the bodv wall. The wall of oesophagus is almost

fused with the thin wall of rhynchodeum. The

epithelium of the oesophagus consiste of very

large cells, with a number of regularly arranged,

large, glanduiar cells.

The radula (Fig. l^D, E) consist only of rwo

marginal teetlt in each row. The teetlt are wish-

hone and composed of two nearly equal limbs.

Ktirilobadalia elonvata Sysoev & Kantor, 1986

(Fig. 8)

The rhynchodeum sphincter is in the anterior

182

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

vgd sg

Fig. 9. — Plicisyrinx decapitata, reconstruction based on longitudinal sections through the foregut.

position but shifted backward. A glandular epi¬

thelium is présent.

The proboscis is rather long, uncoiled and occu-

pies mosr of the rhynchodeum. The proboscis

diameter does not change along its length, The

buccal tube is very narrow from the mouih to

the First sphincter. At the tip ot the proboscis

there are twn anterior sphincters, the First one

being smaller that the second, There is a sac-like

buccal enlargement between the sphincters. lt is

lined with a tall epithelium, ditlering from the

resr of the buccal tube epithelium. Buccal tube

walls are very thin and rather strongly folded

inside the proboscis.

The proboscis diameter is very large, but the pro¬

boscis walls are not very muscufar, the wall thick-

ness being about 7% ol the total diameter. The

proboscis mostly consists of longitudinal muscles.

The buccal mass is large, situated behind the

proboscis. Buccal lips are long and very niuscu-

lar, whilst the walls of the buccal mass are weakly

muscular. In the radular sac opening there are

thin and long lips. Salivary ducts are very short,

straight and open into a deep buccal sac. Salivarv

glands arc large, with an anastomosing-tubular

morphology. The vehom gland opens near the

radular sac by a ver)' short ducr.

Venom gland histology does not change. The

muscular bulb consists of two layers of large lon¬

gitudinal muscle fibres separated by a very thin

laver of structured connective tissuc with a thin

innermost laver of circular fibres. Ihc lumen of

muscular bulb is very broad. The ocsophagus

diameter does not increase markedly after the

nerve ring.

I hc radula consists of rwo rows of marginal cccth

only. The teeth arc elongated, leaf-shaped, slight-

ly curved in profile and consist of two flattened

parts tightly adjoining each other (Sysoev &

Kantor 1986).

ZOOSYSTEMA • 1999 • ZI (2)

183

Medinskaya A. I.

Fig. 10. — Megasurcula carpenteriana, reconstruction based on longitudinal sections through the foregut.

Plicisyrinx decapitata Svsoev & Kantor, 1986

(Fig. 9)

The rhynchodeum sphincter lies in the anterior

position and shifted backward. The rhynchodeal

wall epithelium difFers from that of the proboscis

wall. The rhynchodeum is not evertible.

The proboscis is long, with virtually the same

diameter along its entire length, but the apical

part is narrower. The sac-like buccal chargement

and epitheli.il pad are absent. The buccal tube

sphincter is anterior, the distance from rhe

mouth to the sphincter is slightly more than the

tooth length. Tecth are ver)' long. The walls ol

proboscis, buccal tube, and oesophagus are very

thin. The proboscis wall thickness ts about 5-

6%, and buccal tube wall thickness is about 2%

of the total proboscis diameter. The basal part of

the proboscis is externally covered by a columnar

epithelium with a rhin cuticle. The apical part is

covered by a cubic epithelium, also cuticülized,

with a very éveil surface. The buccal tube is lincd

with a rather low columnar epithelium, which

becomes larger after rhe beginning of buccal

mass. The oesophagus is lined with a columnar

epithelium without cilia.

The buccal nuss is large, muscular, hegins inside

the proboscis and slightly coiled. Buccal lips are

long and muscular. The radular sac opens our-

side the proboscis and the buccal sac is absent.

Salivary ducts are rather short, of small diameter,

not very coiled, opening rather far from the

radular sac, on both its sides, dorso-laterally and

ventro-laterally. Salivary glands are very small

and intermédiare betwcen acinous and tubular.

This type of salivary glands was designated as

modified acinous (Kantor <7 al, 1997). The

venom gland opens neax rhe radular sac, at the

rear boundary of buccal mass. A very short duct

is présent. Venom granules appear ncar the gland

opening. The venom gland histology doe.s not

change along its length. The muscular bulb is

very large, eousisung of nvo lavers of large longi¬

tudinal muscle fibres separated by a layer of

structured connective tissue, with a rhin inner-

most layer of circulai- fibres. The lumen of the

muscular bulb is very broad. 1 he oesophagus is

medium elongated. Oesophagus diameter does

not merease posterior to the nervous ring.

The radula consists of two rows of small, llatten-

ed, slightly curved, wishbone teeth (Svsoev &

Kantor 1986).

Megasurcula carpenteriana (Gabb, 1865)

(Figs 10; 11)

The rhynchodeal sphincter is in the anterior

position, very large. The sphincter is formed by

longitudinal muscles. Longitudinal muscle

bunches pass from the sphincter downward and

184

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

Fig. 11. — Megasurcula carpenteriana, buccal mass (reconstruction based on longitudinal sections through the foregut).

attach to the rhvnchodeum wall. The rhyncho-

deum wall epithelium is not glandular. The

rhynchodeum is not evertible.

The proboscis is long, wide and coiled inside the

rhynchodeum. At the tip of the proboscis, there

are two anterior sphincters and a sac-likc buccal

enlargement between thent. The epithélial pad is

absent. The buccal tube wall between the two

sphincters i.s underlain by a layer of large circular

fibres, i.e. there is a kind of long and thin

sphincter. The buccal tube is folded, with very

thin walls. Proboscis walls are rather muscular

ZOOSYSTEMA • 1999 • 21 (2)

and of the same thickness along the entire lengtlt.

The wall thickness is about 9% of the total pro¬

boscis diameter and the buccal tube wall thick¬

ness is about 5%. Between the proboscis base

and the beginning of buccal mass, the buccal

tube is strongly coiled. very thin, and forms a

rather long loop, which passes along the buccal

mass like a wide and thin ribbon.

The buccal mass is large, very muscular, lient and

situated outside of the proboscis. Buccal lips are

absent. The radular sac has a wide opening, the

buccal sac is absent. Salivary ducts are long,

185 I

Medinskaya A. I.

Fig. 12.— Carinoturris polycaste, reconstruction based on longitudinal sections through the foregut.

slightly coiled and open on both sidcs of radular

sac. Salivary glands arc large and acinous. The

venom gland opens bv a short duct at the border

of buccal mass. The venom gland histology does

not change along its length. The muscular bulb

was not located on the slides. The élongation ol

oesophagus is well-expressed, the oesophagus

forms a long loop near the buccal mass. The

oesophagus diameter does not increase after the

nerve ring.

Radular teeth are wishbone, with the distal limb

of the base dctachcd (Powell 1966).

Carinoturris polycaste (Dali, 1919)

(Fig. 12)

The rhyuchodeum sphincter is in the anterior

position, medium-sized, situated around the

rhynchostome. The rhynchodeum wall epithe¬

lium diflers from the proboscis wall external epi¬

thelium. The rhynchodeum is not evertible.

The proboscis is long, with almost no change in

diameter along its length. The apical part is coil¬

ed inside the rhynchodeum. The buccal tube

from the mouth to the first sphincter is very nar-

row. At the apical end of proboscis there are two

anterior sphincters, the first one is very small,

with a marked bend and a sac-like buccal enlar-

getnent behind it. The second sphincter is larger

and situated behind the sac-like buccal enlarge-

mtnr. There is a large and very high épithélial

pad within the enlargement. The proboscis wall

thickness is about 2ü% of the total proboscis dia¬

meter, the buccal tube wall thickness is only

about 4%. The buccal tube is lined with a rather

low columnar epithelium, which becomes taller

after the buccal mass beginning. The oesophagus

is lined with a columnar epithelium.

The buccal mass is rather large, not very muscu¬

lar and situated ourside the proboscis. The rhick-

rtess ol the buccal mass wall différa only slightly

Iront that of the buccal tube. An alteration of

muscle structure of the wall occurs far from the

venom gland opening, as if the venom gland

186

ZOOSYSTEMA ■ 1999 • 21 (2)

Foregut anatomy of Cochlespirinac

Table 2. — List of characters and States used in cladistic analysis.

1. Epithelium of posterior rhynchodeal wall:0, glandular; 1, not.

2. Proboscis length. 0, long; 1, short.

3. Proboscis wall thickness (in % of proboscis diameter): 0, more than 10%; 1, less than 10%.

4. Proboscis tip. 0. typioal: 1, Camtas-wWe

5. Epithelial pad at lip of buccal tube: 0. présent; 1, absent.

6. Anterîor buccal tube sphincters: o, one; 1, two; 2. absent

7. Sac-like chargement of ttie buccal tube; 0. présent, 1, absent.

8. Shifting buccal tube sphincter; 0 absent 1, présent.

9. Buccal lips. 0. absent; 1. uninvertible; 2, invertlble.

10. Position of the buccal mass; 0, within the proboscis; 1. posterior to the proboscis base; 2, at the proboscis base.

11. Buccal mass Sharp; 0, not curved. 1 curved

12. Elongation of the oesophagus. 0, absent; 1. présent.

13. Increasing of the oesophagus diameter 0, absent; 1, présent.

14. Salivary glands: 0, acinous; t, anastomosing tubular; 2, modified acinous.

15. Venom gland duct: 0 présent; 1, absent. 2, modified,

16. Position of ooening of venom gland lato oesophagus: 0, at the border of buccal mass; 1, behind buccal

mass.

17. Number of muscular layers of lhe muscular bulb: 0, three layers with the connective tissue; 1, one or two layers

without connective tissue.

18. Orientation of fibres in outer two layers of muscular bulb: 0, similar orientation; 1. opposite orientation.

opens in the middle of lhe buccal mass. Buccal

lips are large and muscular, with a very narrovv

space between them. I he radular sac has a wide

opening, and the buccal sac ts absent. Salivary

ducts are short, scraight and open on both sides

of radular sac. Salivary glands arc very small and

acinous. l he venom gland opens near the radu¬

lar sac opening, with a very small duct. l he

venom gland histology dues nor change along its

length. The muscular bulb is composed ot two

layers of large longitudinal muscles separated by

a laver of unstructured connective tissue, with an

innermo.st laycr of circular fibres. Ail muscle

fibres arc very thin. l he lumen of the muscular

bulb is rather wide. l he oesophagus is medium

elongated. Oesophagus diameter does not increa-

se behind the nerve ring.

CLADISTIC ANALYSIS AND RELATION-

SHIPS WITHIN THE SUBFAM1LY

The analysis was based on 18 characters recogni-

zed in previous Works on Conoidea (Taylor et ai

1993; Kantor et ni. 1997) (Table 2). Some cha¬

racters wcte somewhat changcd, taking into

account the peculiarities of tlie given subfamily:

Character 3. Lhe ratio between the proboscis

wall thickness and the proboscis diameter appear-

ed to be important in distinguishing some spe-

cies, winch possess a large proboscis, but the wall

thickness does not exceed 10% of diameter, and

there are large longitudinal muscles passing along

the buccal cube.

Character 4. It emphasizes the présence of a

thin-walled stalk at rhe proboscis apical end in

some species.

Character 8. lt considers that group of sphinc¬

ters which cannor be assigned neither to anterior

nor to posterior (sce Discussion).

Character 13. It considère the sharp alteration in

the histological structure of rhe oesophagus wall

in some species.

Character 15. An additional siate two was intio-

duced for MtmhaUcna philippinarum w h ose

venom gland changes its histology far behind the

nerve ring whilst the duct ts apparently absent.

The data on some species of Aforia Dali, 1889

were incomplète, bccau.se only the very anterior

parts of digestive System were sectioned.

The analysis was performed with Ccmmula

deshayesi (Douinet, 1839) taken as an outgroup

(Table 3). The data on t h is species were extracted

(rom Kantor et al. (1997). Characters 4, 8. 13,

14 and 16 were rreated as ordered.

A heurisric seareh produced 12 equally pareimo-

nious trees (47 sceps, consistency index = 0.489,

rétention index = 0.718, homoplasy index =

ZOOSYSTEMA - 1999 - 21 (2)

187

Medinskaya A. I.

Table 3. — Distribution of character States for the cochlespirine gastropods examined. Details of characters are listed in Table 2.

Character

Gemmula deshayesi

Aforia inoperculata

Aforia circinafa

Aforia lepta

Aforia moskalevi

Aforia aulaca alaskana

Aforia crebristhata

Aforia abyssalis

Aforia kupriyanovi

Antipianes sanctiioanriis

Carinoturrls polycaste

Cochlespira pulchella

Cochlespira radiata

Comitas iïiurrawolga

Comitas onokeana vivens

Kurilohadalia elongata

Leucosyrinx pyramidalis

Leucosyrinx vernlli

Marshallena philippinarum

Megasurcula carpenteriana

Plicisyrinx decapltata

0.511). The trees were rather uniforra, and most

branches were supported in ail trees. The résolu¬

tion of the analysis was poor raostly in branches,

consisting of numerous species of Aforia and in

that including two nearly identical morphologi-

cally species of Cochlespira. This, however, seems

rather natural, because anatomic-al data used in

the analysis, usually do not allow to distinguait

species of a genus.

A strict consensus tree is shown in Figure 15.

Table 4. — Synapomorphies for irrterior nodes, Nodes numbe-

red as in Figure 15.

Node

Synapomorphies

(Character: State)

3(1)

9(1)

5(1)

9(1), 14(1)

15(1). 17(1)

4(1), 6(2). 8(1)

3(1)

16(1). 18(1)

10(2)

5(1)

2(1)

9(2), 11(1)

5(1)

The synapomorphies for internai nodes are listed

in Table 4.

The main resulc of the analysis was the division

ot the taxa studied into three large groups, which

correspond wcll to those bascd on estimation of

kcy anatomical peculiarities (sec below). 1 hese

groups may represenr different evolutionary

lineages within the Cochlcspirinae, but any taxo-

nomic conclusions seem to be prématuré, due to

a rather small percentage of cochlespirine généra

covered bv the study.

DISCUSSION

Anatomicai. variation

The subfamilv Cochlcspirinae is not very diverse

in foregut morphology. It is possible to recognize

three types of the foregut organi/ation. The most

important characters are the position and struc¬

ture ot venant gland with rnuscular bulb, while

ail other characters, such as die position and

structure of buccal mass, the buccal lips, the sali-

vary glands, and the sphincters of buccal tube are

eîther uniforra (e.g., salivary glands), or too

variable, even within the genus (e.g., the number

of buccal tube sphincters). Therefore, the ana-

188

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinac

lysis of structure of the vvhole digestive System

may be based only on the complex of characters.

The présence of glandular epithelium in the

rhynchocieum is characteristic of the majority of

cochlespirines, except Aforia lepta and Mega-

surcula carpenteriana. The sanie conccrns the

protraction of proboscis, which is permitted by

eversion of the posterior part of rhynchodeum.

The rhynchodeum walls in the posterior part are

not attached to the body walls. During the pro¬

boscis protraction, rhey move rogerher with the

proboscis and become the continuation of its

walls, thus making the proboscis substantially

longer. In rnost cases, exceptüng rnost Aforia spe-

cies, there is also a loop of oesophagus .situated

behind che buccal mass but anterior to clic nerve

ring. The presence of che loop is connected with

proboscis profraction and provides an additional

length to avoid the passage of structures, situated

behind the nerve ring, through the ring during

the proboscis protraction (Fonder 1973). The

proboscis is u.sually long, except in some species

of Aforia. 1 lie latter, however, u.sually possess a

loop of oesophagus, which may indicate that the

proboscis is in a retraefed position.

The structure of rhe apical part of the proboscis

is rather variable. No species of Cochlespirinac

was found to possess intermediate or posterior

sphincters, Antenor sphincters may be absent, as

in Aforia lepta\ ali other species srudied possess

either one (e.g. Aforia circinata, Plichyrinx deçà -

pitata , Marshallena philippinartttn , Antiplanes

sanctïioannis ), or two anterior sphincters. The

sac-like en largement of buccal mass is usually

présent, except in three species (PlicisyrinX deen-

pitata , Aforia lepta, and Marshallena philippine!-

mm), while rhe epithélial pad is presepr in only

half of the examined species. Different combina¬

tions may be found within a genus and e\'en sub-

genus and, as in other subfamilies (Taylor et al.

1993; Kanror et al. 1997), there is no çorrespon-

dence with other anaromicai characters of rhe

foregut.

The apical part of the proboscis i.s of spécial

interest in four species; Co mi tas murrawolga,

C. onokeana, Antiplanes sanctïioannis and

Marshallena philippniarum. In this group of spe¬

cies the proboscis narrows abruptly in the apical

part. This resuhs in the formation of a thin stalk

which is a continuation of the proboscis and is

two to four cimes longer thac the radular toorh

length. Inner and outer walls of the stalk are very

thin, bilayered, formed by longitudinal and cir¬

culai - muscles, and lined with rhe saine epithe¬

lium. The wider part of the proboscis contains

one or two buccal sphincters (ail species), a sac-

like enkirgement (Cotaitas finlay, 1926) and an

épithélial pad ( Carmins .nid Ami planes sanctiiaun-

nis). In both species of Comitas, at some distance

from the stalk the exremal columnat - epithelium

abruptly changes to a lower cubic epithelium,

identical to the inner epithelium of the buccal

tube (Fig. 5A, B). There is no .such change in the

epithelium in Antiplanes and Marshallena,

though in Antiplanes the columnat epithelium of

the outer proboscis wall gradually transfers kilo

the cubic onc on the outer wall of the stalk.

The stalk may vary in length in different spéci¬

mens of Comitas and Antiplanes. When the stalk

length changes, rhe position of buccal sphincters

in relation to rlie proboscis tip also changes

(Fig, 5A, B). The distance from the sphincter to

the proboscis tip is abolir four radular tooth

lengths in Figure 5B, whereas it is about two

tooth lengths in Figure SA. Figure 6 shows a spé¬

cimen of Antiplanes with the longest stalk, when

the sphincters are rnost close to the proboscis tip

and the buccal mass lies inside the proboscis

base. Another specimen of Antiplanes (not figu-

red) had a shorter stalk, the distance betwen

sphincters and the proboscis tip was greater, and

the buccal mass was shifted from the proboscis

base ourwardx.

h is problemaric to ascribe thèse sphincters to a

category. h was suggested to designate sphincters

as anterior if the distance from a sphincter to the

mouth opening does not exteed 2.5 tooth

lengths (Kantor et ai 1997). In the situation

tinder considération, the position of sphincters

may va/y, taking either anterior or posterior posi¬

tion Nevertheless, the presence of a sac-like

enkirgement and an épithélial pad indieates that

the sphincters are used for the tooth fixation

and, therefore, are functionally anterior. Such

sphincters, varying in position but functioning as

anterior, are designated here as shifting sphinc¬

ters.

When discussing the structure of anterior part of

ZOOSYSTEMA ■ 1999 • 21 (2)

189

Medinskaya A. I.

con

Fig. 13. — A Cochlespira pulcliella, oesophagus portion near the nerve ring, with lhe venom gland opening; B. Megasurcula car-

penteriana. buccal mass, with lhe opening of lhe venom gland duct.

the proboscis, we avoidcd che use of the term

“mouth”, hecause it is dchned functionally and,

in this case, does noc coïncide with anatomical

tip of the proboscis. The mouth is usually cha-

racterized by a change of ouier epithelium of che

proboscis vvall for che inner epithelium of che

buccal tube, and by an alteration of the wall

muscular structure. Similar changes occur at the

base of che scalk in the species considérée! This

fact and identity of the epithelium of inner and

outer wall,s of the stalk with the epithelium of

the resr pari of che buccal tube allow to suggest

that the stalk représenta an everted parc of che

buccal tube. Howcver, the absence of a distinct

change in epithelium in Marshallerm philippina-

rum dcies not permit to State reliahly that che

stalk is bomologous in ail lour species.

The buccal mass mav be situated eithet at che

proboscis base or outside ic. Therc arc no species

of Cochlespirinae examined wliich possess the

buccal mass inside the proboscis. Buccal lips are

absent iu Comitas onokeana vtvens , Leucosynnx

verrilli and Megasunula carpenteriana , în ail

others they are présent. Both species ol Cot h In¬

spira, one of Aforia, Leucosyrinxpyramidal:.'!, and

Comitas murrawolga hâve inverred buccal lips. It

is very interesting that in two Comitas species,

one lacks the buccal lips, and another possesses

inverted buccal lips. In other respects, these spe-

cies are virtually identical to each other anatomi-

cally. This shows that the presence and che posi¬

tion of the buccal lips may nor be good charac-

ters for the analysis of anaromy, and may vary

even witbin a genus.

Saltvary glands, except in two species, are the

normal acinous type. Anastomosing tubular

glands are présent in Kurilohadalia , and the

modified acinous type — in Pliehyrinx. These

types of salivary glands were fitst dcscribed in the

species of the subfamily Crassispirinae (Kantor et

al. 199%

The structure of the venom gland it.self is sintilar

in ail the species scudied, except Marshallerm phi-

lippinamm. The venom gland does not change

its hîstology in the passage rhrough the nerve

ring and opens directlv or by a small duct in the

oesophagus. l he location of rhe venom gland

opening corrélâtes with the structure of muscular

bulb. When the venom gland opens behind the

buccal mass, rhe duct is absent and the muscular

bulb is composed of two oppositely oriented

muscle layers, with no connective tis.sue laver.

When the venom gland opens at the buccal mass

border, a very small duct may be présent and the

muscular bulb is composed ot rhree muscle

lavers. Two outer layers hâve a similar orientation

and are separated by a connective tissue layer.

1 he venom gland may also open at the border of

rhe buccal mass wiihout a duct. In this situation

the muscular bulb is usually composed of two

190

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

Fig. 14. — Types of foregut organization. A, Atoria spp,, Leucosyrinx pyramidalis, Cochlespira pulchella, C. radiata ; B, Antiplanes

sanctiioannis, Marshallena philippinarum, Comitas onokeana vivens, Comitas murrawolga', C, Carinoturris polycaste, Kurilohadalia

elongata, Leucosyrinx verriili, Megasurcula carpenteriana.

similariy orientée! muscle layers, vvithout a

connective tissue layer (e.g. Comitas sp., Anti-

planés sanctiioannis). Marshallena philippinarum is

the exception, possessing the muscular bulb com¬

posée! o! a single muscle layer and the venom

gland of a very peculiar structure (see description).

It is necessary to define the terni “venom gland

duct” (Fig. 13A, B). We consider the venom

gland duct being présent if there is an abrupt

narrowing bclorc the opening and at le-ast a small

part of the gland Iacks venom granules. If the

venom gland has a large opening and the venom

ZOOSYSTEMA • 1999 • 21 (2)

191

Medinskaya A. I.

granules are présent just néar the opening, we

consider rhe venuni gland duel: lo be absent.

Observations lierein suggest tbat ihe site and the

mode of the venom gland opening, and the

structure of the muSculat bulb are closely interre-

lated and very important. In any case, it is now

évident that if the venom gland duct is absent,

the number of muscle layers ol the muscular

bulb is reduccd to tvvo (and cven to one in

Marshallena). When the venom gland duct is

présent, irrespective of ics lcngtli, the muscular

bulb is composed ol three layers. For example,

some Cochlespirinae (c.g. Carinohmis polycaste ,

Kurilohadalia elongata , Leucosyrinx verril/i,

Megasurcula carpenteriana, Plicisyrinx dealpiltila)

hâve a very short duct, whereas ail Crassispirinac

(Kantor et al. 1977) hâve a long duct, and a very

similar threedayer structure of the muscular

bulb.

As stated above, one of the rnost important crite-

ria in distinguishing the rhree morphological

groups in Cochlespirinae unexpectedly became

the structure of the venom gland and the muscu-

lar bulb, though similarities and distinctions in

the foregut organisation should be also taken

into account.

The first group indudes 11 species. The princi¬

pal charactcrs arc the position of the venom

gland opening (beyond the buccal mass) and the

structure of the muscular bulb composed of rwo

opposirely orîented muscle layers (Fig. 14A).

This group consiscs of A for ta spp., Leucosyrinx

pyramidalis , Cochlespim pulchella, C. radiata, The

three latter species hâve a well-developed oeso-

phageal loop and the distance from the buccal

mass to the nerve ring and, correspondingly, to

the place of the venom gland opening is rather

long. In ail species ol Aforia this distance is

shorter. In fact, some Aforia species (e.g„ A. nios-

kalevi , A. abyssahs) bave an oesophageal loop but

it is not very long. In A circinata and A. crebris-

triata, the distance betvveen the buccal mass and

the nerve ring is very short, However, in ail

Aforia species the nerve ring is situated at the

same level as the buccal mass, but Jaterally. As a

resuh, the ocsophagus curves behind the buccal

mass and passes along the buccal mass for some

distance. Th us even in species lacking the ocso-

phageal loop, the oesophagus deviates from the

buccal mass at an angle of about 90". In many

species (e.g. Leucosyrinx pyramidal]s, Aforia circi¬

nata, A. lepta ), the end of the buccal mass is not

clearly differentiated histologically. The buccal

mass wall gradua!!)' becomes thiimer, almost

without a change in histology. A distinct change

occurs only near the nerve ring and behind it,

wlicre oesophagus diameter greatly incrcases.

This inc reasc is characteristic ol Leucosyrinx pyra¬

midalis and ail Aforia species, including chose

havinga weil-differentiatcd buccal mass.

The position of the venom gland opening

behind the buccal mass is probably the resuh of

an élongation ol oesophagus at the expensc of

rhe rear part of the buccal mass. This may be evi-

deneed by the absence of clearly differentiated

post-erior end of the buccal mass in some spe¬

cies. h is possible to assume that rhe buccal mass

irself exrends to rhe nerve ring, and the oesopha-

gus expansion after the ring wirh the changing

oesophagus wall histology represents rhe end of

the buccal mass. However, both species of

Cochlespim , winch are superfïcially very similar

to Aforia , bave a well-differenriared buccal mass,

a long oesophagus loop, and l.tck rhe oesophagus

expansion behind the nerve ring. That is, in

these species the élongation ol oesophagus took

place, but il occurred betvveen the buccal mass

and the venom gland opening.

Aforia lepta is a spécial case, because it complete-

ly lacks an oesophageal loop, its venom gland

opens at rhe end of rhe buccal mass, and the

nerve ring is situated just behind. However, the

muscular bulb is also composed of rwo opposite-

ly oriented layers, and the oesophagus expansion

is présent. This type of structure looks very pecu-

liar, but it is necessary to rake into account that

the original description (Sysoev & Kantor 1987)

was based on a single juvénile specimen, and it

canner be excluded that rhere may be rather sub-

stantial ontogenetic changes.

The second group consists of Antiplanes sanc-

tiioannis,. Cornu as munawolga , C. onokeana

ta tiens, and Marshallena phtlîppinarum, wirh the

latter species occupyïng a spécial position. This

group is characrerized by the structure of the

muscular bulb composed of rwo muscular layers

wirh similar orientation (or one muscular layer in

Marshallena philippinarum)\ and the venom

192

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

_ Gemmula deshayesi

- Aforia inoperculala

- Leucosyrinx pyramldalis

- Cochlespira pulchella

-Cochlespira radiata

-Aforia circinata

-Aforia lepta

-Aforia moskalevi

-Aforia aulaca alaskana

-Aforia crebristriana

-Aforia abyssalis

-Aforia kupriyanovi

-Antiplanes sanctifoannis

-Comitas murrawolga

-Comifas onokeana vivens

-Marshallena philippinarum

-Carinoturris polycaste

-Kurilohadalia elongata

-Plicisyrinx décapita fa

-Megasurcula carpenteriana

-Leucosyrinx verrilli

Fig. 15. — Strict consensus tree.

gland opening at the boundary of the buccal

mass, without a duct (except in Marshallena phi¬

lippinarum) (Fig. 14B). An additional character

is the structure of the proboscis apical part, with

a stalk and shifting sphincters. Marshallena phi¬

lippinarum vvas assigned to this group for several

reasons. In spite of the unique structure of the

venoin gland (see description) and the presence

of only one layer in the muscular bulb, the latter

is very siniilar to that ol ail other species of this

group, and the structure of the apical end is also

approximately the same.

The third group includes Carinoturris polycaste,

Kurilohadalia elongata, Leucosyrinx verrilli,

Megasurcula carpenteriana, and Plicisyrinx decapi-

tata (Fig. 14C). These species hâve no clearly

expressed features, and cannot be assigned to the

two other groups. In general, besides the three-

laycred muscular bulb with very large muscle

fibres, they are characterized by a weakly coiled

proboscis with a large and uniform diameter.

The proboscis diameter is mostly occupied not

by walls (less than 10% of the total diameter)

but by longitudinal muscle bunches. The venom

gland does not change its histology and opens by

a s'tnall duct at the border of the buccal mass.

The muscular bulb is composed of three muscu¬

lar layers, two outer having a similar orientation

and separated by a layer of connective tissue. The

exception is Carinoturris polycaste, which pos-

sesses a very thin proboscis Wall. Its muscular

bulb is composed by very thin fibres. The size of

ZOOSYSTEMA • 1999 • 21 (2)

193

Medinskaya A. I.

Fig. 16. — Radulae of Cochfespirinae. A, B, Cochlespira pulchella\ C, Cochlespira radiala; D, E, Leucosyrinx (Sibogasyrinx) pyrami-

dalis.

the fibres is very important, as it will be shown Kurilohadalia elongata and modified acinous in

below. Other types of salivary glands also occur Plicisyrinx clecdpitatd, previously recorded in the

in this group, i.e. anastomosing tubular in Crassispirinae (Kantor et al. 1997).

I 194 ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochiespirinae

Fig. 17. — Radulae of Cochiespirinae. A, Comitas onokeana vivens; B, Comitas murrawolga - middle part; C, Comitas murrawolga,

initial part; D, E. Marshallena philippinarum.

Shell and radular morphology

Shell characters appeared ro be r.uhcr poorly cor-

related with features of foregut anatomy. It may

be only mentioned rhat menibers of the first

group, separated by anaromical characters and in

the cladisric analysis, possess a typical cochlcspi-

rine elongated fusiform shell with a long anterior

canal and weak axial sculpture. Species of the

second group hâve stouter shelU with shorter

canals and well-developed sculpture (except for

Antiplanes). The third group is very hetero-

geneous in shell characters corresponding to the

wide anaromical variation. Ilowever, no shell

character was found to strictlv correlate with a

feature of the foregut anatomy, or a set of anato-

mical characters.

The radular morphology of cochlespirines had

previously been assigned to the wishbone tooth

type (Taylor Pt ni. 1993). Robust, short, and curv-

ed marginal teeth with a knîfe-like cutting edge

were found in Antiplanessanctïwannis (Kantor Si

Sysoev 1991; Taylor et ni. 1993). Teeth of

Afp nu spp. (Sysoev 8 1 Kantor 1987) are similar

to teeth al Antiplanes.

Among the species stndied, more or less typical

wishbone teeth were found in Leucosyrinxpyra¬

midale (Fig. I (SD, F.) and Marshallena philippi-

narum (Fig. 17D, E). They are quite comparable

ro the marginal teeth of many Crassispirinae (e.g.

Inquisitar- and Hindsiclava- type teeth - see

Kantor et al. 1997).

At the saine time, the marginal teeth of both spe¬

cies of Comitas studied seem to be not true wish¬

bone. At an early stage of formation, they

ZOOSYSTEMA • 1999 • 21 (2)

195

Medinskaya A. I.

resemble a broad liât plate thickened apically and

on outer si des (Fig. 17C). Later, the teetb become

more similar to wishbone: die latéral thickenings

become more distinct and appear like the separate

limbs ut a typical wishbone tooth. This process is

more pronouneed in C. mumiu'olga (Fig. 17B,

C), whereas in (.. onokedna the teeth remain fiat

and poorly dilferentiated (Pig. 17A). Never-

theless, a complété séparation of tooth limbs

does not occur in any of Comitas species.

The teeth o( species of Cuchléspirtt are somewhat

different. In C. pulcheltd, they are of a normal

wishbone form, with a narrow limb (Fig. 16A, B)

whereas the teeth of C. radiata (Fig. 16C)

consist ot a longitudinally tbldcd plate, torming a

central canal, and re.semble enrollcd hollow

teeth, recogni/.cd by Taylor et al, (1993) for

Pihbryspira. It may bealso added rhat the diversi-

ty of Cncblespim teeth seems to be rarher high, as

indicared by drawings of radula of different .spe¬

cies (Powell 1966: figs 17-70).

The central tooth may be variously developed:

front a well-tormed plate, with a strong cusp and

clearly defined boundaries, as in Cochlespira

(Figs 16A-C), to a weak formation, vvhich looks

like a membrane fold rather rhan a toodi (e.g.,

Comitas unokeana , Fig. 17A). h should be men-

tioncd chat the central part of the radula of

C. onokedna at early stages of formation shows

two separate plates instead of a single tooth. This

may indicate that the central "tooth" is actuallv a

producr of fusion of two teeth, and supports the

idea of Kantor &t Sysoev (1991) on the origin of

central tooth in Aniiplanes.

No corrélation was fuund between the anatomy

of digestive System and radular morphology in

any of the species studied.

COMPARISÜN WITH OTHER

SUBFAMILIES OF TURRIDAE

This article is the second in the sériés of Works

concerning Turridae, and in the first paper

(Kantor et al. 1997) short descriptions ofall sub-

famiües werc given. Therefore, hcre we will

mainly compare the two conchologically similar

and best studied subfamilies: Cochlespirinae and

Crassispirinae.

The subfamily Crassispirinae is characterizcd by

very diverse structure of proboscis, buccal mass,

and sulivarv glands. The proboscis structure is so

variable not only within the subfamily, but even

within the généra and subgenera, chat a total of

13 types of foregut organization hâve been reco-

gnized. The must characteristic features of the

subfamily are as follows: the présence of the

rhynchodeunt glandular epithelium, the ability

ot the rear part of rhynthodeum to even during

the proboscis protraction, che position of the

oesophagus élongation usuallv between the buc¬

cal mass and the nerve ring, with a more or less

long oesophagus loop, the présence of non-

glandular venom gland duct, and the change in

its hisrology behind the nerve ring, the presence

of threedayered muscular bulb, with a layer of

connective tissue (the bulb is composée! by small

fibres and the two outer layers hâve a similar

orientation) (Taylor et al. 1993; Taylor 1994;

Kantor et al. 1997).

The firsr two characters are présent also in

Cochlespirinae. Thus, the most characteristic

features of this Family become as follows: the

absence of change in the venom gland histology

behind the nerve ring in ail groups; the complété

absence of venom gland duct and, correspondis

the two-layered muscular bulb, without any

connective tissue, or the presence of a small duct

and the three-layered muscular bulb, like in cras-

sispirines. In cochlespirines the muscular bulb,

with a similar organisation and arrangement of

muscle layers, is cotnposed of very large fibres,

distinguishable even ai a small magnification,

whereas in crassispirines they arc very small and

form a compact layer.

Représentatives of both subfamilies are rather

well distinguishable conchologically, when typical

forms are considered. Species of the subfamily

Crassispirinae are generally characterized by a

small clavifbrm shell with a rather well-developed

sculpture, while the shell of typical cochlespirines

is fusiform, with a long anterior canal and a

rather weak sculpture. But many représentatives

(e.g., Plirhyrinx) represent intermédiare variants

in conchological characters, and it is thus diffi-

cult to separate reliably these subfamilies only by

the shell.

Anatomy of six species of Clavatulinae has been

196

ZOOSYSTEMA • 1999 • 21 (2)

Foregut anatomy of Cochlespirinae

described: Turricula nelliae spurius (Hedlev,

1922), Turricula javana (Linnaeus, 1767),

Toxtdionellit tumida (Sowcrby, 1870), Clionella

sinuata (Boni, 1778), Cia vaut!a caerulea

(Weinkauff, 1875), C. mûri cala (Lamarck, 1822)

(Kantor 1990; Taylor 1985, 1994; Taylor et ai

1993). The main différence of Clavarulinae from

Cochlespirinae and ail other turrid subfamllies is

the position of the buccal mass inside the pro-

boscis, near the mouth. As in Cochlespirinae, the

buccal sphincters in Clavarulinae mav be either

présent or absent. Oeasophageal loop is similar

to that in sotne Cochlespirinae and Crassis-

pirinae. Vcnom gland opens in rhe buccal mass.

The structure of muscular bulb is somewhar dif¬

ferent from that in Cochlespirinae: the bulb is

composée! of two similarlv oriented layers, with a

connective tissuc between layers.

A detailed comparison with Turrinae is currently

rather difficult, due to insufficient data on this

subfamily, Features characteristic of Turrinae

often occur also in Cochlespirinae. For examplc,

oesophageal loop may be absent, venom gland

does not change its hisrology anterior to the

nerve ring and opens ar the buccal mass bounda-

rv by a short duct. Howcver, the structure of

muscular bulb differs from that found in ail

Cochlespirinae: its bulb is composed of two simi-

larly oriented layers, with a connecrive tissus bet¬

ween layers. Another distinguishing character of

Turrinae is the rhynchodeum lined with gland-

ular epithelium along its entire length (Leviten

1970, cit. after Taylor et al. 1993; Taylor et al.

1993; Taylor 1994).

SYSTEMATtC CONCLUSIONS

The data obtained conftrm that the main part of

the subfamily Cochlespirinae represents an ana-

tomically distinct group, differing from other

subfamilies of Turridae in several minor but

constant characters. At the same time, the subfa-

mily in its currently adopted volume seems not

to constïtute a monophyletic group.

A spécial note should be made on the systematic

position of the five species belonging to the third

anatomical group. The cladistic analysis showed

that four of the five species ( Kurilohadalia elon-

gaia, Leucosyrinx verrilli, Megasurcula carpenteria-

nd , and Plicisyrinx decapitata) are clearly separa-

red in ail cladograms, whercas Carinoturris

polycaste occupies an isolared position and is a

sister-group of ail species from the first and

second anatomical groups. The group of the four

species is deftned on the cladogram by a single

synapomorphy (relatively very thin walls of the

proboscis), but evolurionary and funcrional

significance of this character is unclear. In fret,

rhe rhird anatomical group is somewhar interme-

diatc in its morphology between Crassispirinae

and Cochlespirinae and possesses some charac¬

ters of both subfamilies. Conchologically, sotne

of tbe taxa are also similar to représentatives of

borh subfamilies and earlier were alternately

referred to rhe Crassispirinae or to the

Cochlespirinae (e.g. Kurilohadalia and Plicisyrinx

- see above)

Two species of the genus Leucosyrinx were refer¬

red to different groups in both cladistic and ana-

tomicaJ analyses. These species belong to

different subgenera: Leucosyrinx Dali. 1889 s. >tr.

(verrilli, type species of the genus) and

Sibugasyrinx Powell, 1969 (pyramidales, type spe¬

cies of the suhgenus). The grcat différences in

anatomy indicate that these subgenera are not

closely relared, and the rank of the subgenus

Sibogasyrinx should be raised to a full genus.

Acknowledgements

The author is very grateful to John Taylor,

Philippe Bouchet, Yuri Kantor, and Alexander

Sysoev fortheir kind donation of rhe macerial for

the study. Thanks are also due to Yuri Kantor for

making SEM photographs, and Alexander

Sysoev. who discussed, corrected and improved

the manuscripr. The very useful comments on

the cladistic section and the linguistic corrections

made by an anonymous reviewer are also grate-

fully acknowledged.

REFERENCES

Kantor Yu. 1. 1990. —Anatomical basis for the origin

and évolution ol the toxoglossan mode of feeding.

Malacologia 32:3-18.

Kantor Yu. I., Medinskaya A. I. & Taylor J. D.

1997. — Foregut anatomy and relationships of the

ZOOSYSTEMA • 1999 • 21 (2)

197

Medinskaya A. I.

Crassispirinae (Gastropoda: Conoidea). Bulletin of

the Natural History Muséum, London (Zoology) 63:

55-92.

Kantor Yu. 1 & Sysoev A. V. 1991. — Mollusks of

the genus Antiplanes (Gastropoda, Turridae) of the

northwestern Pacific Océan. Nautilus 105:

119-146,

Leviten P. J. 1970. — The Structure and Function of

the Alimentary System o/Tolystira albida (Gastro¬

poda: Toxoglossa). M.Sc. Thesis, University of

Miami, Miami (USA).

McLean J. H. 1971. — A revised classification of the

family Turridae, with the proposai of new sublami-

lies, généra, and subgenera from the eastern Pacific.

Veliger 14: 1 14-130.

Ponder W. F. 1973. — Origin and évolution of the

Neogastropoda. Maldcologia 12: 195-338.

Powell A. W. B. 1942. — The New Zealand Recent

and fossil Mollusca of the family Turridae. Bulletin

of the Auckland lmütute and Muséum 2, 192 p.

Powell A. W. B. 1966. — The molluscan families

SpeightÜdae and Turridae. Bulletin of the Auckland

Institute and Muséum 5, 184 p.

SwofFord D. L. 1993. — PAUP, Phylogenetic Analysis

Using Parsimony. Smithsonian Institution, Labora-

tory of Molecular Systematics, Washington D.C.,

260 p.

Sysoev A. V. Kantor Yu. I. 1986. — New and rare

deep-sea species of the subfamily Turridae

(Gastropoda, Toxoglossa) from the northern part

of the Pacific. Zoologicheskij Zhurnal 65:

1457-1469 (in Russian).

Sysoev A. V. & Kantor Yu. I. 1987. — Deep-sea gas-

tropods of the genus Aforia (Turridae) of the

Pacific: species composition, systematics, and func-

tional morphology of the digestive System. Veliger

30: 105-126.

Sysoev A. V. & Kantor Yu. I. 1988. — Three new

species of deep-sea mollusks of the genus Aforia

(Gastropoda, Toxoglossa, Turridae). Apex 3: 39-46.

Taylor J. D. 1985. — The anterior alimentary System

and diet of Turricula nelliae spurius (Gastropoda:

Turridae): 175-190, in Morton B. & Dudgeon D.

(eds), Proceedings of the Second International

Workshop on the Malacofauna of Hong Kong and

Southern China, Hong Kong, 1983. Hong Kong

University Press, Hong Kong.

Taylor J. D. 1994. —- Foregut anatomy of the larger

species of Turrinae, Clavatulinae and Crassispirinae

(Gastropoda, Conoidea) from Hong Kong:

185-212, in Morton B. (ed.), The Malacofauna of

Hong Kong and Southern China. III. Proceedings of

the Third International Workshop on the

Malacofauna of Hong Kong and Southern China.

Hong Kong University Press, Hong Kong.

Taylor J. D.. Kantor Yu. 1. & Sysoev A. V. 1993. —

Foregut anatomy, feeding mechanisms, relationships

and classification of the Conoidea (= Toxoglossa)

(Gastropoda). Bulletin of the Natural History

Muséum , London (Zoology) 59 (2): 125-170.

Submitted on 12 January 1998;

accepted on 14 October 1998.

198

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology and systematics

of Brazilian Terebridae (Mollusca, Gastropoda,

Conoidea), with descriptions

of three new species

Luiz Ricardo L. SIMONE

Museu de Zoologia da Universidade de Sâo Paulo,

Cx. Postal 42694, 04299-970 Sâo Paulo, SP (Brazil)

lrsimone@usp.br

Simone Luiz Ricardo L, 1999. — Comparative morphology and systematics of Brazilian

Terebridae (Mollusca, Gastropoda, Conoidea), with descriptions of three new species.

Zoosystema 21 (2) : 199-248.

KEYWORDS

Gastropoda,

Terebridae,

anatomy,

systematics,

proboscis,

homology,

Brazil.

ABSTRACT

A detailed morphological study is provided ot the Brazilian Terebridae

HastuLa cinerea (Born, 1778), H. hastata (Gmelln, 1791), Terebra bnisiliensis

Smith, 1873, T. crassireticula n. nom. {replacing rite junior homonym

71 reticulata Simone & Verissimo, 1995), T. gemmulata Kiencr, 1839,

T. leptapsisn. sp., T. tauritia (Lightfoot, 1786), and T, spirosulcatu Simone &

P. M. Costa n. sp. Terebra sterigma n. sp. is represented by shells only.

Anatomically, the terebrids appear to be cliaracterized by the réduction of the

cephalic tentacles, the anierior extremiry of the ctenidial vein being promi¬

nent and without gill filaments, and the rhynchodeal introvert and the anus

being situated very posteriorly in the palliai cavity. The species of Hastula

examined are characterized by an enlarged font and by the complexity ot the

osphradium filaments. The species of Terebra examined are characterized by

the eye situated at the tip of the tentacles, and also by a tendency for enlarge-

ment of the introvert, and réduction of the probosds and the venom appa-

ratus with their en tire loss in some species (T gemmulata and T. brasiliemis).

The accessory proboscis structure is présent in lltree ot the Terebra species

studied, and it is hypothesized that it may be a diaracter of the lamily, as it is

présent in some Hastula species, and it may hâve been secondarily lost in

several species of the family. Based on anatomical similarities, it is hypothe¬

sized that the conoidea n proboscis is derived front a pleurembolic proboscis,

homologous to the buccal mass part of this proboscis type, and the rhyncho¬

deal wall is homologous to the remaining régions of the pleurembolic pro¬

boscis.

ZOOSYSTEMA • 1999 • 21 (2)

199

Simone L. R. L.

MOTS CLÉS

Gastropoda,

Terebridae,

anatomie.

systématique,

homologie,

proboscis,

Brésil.

RÉSUMÉ

Anatomie comparée et taxinomie de Terebridae du Brésil (Mollusca, Gastropoda,

Conoidea) ; description de trois espèces nouvelles.

Huit espèces de Terebridae du Brésil font l’objet d’une étude anatomique

détaillée : Hastula cinerea (Born, 1778), H. hastata (Gmelin, 1791), Terebra

brasiliensis Smith, 1873, T. crassiretiada », nom. (nom de remplacement

pour T. mutdaut Simone & Verissimo, 1995. invalide pour homonymie pri¬

maire), J. gemmulata Kiener, 1839, T. leptapsis n. sp„ 7. laurina (Lightloot,

1786) et i, spirusulcata Simone & P. M. Costa n. sp. Terebra sterigma n. sp.

est représenté uniquement par des coquilles vides. Au plan de leur anatomie,

les Terebridac sont caractérisés par une réduction des tentacules céphaliques ;

le vaisseau efférent de la branchîc dont l'extrémité antérieure, proéminente,

ne porte pas de filaments branchiaux ; l’introvers du rhynchodeum et l’anus

situés très en arrière dans la cavité palléale. Les espèces d Hastula examinées

sont caractérisées par le développement du pied et la complexité des fila¬

ments de l'osphradium. Les espèces de Terebra sont caractérisées par la posi¬

tion des yeux à l’exrrémiré des tentacules, par une tendance au

développement de l'introvers, et à la réduction du proboscis et de l'appareil à

venin, allant jusqu'à leur disparition totale chez certaines espèces ( T. gemmu¬

lata et T. brasiliensis). Des structures accessoires du proboscis sont présentes

chez trois des espèces de Terebra étudiées, et il est possible qu’il s'agisse d’un

caractère familial, car il esr présent chez certaines Hastula, et peut avoir été

secondairement perdu chez plusieurs autres espèces. Lu se fondant sur les

similarités anatomiques, il est fait l’hypothèse que le proboscis des Conoidea

dérive du proboscis de type pleurembolique et est homologue de la masse

buccale des espèces possédant ce type de proboscis, la paroi du rhynchodeum

étant homologue des autres parties du proboscis pleurembolique.

INTRODUCTION

The Terebridae comprise more than 300 species

in seas around ibe world (Bratcher &t

Cernohorsky 1987; Taylor 1990). Lltey arc cas il y

differentiated frorn die odier Conoidea by their

elongated, multiwhorled shell. The systematies

within the Family has, hoxvcver, been problc-

matic, with mosr species still placed in the

genus Terebra Bruguière, 1789 (Bratcher &

Cernohorsky 1987).

The Brazilian Terebridae vvere revised by

Matthews et al. (1975) who recognizcd seven

species of Terebra: T. taurina (Lightfoot, 1786);

T. gemmulata Kiener, 1839; T, brasiliensis (Smith,

1873); 7. concava Say, 1822; T. dislocata Say,

1822; T. protexta (Conrad, 1846) and T doello-

juradoi Carcelles, 1953 and tltree of Hastula H.

& A. Adams, 1853: 77 cinerea (Born, 1778);

H. hastata (Gmelin, 1791) and H. salleana

(Deshayes, 1859). Three more species hâve been

described since: T. riosi Bratcher & Cernohorsky,

1985; T. imitatrix Auffenberg & Lee, 198S (ail

présent in Rios 1994) and 7. reticulata Simone &

Verissimo, 1995. An extensive anatornical des¬

cription of a Brazilian terebrid is provided by

Marcus & Marcus (1960) for H. cinerea, and

briel data is provided in Auflenherg & Lee

(1988) and in Simone ôè Verissimo (1995).

Taylor (1990) provided a complété and compré¬

hensive history and commenfary on the tere-

brids. He summarized the current informai (for

systematies) classification ol the terebrid foregut

anatomy types (la, Ib, lia, Ilb, 111) found in the

literature (e.g., Miller 1971; Taylor & Miller

1990) with several interesting examples. The

classification ol the Conoidea (= Toxoglossa), as

a whole. has also been analysed in sorae recent

papers, ol which Taylor, Kantor Sysoev (1993)

is considered the most important. This paper

200

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

discussed each foregut cbaracter, erected a uni-

form terminology, and concluded witli a phylo-

genetic analysis of rhc sapcrlarntly based mainly

on :hc shell, thc operculum and forcgut anato-

rny. Among thc several conoidcans .malysed, thc

authors smdied seven terebrids, giving as synapo-

ntorphies of the family thc following characters:

(1) rhynchodeal inrrovert; (2) accessory proboscis

structure; (3) terebriform shell; (4) absence of an

anal sinus tn rhc shell apcrrure and (3) more than

ni ne teleoconch whorls.

Although the foregut anatonty is fundamental

for the understanding of the conoideans, and irs

characters hâve followed in importance that of

the shell in systematics, ir has obscured the

importance of other structures such as the repro¬

ductive System and the mande organs. Papers

with anatomical data on structures other than

the foregut are very few (e.g., Marcus & Marcus

I960).

This paper is part of a larger research project

concerntng the inter-relationship ot sortie

Caenogastropod grotips, based mainly upon

comparative morphology. Sonie Brazilian species'

of terebrids were selccted for detailed morpholo-

gical studv with rhc following objectives: (1) to

présent a holistic morphological study of each

species providing a basis for a discussion ol seve¬

ral characters including and beyotid those of the

foregut and providing a basis for a future phylo-

genetic analysis of the family; (2) to cstablish the

morphological characters of species Iront a géo¬

graphie area from which they hâve been poorly

studied, providing data for future révisions.

MATERIALS AND METHODS

A detailed list of thc matcrial examined follows

each species description. Specimens largely

belong to muséum collections or were collected

especially for this study. Thcse collections are

deposited in different institutions Ii.sted below

(with other abbreviations) in Brazil and France.

Several lots were obtained in following projccts:

(1) Marion-Dufresne Expédition “MD55”;

(2) Integrated projcct “Coastal Rational Utiliza-

tion of the Brazilian Tropical Région: Sâo Paulo

State”, Instituto Oceanogrâfico da Universidade

de Sâo Paulo (IOUSP); (3) “Oceanic Environ -

mental Monitoring of the Campos Bay", IOUSP,

GEOMAP. EUNDESPA.

The specimens were dissectcd by standard tech¬

niques, under a stcrcomicroscope. with the speci¬

mens immersed under water. The specimens

were extracted from their shells by means of

décalcification or the shells were broken. Serial

sections of anterior régions were rnade by stan¬

dard histological techniques, and stained with

Mallory's triple stain. Some structures were also

cleared in créosote after déhydration in an étha¬

nol sériés and stained in carminé Anatomical

terminology follows Marcus &T Marcus (I960)

and for the foregut, Taylor et al. ( 1993).

Abbreviations used in the figures

aa anterior aorta

ac anterior extremity of crenidial vein

ag albumen gland

al anal gland

an anus

ap accessory proboscis structure

as accessory salivary gland

at duct of accessory salivary gland

au auricle

az anal papilla

bd vetiom bulb duct

bm buccal mass

bs blood sinus

bt buccal tube

ca capsule gland aperture

cc chamber anterior to glandular tissue of

capsule gland

cl furrow formed by columellar fold

cg capsule gland

ci transverse muscle fibres

cm columellar muscle

cv ctenidial vein

dd duct digestive gland

dg digestive gland

ey eye

fl foot latéral furrow

fs foot sole

f't foot

g' g 111

gp gonopericardial duct

he haemocoelic cavity

hg hypobranchial gland

ZOOSYSTEMA • 1999 ■ 21 (2)

201

Simone L. R. L.

ia rhynchudeal introvert distal aperture or

rhynchostome

ib inner wal! of buccal tube

ig Lngesting gland

il inner layer of muscles of venom bulb

in intestine

ip insertion of introvert in foot muscles

kd dorsal lobe of kidney

ki kidney massive tissue

km membrane between kidney and palliai

cavity

ks ventral septatc lobe of kidney attached to

intestine

lf longitudinal muscle fibres

ml muscle unitiug oesophagus with dorsal

wall ofhaemocoel

m3 muscle connecting oesophagus with pos-

terior extremily of rhynchodeal wall

mb mantle border

mn mantle notch

mo ntouth

mt muscular tissue

ne nephrostome

ng nephridial gland

nr nerve ring

nv nerve

ob outer wall of buccal tube

oe oesophagus

oep oesophageal pouch

ol outer laver of muscles of venom gland

op operculum

os osp h radium

ov oviduct

pb proboscis

pc pericardium

pd pénis duct

pe pénis

pf peda! gland

pg nnterior furrow of pedal glands

ph petiis distal chambcr

pm muscles in pénis base

pp pénis papilla

ps palliai sperm duct

pt prostate

rc receptaculum seminis

te rectal septum

ri rhynchodeal introvert

rm retractor muscle of proboscis

ro rhynchodeal introvert proximal aperture

rs radular sac

rt rectum

rw rhynchodeal wall

sd salivary duct

sg salivary gland

si siphon

sn snout

sp siphon projections

st stomach

te cephalic tentacles

tg tégument

tl terminal pouch-like pad of male

tp terminal pouch

ts testis-seminal vesicle

tt radular tooth

vb venom bulb

vd viscéral vas deferens

vc ventricle

vg venom gland

Abrreviations of institutions

IBUFRJ fnstituto de Biologia, Universidade

Fédéral do Rio de Janeiro, Brazil

MNHN Muséum national d’Histoire naturelle,

Paris, France

MNRJ Museu Nacionai da Universidade

Fédéral do Rio de Janeiro, Brazil

MORG Museu Occanogràfico da Fundaçâo

Universidade de Rio Grande, Brazil

MZSP Museu de Zoologia da Universidade

de Sâo Paulo, Brazil

Genus Hastula H. & A. Adams, 1853

Hastula cinerea (Born, 1778)

(Figs 1 A, B; 2; 3A; 4; 5; 6A, B; 7B; 8B)

Sy nonymy: cf. Matrhews et al. (1975: 98) and

Bratcher & Cernohorsky (1987: 191).

Others: Hastula cinerea - Rios 1985: 131

(fig. 590). - Calvo 1987 (fig. 160). - Rios 1994: 181

(fig. 841).

Mat TRIAI. IXAMINRD. AJagoas. Brazil, Maceiô,

Riacho Dote, 16.XII.1973, Menezes coll., 19 spéci¬

mens (MZSP 25069).

Rio de Janeiro. Brazil, Cabo Frio, Conchas Beach,

6.1.1973, Marini coll., 6 specimens (MZSP 24673).

Sâo Paulo. Brazil, Uhatuba, Tahatinga Beach,

8.1.1972, Marini coll., 23 specimens (MZSP

202

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 1. — Shells; A, Hastula cinerea, normally pigmented Shell: B, same species, melanic form; C, D, Hastula hastata, two spéci¬

mens in frontal view; E. Terebra spirosulcata Simone & P. M. Costa n. sp., dorsal view of paratype MZSP 25213; F, same, frontal

view; G, Terebra gemmulata , frontal view; H, same, dorsal view. Scale bars: A, C, D, F-H, 5 mm; E, B, 10 mm.

ZOOSYSTEMA • 1999 • 21 (2)

203

Simone L. R. L.

Fig. 2. — Hastula cinerea, anatorny; A, 6 head-foot, mantle removed; B, mantle organs and anterior région of viscéral mass,

ventral-interior view; C. detail ot 2A. tentacle région; D, transverse section ot middle région ol palliai cavity roof; E, ï head-foot with

protracted introvert, mantle removed; F. anterior région of viscéral mass with kidney and pericardium dissected and with interior

structures exposed. Scale bars: 0.5 mm.

25083). — Sâo Sebasriào, Baraqueçaba Beach,

13.VII.1996, Simone coll., I specimen observed alive

(MZSP 28599). — 26.X.1996, Simone-coll., 1 spéci¬

men observer! alive (MZSP 28600). — 16.V.1997,

Simone coll., 4 specimens observed alive (MZSP

28605). — Guaruja, 20.1X.1968, Costa coll., 42 spe¬

cimens (MZSP 25189).

Distribution. — From Florida, USA, to Santa

Catarina, Brazil.

204

ZOOSYSTEMA ■ 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 3. — Opercula in SEM (outer view except those indicated); A, Hastula cinerea\ B, H. hastata ; C, same species, internai view;

D, Terebra gemmulata-, E, same species, internai view; F. T. crassireticula', G. T brasHiensis', H. T. spirbsulcata. Scale bars: A, G,

0.2 mm; B E. H, 0.5 mm; F, 1 mm.

HABITAI’. — lntertidal, sandy bottoms, beaches with

waves of medium to high energy. Migrâtes during the

tides using the extended foot to be carried shorewards

by the wave and the sharp shell to anchor when the

waves retrear.

Description

Shell( Fig. IA, B)

A description ot the shell can be found elsewhere

(Marcus & Marcus, 1960: 27-28: Matthews

et ai, 1975: 98-99, figs 29-31; Bratcher &

Cernohorsky, 1987: 191-192, figs 233a-i).

Melanic forms, i.e., with homogeneous dark

brown shell, occur at a low frequency (Fig. 1 B).

Head-foot (Fig. 2A, C, E)

Colour homogeneous créant, with pale gray spots

on anterior structures. Head weakly differendat-

ed from head-foot axis (Fig. 2A, E). Tentacles

ZOOSYSTEMA • 1999 • 21 (2)

205

Simone L. R. L.

Fig. 4. — Haslula cinerea, anatomy. A. palliai oviduct, ventral view, including a transverse section of the anterior third; B. pénis, ven¬

tral view. Scale bars: 1 mm.

very short, almost vestigial. Eyes very small, dark,

located süghtly below tentacle base (Fig. 2C).

Basal, proximal introvert aperture rather broad,

transver.se, located arttciiorly and ventral to ten-

tacles. Font large, occupying more tban hall a

whorl, without clear divisions; sole with loidcd

borders Cnltimell.ir muscle of about 1.3 whorl,

rather thin, Other details in Marcus & Marcus

(1960:28-30).

Operculum (Figs 2A, E; 3A)

Minute, unguiculate, pale brown, located in

middle région oF posterodorsal foot surface,

nucléus terminal (Figs 2A, E; 3A). Occupies

small part o( aperture.

Mande organs (Figs 2B, D; 4A)

Mande border simple, nor pigmenred. Siphon

well-developed, pale cream, with borders cntirely

edged bv small lobed papillae (Fig, 2B). Mande

cavity ex tends about two whorls. Ü.sphradium

bipectinate, elliptical, about one third of gill

length; with several filaments unitorm on horh

sides of osphradial ganglion; right filaments

similar in size to left filaments; each filament

scalloped by about four decreasing digitations

(Fig. 2D). Gill rather elliptical and long, about

four fîfths of palliai cavity length; anterior end

radier far from mantle border; in anterior extre-

mitv only ctenidial vein présent, in form of a

small seprunt (Fig. 2B: ac); filaments begin gra-

dually at sorte distance from anterior end; each

filament trianguJar, apex varies from central to

located on the right, right margin varies from

almost straight to convex; gill postenor extremity

ver)' dose to pericardium. Ctenidial vein narrow

and uniform ail along its lengrh, excepi for its

broader anterior extremity. A proportionally

broad space berween gill and rectum. Hypo-

branchial gland thin. located in postenor half of

cavity, at left of rectum, pale cream. Anal gland

with snmc sleiidcr, irregular acina immersed in

right margin ol hypohranchial gland close to rec¬

tum (Fig. 4A), but nut close to anus, purple.

Palliai gonoduers rim along right margin ol post-

erior half ol palliai cavity. Rectum narrow, lying

ventral io.and to Idt ol palliai gonoduers. Anus

close to anterior extremity of palliai oviduct in

206

ZOOSYSTEMA * 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 5. — Haslula cinerea, anatomy: A, foregut removed from nead-toot, dorsal view: B, same. ventral view: C, rhynchodeal wall,

proboscis and buccal tube partially opened longitudinally, ventral view: D, detail of apical extremity of proboscis opened longitudinal-

ly, showing gripping of tooth: E, buccal mass and oesophagus opened longitudinally, internai surface exposed; F, stomach, ventral

view, digestive gland partially removed. Scale bars: 1 mm.

ZOOSYSTEMA • 1999 • 21 (2)

207

Simone L. R. L.

Fig. 6. — Radular teeth, SEM; A B, Hastula cinerea:

C. Terebra crassirelicula ; D. E, H. haslata ; F, G. T. taurins-,

H, I, T. spirosulcata. Scale bars: A. B, F, G, 100 (jm; C, 40 pm;

D, E, 20 |jm: H. I. 50 pim.

females or prostate in males; bearing a small,

ventral papilla on its border (Figs 2B, 4A).

Anterior half of palliai right margin without

inner structures. Other details in Marcus &

Marcus (1960: 30-32).

Circulatory and excrecory Systems (Fig. 2B, F)

Heart of medium si/.c, locatcd in left région of

posterior limir of palliai cavity. Auricle just

behind end of gill and anterior to ventricle.

Aortas run along anterior surlace of digestive

gland. Kidney occupies almost half a whorl, flat-

rcncd, located in right région ol posterior liinit of

palliai cavity. Kidney with two main glandular

masses, separated by a flatcened chamber

(Fig. 2F). Dorsal glandular mass rhick (kd), clear

brown, bulging into palliai cavity. Ventral glan¬

dular mass (ks) bordering rectum, pale créant,

bearing several. uniform transverse septa.

NephridiaJ gland pale cream, triangular in sec¬

tion, bordering dorsal margin of membrane bet-

ween kidney and pericardial chambers (ng).

Nepbrostome a transverse slit in middle région of

membrane between kidney and palliai cavity.

Other details ru Marcus & Marcus (1960: 33-34).

Digestive System (Figs 2E; 5; 6A, B; 7B; 8B)

Rhvnchodeal introvert occupies about half of

remainder of foregut length, cylindrical (Figs 2E;

5A, B); muscular walls rhick, cumpostd of tvvo

closcly connccted muscular layers. Distal apertu-

re of rhvnchodeal introvert (rhynchostome) a

small longitudinal slit, preccded by a rhick

sphincter. Inner rhynchodeal wall very thin,

transparent (Fig. 5A, B), covering inner surface

of anterior hall of haemocoel, connccted to it by

small muscle fibres mainly located near rhyncho¬

stome. Proboscis conical, ol about sanie length as

rhvnchocoel. musculat and rhick (Fig. 5A-C); its

base Itised with that of rhynchodeal wall. A pair

of large retractor muscles originate in mid-lateral

région of inner surface of haemocoel and are

inserted at rhynchodeal waii-proboscis transition,

with a small part also inserted into buccal mass

(Fig. 5C). Buccal mass spherical, with a long,

broad buccal tube (Fig. 5C). Buccal tube with

two muscular walls, connccted with each other

only at anterior extremity near mouth. Several

small transverse muscle fibres connecting inner

surface of proboscis wall with outer surface of

oLitcr buccal tube, Outer buccal tube wall thick,

bearing mainly transverse muscle fibres. Inner

buccal tube wall thin, semi-transparent, bearing

mainly longitudinal muscle fibres, most of its

inner surface smooth, with a few longitudinal

folds. Anterior extremity of buccal tube, wherc it

grips radular tooth, with a horseshoe-.shaped

inner lold (concavity anLerior), with two post¬

erior, longitudinal, muscular projections attached

to dorsal surface, and a central, short, digitilorm

projection notching radular tooth base (Fig. 5D)

(see also Marcus & Marcus 1960: lig. 7).

Posterior extremity of outer buccal tube wall

contours proboscis base, connccted with it

throughout ils entire circutnfercnce by a thin

layer of muscle fibres (Fig. 5C), but not fused

with each other; buccal tube also inserting in

208

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphoiogy of Brazilian Terebridae

Fig. 7. — Schematîc piclures; A, composite-synoptic diagram of the terebrid foregut based on species studied herein. ventral view,

not to scale or proportion; B G, diagram of foregut in dorsal view of studied species following the model in the current literature, not

to scale or proportion: B, Hastula cinerea, C, H. haslala, Terebra taurina: D, T. brasiliensis ; E, T. spirosu!cata\ F, T. crassireticula,

T. leplapsis; G, T. gemmulala.

retractor muscle pair (rm), and in inner surface terminally in mid-ventral région of buccal mass

of haemocoel. Posterior extremity of inner buccal (Fig. 5F.). Radula composed of about twetity

tube wall fused directly to buccal mass. Pair of single, hollow, marginal teeth; each tooth coni-

retract-or muscles (rm) also inséra in buccal cal, base barbed, tip narrowing gradually, circular

mass latéral surface (Fig. 5C). Radular sac long, in section (but not altogether fused); lenghr

curved, with several radular teeth, opening sub- about 650 pm. Tip sharply pointed, with a

ZOOSYSTEMA • 1999 • 21 (2)

209

Simone L. R. L.

small, narrow orifice (Fig. 6A, B). SalLvary glands

form pair of hemi-spheres ofglandular, whire tis-

sue connccted to each other; cheir ducts contour

oesophagus and insère at anterior and posterior

side of base of radular sac aperture (Fig. 5C, E);

gray, indescent. Venorn gland very long and

convolute, about hall: anterior to and hall poste-

rior to nerve ring, no apparent change in its

inner tissue along its length; inserts in buccal

mass close to and ar left of radular sac aperture.

Muscular bulb radier long, conical, with broad

distal région (Fig. 5A, B); vvall conrposed of two

layers of muscular tissue, inner layer slightly

thinner rhan oufer layer. A sketch ol foregut

structures shown in Figure 7B. Inner surface of

buccal mass with several longitudinal, lovv, sube-

quai folds (Fig. 5F), some of them converge to

aperture ofvenom gland and of radular sac; these

folds become discontinuons at anterior end of

oesophagus, marked by a sudden interruption of

folds and a réduction of their number (Fig. 5Fl).

Oesophagus a long, narrow tube with several

longitudinal folds similar to those of buccal mass

(Fig. 5E), folds tall in some spécimens (Fig. 8B);

no interior glands présent. Stomach simple, cur-

ved (Fig. 5F), loc.ated hajf a whorl posterior to

kidney and immersed in digestive gland. Duct to

digestive gland single, Jocatcd in mid-ventral

région of stomach; a fier a short distance bifitr-

cares into anterior and posterior branches.

Digestive gland with about 5.5 whorls posrerittr

to stomach and hall a whorl anterior to it, close

to pericardium level (Fig. 2B), presscd between

kidney and oesophagus, beige. Intestine narrow,

with thin walls and an almost smootli inner sur¬

face; runs to left ol and ventral to margin of kid¬

ney. Rectum and anus descrihed above. Other

details in Marcus &i Marcus (1960: 37-44).

Génital System (Figs 2A; 4A, B)

Male. Prostate thick-wallcd, of about same ieng-

th as rectum, running ventrally to it, attached to

palliai floor, with a small longitudinal aperture in

its posterior ventral région. Vas deferens, aiter

prostate, runs along floor of right margin of pal¬

liai cavity. Pénis moderatelv long (Fig. 2A), cylin-

drical; pénis duct slightly convoluted, near left

margin ol pénis. Distal end ol pénis with a

concavity turned to right, in its centre a rather

large papilla vvhere pénis duct opens (Fig. 4B).

Pénis papilla close to right margin. Other details

in Marcus & Marcus (1960; 44-45, figs 14, 15).

Female. Palliai gonoduct with a wcll-develaped

terminal pouch (Fig. 4A), with a longitudinal

lurrow turned to right close to columella, occu-

pying about hait total length of palliai gonoduct.

Other details in Marcus & Marcus (1960: 45-47,

figs 16-18).

/Verrous System

As described by Marcus & Marcus (1960: 34-37,

% 5).

Hastula hastata (Gmelin, 1791)

(Figs IC. D; 3B, C; 6D, E; 70 8A; 9-11)

Synonvmy: ef. Matthews et al. (1975: 101) and

B râtelier 8c Cernohorsky (1987: 184).

Othcrs: Hastula hastata — Rios 1985: 131

(fig. 591). Calvo 1987: 171 (fig. 161). - Rîos 1994:

181 (fig. 892).

Ma I h Kl Al. EXAMINE». — Bahia. Brazil, Salvador,

Itapuâ Bcach, 27.11.1997, Simone co11., 2 specimens

observed alive (MZSP 28426).

Rio de Janeiro. Brazil, Büzios, Ferradura Beach,

17.1.1994, Simone coll., 30 specimens (MZSP

28713). — llha Grande Bay, RV Emilia , stn 288,

24 m, 30 . VII. 1966, 1 9 (MZSP 24671). — RV

Emilia , sin 269, 30.2 ni, 19.VI1.1966, 1 9

(MZSP 24669).

MlASUlUAll.Nls. — MZSP 24671: 32.1 x 8.1 mm.

Distribution. — From Florida, USA, to Santa

Catarina, Brazil,

HABITAT. — lnfratidal, crawling on sand or sandy

mud.

Description

Shell { Fig. IC, D)

A description of the shell can be found in

Matthews et al. 1975: 101 (figs 35-37);

Bouclier &c Cernohorsky, 1987: 184 (fig. 222a,

b). The shell bears two spiral colour bands on

each whorl, pale brown and beige, some speci¬

mens cntirely beige. Periostracum rhm, opaque,

black. Ourlinc rather short and cylindrical, last

few whorls being of about the same diameter.

Sculpture of well-developed axial ridges.

210

ZOOSYSTEMA - 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Head-foot{ Fig. 9 A, D)

Colour homogeneous cream. Head Iittle differ-

entiated from hcad-foot axis (Fig. 9A). Tentacles

short, poiutcd, directed laterally (Fig. 9D). Eyes

small, dark, locatcd at tentacle bases (Fig. 9D).

Basal, proximal introvcrt apcrcure rather broad,

transverse, anterior and ventral to ren racles. Foot

large, occupytng more than half a whorl, wtth a

shallow furrow bctwecn mcsopodium and epipo-

dium (Fig. 9A); sole wi’tb folded borders.

Columellar muscle occupying about two whorls,

rather thin. Males with pénis in central-right

région posterior to head.

Opèreulum (Fig. 3B, C)

Large, unguiculate, pale brown, nucléus terminal

(Figs 3B, C). Occupying entire shell aperture.

lnner surlace with a large oblique scar near inner

and posterior margins (Fig. 3C).

Mantle or pans (Figs 9B, C; 10)

Mantle border simple, not pigmented. Siphon

well-developed, long, pale cream, with smooth

borders; both sides ol siphon base with well-

developed siphon appendices, standing out Irom

remainder of border (Fig. 9B: sp), sometimes

both extend through shell aperture. Mantle cavi-

ty occupying about two whorls. Osphradium

elliptical, about une rhird of gill length; with

several uniform filaments on both sides of

osphradial ganglion; right filaments similar in

.size to left filaments; each filament scalloped by

about threc digitations (Fig, 9C). (iill long and

narrow, extending about nine tenths of palliai

cavity length; anterior extremity close to mantle

border, with only a short portion of ctenidial

vein présent, in form of a very small septum

(Fig. 9EL ac); filaments hegin graduai ly a short

distance from anterior extremiry; each filament

triangular, apex located at rigln, right margin

almost straight; gill posterior extremity very close

to pericardium, Ctenidial vein uniformly narrow,

except for a broader anterior extremity. A pm-

portionally broad spacc bctwecn gill and rectum.

Hypobranchial gland thin, locatcd in posterior

two thirds of cavity, at lefi ol rectum, pale cream.

Anal gland with some slcnder, irrcgular acina

immersod in right margin ol hypobranchial

gland in anterior half of rectum, distant from

anus (Figs 9B; 10), purple. Palliai gonoducts run

along right margin of posterior two thirds ol pal¬

liai cavity, attached also to palliai floor. Recrum

narrow, lyîng dorsally to prostate in males,

ventrally and to left région of palliai oviducts in

féru a les; in females, before anus, rectum crosses

to dorsum and remains attachée! to mantle, Anus

chose to anterior extremity of palliai oviduct in

females (Fig. 10) or prostate in males (Fig. 9B);

bearing a small, ventral papilla at its border.

Anterior third oi palliai right margin without

inner structures.

Circidatory and cxcretory Systems (Fig. 9F)

Heart of medium size, similar to that of prece-

ding species. Kidney of almost one quarfer

whorl, flattened, located in right région of post¬

erior limit of palliai cavity. Kidney apparendy

without large inner chambers and free from rec¬

tum; internally with only a flattened chamber

and a large, massive glandular mass, triangular in

section, pale cream. Nephridial gland small, pale

cream, section triangular, bordering dorsal mar¬

gin ol membrane bctwecn kidney and pcricardial

chambers. Ncphrostomc a transverse slit in

middlc région ol membrane bctwecn kidney and

palliai atvity.

Digestive sys terri

(Figs 6D, E; T C; 8A; 9A, E, F; 1 IB)

Organizafion rather similar to that of prcccdiug

species. Rhynchodeal introvert occupying about

half o( remainder ol foregut length, cylindrical

(Figs 9A; 11 B); walls with thick muscles, double.

Rhynchodeal introvert distal aperture (rhyncho-

stome) large; with a thin sphincter, Inner rhyn¬

chodeal wall very thin, transparent, covering

inner surface of anterior half ol hacmocoel as in

preceding species (Fig. 1 IB). Prohoscis cortical,

ol about sanie length as rhynchodeal, connccted

at its base to rhynchodeal wall and rétracter

muscles; prohoscis distal région very narrow and

long, gcncrallv preserved part protracted; a mas-

cular thickcning grips a tooth close ro prohoscis

tip. Buccal mass cylindrical and elongated, with a

long, broad buccal tube that also bccomes very

narrow distal ly likt prohoscis. Buccal tube with

outer muscnlar wall, consisring mainly ol longi¬

tudinal muscle fibres; inner wall thin, with seve-

211

ZOOSYSTEMA - 1999 • 21 (2)

Simone L. R. L.

Fia. 8. — Interasting aspects of serial sections, semi-diagrammatic représentations; A, Haslula hastata, transverse section a! the

basal level of the proboscis, tiead tégument removed; B. H. cinerea transverse section ot anterior région af oesophagus showing

internai folds; C, Terebra gemmulata, antero-postenor section in middle région of right tentacle, just at the level ol the eye. anterior-

dorsal extramlty ol lùregut also shown; D. E, T■ spirosulcata, transverse sections of venom gland. D. proximal (anterior to nerve

ring), E, distal région: F. T. brasiliensis, accessory proboscis structure, transverse section ol middle région: G, 7 taurina, proximal

(anterior) extremity of venom bulb and distal extremity ot venom gland sectioned just at their duct level. Thickness of section: 7 pm,

Mallory stain, Scale bars: 100 pm.

212

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology oi Brazilian Terebridae

Fig. 9. — Haslula hastata, anatomy; A, 6 head-foot, mantle removed, inlrovert protracted; B, manlle organs, ventral-internal view;

C, transverse section ot rmddle région of palliai roof; D. detail of A in région ol tentacles; E. transverse section of viscéral mass hait a

whorl posterior to the stomacn; F, viscéral mass anterior région, ventral view, digestive gland near stomach partially removed, kidney

and pericardluin opened. Scale dais. A, 1 mm, B-F. 0.5 mm

ral longitudinal, internai, tall folds (Fig. 8A).

Anterior extremity of buccal tube smooth intern¬

ai ly. A pair of retractor muscles (rm) is also insert-

ed in latéral surface of buccal mass. Two acces-

sory salivary glands, one on each side of probos-

cis base, close to insertion of retractor muscles

(Fig. 11 B: as); each gland elliptical, long, occupy-

ing about one quarter of length of rhynchodeal

ZOOSYSTEMA • 1999 • 21 (2)

213

Simone L. R. L.

Fig. 10. — Hastula hastata palliai oviduct, ventral view, pari of its adjacent régions of mantle and viscéral mass also shown. Scale

bars: 1 mm.

cavity length; their ducts run in outer wall of

buccal tube (Fig. 8A), fusing witb cach amer in

middle level rtf proboscîs. iladular sac ver)' small,

elongated, curvtd, with about ivvcnry radular

teeth, opening sub-terininally in ventrtd-mid

région of buccal mass (Fig. 11 B). Radula with

very small teeth, about 120 pin in length; each

tooth wirh two distinct régions (Fig. 6D, E):

basal région broader than disral one, hollow, base

barbed, circular in section (but not altogether

fused); distal région about same length as basal

one, narrnw, almost solid except for a longitudi¬

nal apertttre in its proximal région, tip sharply

pointed. Salivary glands similar tu tltose of prece-

ding spccies. Venom gland ver)' long, narrow and

convolutc (Fig. Il B), about Fiai f anterior and

half posterior to nerve ring; without apparent

change of inrter tissue along its length; inserted

in buccal mass close and ar left of radular sac

aperture, Muscular bulb elliptical, short, almost

circulai' in section; with two muscular layers,

inner layer about half thickness of outer laver.

Inner surface of buccal mass similar to that of

preccding spccies. A sketch of foregut structures

is shown in Figure 7C, E.

Oesophagus a long, narrow tube, anterior région

flattened and with smooth inner surface, poste¬

rior région narrow with several longitudinal folds;

no interior glands. Stomach simple, curved,

located half a whorl posterior to kidney, irnmer-

sed in digestive gland, almost restricted ro left

half of this région (Fig. S>F). Duct to diges-tive

gland single, locatcd in mid-ventral région of sto¬

mach. Digestive gland (Fig, 9F, F) occupying

a bout 5.b vvhorls posterior to stomach and half a

whorl anterior to it, beige. Intestine narrow, with

thin walls, inner surface almost smooth, sigmoid;

rttns left and ventral to kidney (Fig. 9F).

Génital System (Figs 9A, E, F; 11 A, C, D)

Male. I estis (Figs 9F; 11A) rather short, located

in peri-columellar région of second whorl poste¬

rior ro stomach, gradually narrowing and heco-

ming a thick wallcd duct which also runs close to

cohtmclla (Fig. 9F), opening to palliai cavity near

and ventral to rectum. Prostate thick-walled,

rather narrow and long, close about half length

of rectum, without détectable aperture. Vas défe¬

rais runs along floor of right margin of palliai

cavity; in its anterior région it is immersed in

head tégument and émerges into hacmocoel a

short distance before its entrante to pénis

(Fig. 11 D); în this area vas déferais thîcker.

Inner région of pénis base stronglv muscular

(Fig. 1 I D), a flattened muscle crosses through

foregut and insertS in ventral surface of haemo-

coel. Sometimes these pénis muscles di.slocate

foregut structures to left. Pénis rather short

214

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology oi Brazilian Terebridae

Fig. 11. — Hastula hastata, anatomy: A, viscéral mass posterior to stomach, partially uncoiled. ventral view: B. foregut, ventral view,

rhynchodeal wall and introvert opened longitudinally, proboscis partially opened longitudinally, nerve ring removed (its position indi-

cated): C. pénis, dorsal view, with a detail ol interior surface of the right wall of Its chamber 0, internai surface of dorsal wall of hae-

mocoel, just in région of pénis, showing pénis duct and muscles. Scale bars: 1 mm

(Fig. 9A) — about hall Icngth of palliai cavity, (Fig. llC). Distal end ol pénis with a concavity

flattened; basal half broadcr ch an distal liait", strongly turned to right, with a rather large

which tapers gradually; pénis duct very narrow, papilla in its centre through which pénis duct

slightly cotivoluted near left margin of pénis opens (Fig. 11C). Right région of this concavity

ZOOSYSTEMA • 1999 • 21 (2)

215

Simone L. R. L.

very deep - about one third of pénis length, pro-

ducing a long chamber with right margin bea-

ring a pair of low longitudinal folds (Fig. 11C:

ph). Pénis papilla doser to right margin of pénis

than to left.

Female, (Fig. 10) Ovary similar in position to

testis. Viscéral oviduct very narrow, running

close to columella, with a small gono-pericardial

aperture; inserted in palliai gonoduct anterior to

ingesting gland. Ingesting gland hcmispherical,

orange, with a short rcceptaculum scminis

connecting it to oviduct. Albumen gland short,

anterior ro ingesting gland. Capsule gland long-

about half of total palliai oviduct length, cylin-

drical, with a central, (lattened duct which opens

as a small sût at right of terminal pouch base.

Terminal pouch elliptical, with a longitudinal

furrow curned to right close to columella and

opening anteriorly; occupying about one third of

total length of palliai gonoduct.

Genus Terebra Bruguière, 1789

Terebra branliensis Smith, 1873

(Figs 3G; 7D; 8F; 12H; 13B; 14E; 15; 16)

Synonyme: cf. Matthews et al. (1975: 91) and

Bratcher & Cernohorsky (1987: 160).

Others: Terebra brusittensis — Rios 1985: 130

(fig. 583). 1994: 179 (fig. 832).

Maferial iXAMINtn. — Rio de Janeiro. Brazil, RV

W. Besnar/l, stn 1475, 23°08’S, 43°46’W, off Rio de

Janeiro, 40 m, 8.111.1971: 1 d, 1 9,3 shells

(MZSP 19403).

Mf.ASUREMENTS. — ln millimeters MZSP 19403:

9 11.0X3.3; <? 9.0 x 2.8.

Distribution. — Rio de Janeiro, Brazil.

HABITAT. — From 20 to 40 m deep. Sandy sub¬

strates.

Description

Shell {Figs 12F.; 13B; 14E)

Shell description fôund in Matthews et al. , 1975:

91-92 ( Figs 9-13); Bratcher & Cernohorsky,

1987: 160 (ligs 186a, b). Sculpture lacking

except for axial subsiltural ribs, giving an almost

smooth surface (Figs- 12E; 13B). In comparison

with other terebrids, T. brasiliensis has few teleo-

conch whoris and a proportionally large proto-

conch (Fig. 14F), giving an appearance ot intma-

turity, but ail specimens are rather similar and

hâve génital glands developed.

llmd-foot (Figs 15B, G; 16A)

Colour homogeneotis yellowîsh cream. Head

weakly differcntiated from head-foot axis

(Fig. 15B). lentacles very short, dorso-ventrally

flattened, tip rounded (Figs I5B, C; 16A). Eyes

small, dark, located in middle of lentacles. Basal,

proximal introvert aperture rather broad, trans-

verst-, located anteriorly and ventrally to ten-

tacles. Four occupying almost halfwhorl; sole

with lolded borders; a shallow furrow scparates

mesopodium from metapodium; furrow of pedal

glands anterior (Fig. 15B). Columellar muscle of

about two whoris, rather thick. Males with pénis

of medium si/.e, originaling in central-right

région posterior to head.

Uperculum (Figs 3G; I5B)

Large, unguiculate, pale brown, nucléus terminal

(Fig. 3G). Occupying entire shell aperture

(Fig. 15 B).

Mande urgam (Figs 15A, D, I ; 16D)

Mande border simple, not pigmented. Siphon

well-developed, ycllowisb cream, with smooth

borders. Mande cavtty of about two whoris

(Fig. 15A). Osphradium elliptical, long, about

rwo tliirds of gill length, with several, unilorm

filaments; right filaments largc-r than left, angu-

lar, more numerous than left filaments

(Fig. 15D); a short portion of anterior left région

of osphradium with filaments lacking (Fig. 15A).

Gill narrow and long, about two thirds of palliai

cavity length; anterior end rather far from (nantie

border, with onlv a short portion of ctenidia!

vein présent, in form of a very small septum

(Fig. 15A: ac); filaments hegin graduallv at sonie

distance from anterior end: cach filament trian-

gular, low, apex about central, margïns almost

straight; gill posterior end far from pericardium.

Cfenidial vein narrow and uniform ail along irs

length, excepr for ils hroader anterior extremity;

a long posterior région Iree from filaments

(Fig. 15A, F). A proportionally narrow space bct-

ween gill and rectum. Hypobranchia! gland thin,

216

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 12. — Shells in frontal view, SEM (only in E the shell is coated with gold); A, Terebra crassireticula n. nom., MNHN; B, Terebra

leplapsis n. sp., holotype; C, same species, paratype; D. Terebra sterigma n. sp., holotype; E, Terebra brasiliensis. Scale bars:

2 mm.

located in posterior half of cavity, ar left of rec¬

tum, pale cream. Anal gland not différentiable.

Palliai gonoducts run along right margin of pos-

terior half of palliai caviry. Recuint narrow, lying

ventral and to left of palliai gonoducts, after-

wards attached to mande (Figs 15A; 16D). Anus

in front of anterior extremity of palliai oviduct in

females or prostate in males; bears no papilla.

Anterior half of palliai right margin without

internai structures.

Circulatory and excretory Systems (Fig. 15E, F)

Heart of medium size, position similat to that of

preceding species. Kidney of almost one quarter

whorl, flattened, located at right of posterior

limit of palliai cavity; without inner chambers,

consisting of a mass of whitish tissue with trans¬

ZOOSYSTEMA • 1999 • 21 (2)

verse furrows; furrows more évident in posterior

région where rectum borders. Nephcidial gland

small, pale cream, triangular in section, borde-

ring dorsal margin of membrane between kidney

and pcricardial chambers. Nephrostome a trans¬

verse slit in rniddle région of membrane between

kidney and palliai cavity'.

Digestive system (Figs 7D; 8F; 15E; 16A-C)

Rbynchodea) introvert large, conical, cylindrical

length almost equal to rhynchococl léngth, mus-

cular walls rhick (Fig. 16A, B). Outer région (in

retracted condition) mainiy of longitudinal fibres

and inner région mainiy of circular fibres; both

muscular layers closely connected to each other

by connective tissue, without any space between.

Distal aperture of rhynchodeal introvert (rhyn-

217 I

Simone L. R. L.

Fig. 13. — Details of shells in SEM; A, detail of shell sculpture at the levai of the penultimate whorl of T. gemmulata: B, same of

T. brasiliensis: C, same of T. spirosulcata, D, f. taurina, columella of penultimate whorl exposed; E, detail of aperture of T. crassireti-

cula ; F, same al 7. leprapsis: G, same ol 7. slerigina. Scale bars: 0.5 mm.

chostomc) very large andwith a latéral expansion parent, covering inner surface of anterior half of

on each side (Fig. 16B); a very large sphincter haemocoel as in preceding species (Fig. 16A-C).

présent. Inner rhynchodeal wall very thin, trans- Accessory proboscis structure large - about same

218

ZOOSYSTEMA • 1S99 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 14. — Shells (B-H in SEM): A. Terebra taurina ; B, detail of protoconch of T. crassireticula ; C, same of T. leptapsis ; D, same of

T. sterigma : E, same of 7. brasiliensis ; F, detail of Shell sculpture at the level of the penultimate whorl of T. crassireticula ; G, same of

T. leptapsis; H. same of T. sterigma. Scale bars: A, 10 mm; B-D, 0.2 mm; E, 0.1 mm; F-H, 0.5 mm.

ZOOSYSTEMA • 1999 • 21 (2) 219

Simone L. R. L.

Fig. 15. — Terebra braslliensis, anatomy, A, palliai organs, ventral-internal view; B, head-foot of male, mantle removed; C. same,

detail of tentade région; D. palliai cavity roof, transverse section of ils middle région; E. anterior région of viscéral mass and adjacent

palliai cavity, ventral view, part ol digestive gland adjacent to stomach removed, pericardium and kidney partially opened; F. whorls

adjacent to posterior limit oi palliai cavity, ventral view, Scale bars; 0 5 mm,

length as rhynchococl, flattencd, originating in

middle-lefr région of rhynchodcal wall inner sur¬

face, in proximal région vcry thin, gradually

becoming tliick and broader (Fig. 16B, C), tape-

ring suddenly at tip. Central région of accessory

proboscis structure muscular and peripheral

région glandular (Fig. 8F). Proboscis very small,

reduced, présent only as muscular ring around

buccal mass (Fig. 16B, C). Buccal mass spherical.

Pair of rctractor muscles reduced. Radular sac

and salivary glands' not présent. Venom gland

and muscular bulb also absent. A sketch of fore-

220

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 16. — Terebra brasiliensis , anatomy; A, head and haemocoel, ventral view, foot and columellar muscle removed; B. same. with

ventral région o! rhynchodeal Wall removed, introvert and nerve ring reflected to show dorsal surface: C. poslerior extremily ol rhyn-

chodeal wall and buccal mass both opened tongiludinally, internai surface exposed; D, palliai oviduct and adjacent rectum, ventral

view; E, pénis, dorsal view. Scale bars: 0.5 mm.

gut structures shown in Figure 7D. Inner surface tive gland. Duct to digestive gland single, located

of buccal mass and oesophagus smooth in middle, ventral région of stomach. Digestive

(Fig. 16C). Oesophagus long and narrow. gland of about 3.5 vvhorls posterior to stomach

Stomach simple, curved, located half whorl pos- and also half a whorl anterior to it, beige,

terior to kidney (Fig. 15E), immersed in diges- Intestine broad, slightly sinuous, with thin walls,

ZOOSYSTEMA • 1999 • 21 (Z)

221

Simone L. R. L.

inner surface almost smooth; runs along left and

ventral margins of kidney (Fig. 15E). Rectum

and anus described above.

Génital systetn (Figs 15B, F; 16D, E)

Male. 1 es ci s similar to tltat of preceding species.

Viscéral vas déférons runs close to columclla;

about one whorl before kidney becoming very

broad, wich thick, i ri discent walls and rather coîl-

ed; narruvving at Içvcl of kidney (Fig. 15F), exit-

ing to palliai cavity in middle région of posterior

limit of cavity, running alcmg floor of palliai cavt-

ty within tégument (Fig. 15B). Prostate not dif¬

férentiable. Pénis about half of palliai cavity

length, flattened (Fig. 15B); rather uniform in

width; pénis duct very narrow, ncarlv straight,

near left margin of pénis (Fig. 16E). Pénis distal

end with a deep concavity turned to right and

with a simili aperture; in its centre a rather large

papilla where pénis duel opeJis.

Female. Ovary siniilar in position to testis.

Viscéral oviduct very narrow, running close to

columella, inserted in palliai oviduct ai right pos¬

terior extretnity ot albumen gland (Fig. 16D).

Ingesting gland rather sphericah posterior to

albumen gland, with a very short receptaculum

seminis connected dîrecrly with palliai oviduct in

left margin. Albumen gland short, thick-walled,

whire; limit between it and capsule gland impré¬

cise. Capsule gland ycllow, about half of total

palliai oviduct length. cylindrical, with a central,

flattened duct (Fig. 16D); this duct narrows

abruptly in riglu posterior extretnity of capsule

gland, Crossing right posterior région of terminal

pouch, bearing several narrow longitudinal folds,

opening as a snvall pore located in about mid-

ventral région of terminal pouch. Terminal

pouch broad, cylindrical, with a snrall terminal

anterior aperture, close to columella; terminal

pouch comprises a deep, blind sac (Fig. 16D);

left lip of aperture svith a long fold which gra-

dually disappears; total length of terminal pouch

about half that of palliai oviduct.

Terebra crassireticula n. nom.

(Figs 3F; 6C; 7F; 12A; 13E; 14B, F; 17)

Terebra reticulata Simone & Verissimo, 1995: 460-

466, figs 1-8 (pre-occupied name) (non Sowerby 1840).

IAti MA'L tkiAL. — SE Brazil. RV Manon-Dufresne,

stn DC75 18°59’S. 37°50AV, 295 m, V.1987. 1 shell

MN11N. — Stn CB92, L9°34'S, 38 < ’55’W,

340-360 m, V.1987, 160 shells MNHN. —

Stn CB96, 21°32’S, 40"09AXA 295 m, V.1987,

46 sliells MN'HN. —Stn CB9S, 2I°35’S, 40°31’W,

900 in, V.1987, 1 erodeci shell MNHN. —

Stn cm 03, 23°36‘S. 42°02‘W, 200-217 m, V.1987,

1 shell MNHN. Stn CB104 . 23"42'S, 42'’07’W,

430 450 m, V.1987, 1 shell MNF IN.

Sâo Paulo. Brazil. off llha BeU, RV W. Besnard,

23°57.3’S. 44 ü 53AV. 75 m. 27.VII.1986, 2 specimens

(MZSP 28393).

DISTRIBUTION. — From Rio de Janeiro to Sâo Paulo,

Brazil.

Habitat, — Deep waters, from 75 to 900 m depth

(commoncst between 200 and 450 m depth), sandy

siibst rates.

Ml ASt'RtMtNTs. — In millimerers, followed by

number of axial and spiral ribs in penultimate whorl.

MNHN srn CB96: 15.5 X 3.9, 19, 3; 17.4 x 3.7,

19, 4.

Remarks

Males and mature females were not examined in

the original description. With additional fixed

material provided from a new collection and a

loan from MNHN the following complement-

ary morphological description is possible.

COMI’l l:MENTARY OESCRINTON

Shell (Figs 1ZA; 13E; I4B. F)

Shell description can be found in Simone &

Verissimo (1995: 462-463. figs 1-3).

Head-foot and operculum (Figs 3F; 17A, C)

As described by Simone & Verissimo (1995:

463, figs 4-5).

Mande organs (Fig. 17B, E, F)

Description found in Simone & Verissimo

(1995; 464, lig. 6) particularly accurate, follow¬

ing data additional: anus located near middle

région of right margin of palliai cavity, anterior

one third of palliai riglu margin wilhout internai

structures (Fig. 17E); gill filaments triangular

with tip about central (Fig. 17B); oSph radium

wirh left filaments smaller and fewer rhan right

filaments.

222

ZOOSYSTEMA ■ 1999 ■ 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 17. — Terebra crassireticula n. nom., anatomy; A, S head-foot, mantle removed: B, palliai cavîty roof, transverse section ot its

middle région; C head and toregut removed, ventral view, rhynchodeal wall partially opened with sortie internai structures expos-ed;

D, pénis and adjacent area of head-foot. dorsal view: E, palliai oviduct and adjacent structures, ventral view: F, part ot viscéral mass

and a short portion ol adjacent palliai cavîty, part of digestive gland adjacent to stomach removed, Kidney and pericardlum opened,

internai surface exposod Scale bars: 0,5 mm

Circula tory and cxvretory Systems (l : tg. 17F) almost one third wliorl, flattcncd, loc.ued in

Hcart of proporlionaly small si œ, similar in posi- right région ol posterior limit of palliai cavity.

tion to that of preceding species. Kidney of Kidney with flattened inner chamber separating

ZOOSYSTEMA • 1999 • 21 (2)

223

Simone L. R. L.

two glandular masses. Dorsal glandular mass

thin, white, with several, irregular glandular

acina. Ventral right mass border attached to rec¬

tum, very thin, with several transversc folds.

Nephridial gland small, pale crcam, section trian-

gular, bordering dorsal margin ot membrane bet-

ween kidney and pericardial chambers.

Nephrostotne a transverse slit in middle région of

membrane between kidney and palliai cavity.

Digestive System (Figs 6C; 7F; 17C, F)

Rhynchodeal introvert occupies about half of

remainder of foregut length. cylindrical

(Fig. 17C), double walls thick and muscular.

Distal aperture of rhynchodeal introvert (rhyncho-

stome) large, précédée) by a moderately rhick-ened

sphincter. Rhynchodeal wall very thin, transpa¬

rent, covers inner surface of anterior half of hae-

mocoelas as does that of preceding species

(Fig. 17C). Accessory probuscis structure small,

white, long, connected to rhynchodeal wall in

middle région of its left side and attached to inner

surface of haemocoel by small ligaments

(Fig. 17C: ap); inner région muscular, outer

région glandular and irregular. Praboscis conical,

clearly shorter rhan rhynchocoel, connected atits

posterior limit to rhynchodeal wall and retractor

muscle; gripping tooth at its tip, similar to

H. cinevea. Buccal mass spherical, with inner and

outer organization similar to that described for

H. cinereil. Radula (Fig. 60 described by

Simone &C Verissimo (1995: 464, fig. 8), each

tooth about 280 pm long. Salivary glands not

hemi-spherical, so a deformed glandular mass.

Venom gland very long, narrow and convoiute,

about half anterior and half posterior to nerve

ring, distal région broader th;m proximal région,

but apparcntly with same inner tissuc organiza¬

tion. Muscular bulb elliptical, short, almost circu¬

lât in section; two inner muscular layers, inner

layer of about half thickuess of outer layer. Inner

surface of buccal mass similar to that o! preced-iug

species. A sketch ol foregut structures shown in

Figure 7F. Oesophagus a long, narrow tube

(Fig. 17F), flattened and with smooth inner sur¬

face in its anterior région, narrow, with several

longitudinal folds in its posterior région; no deve-

loped inner glands. Stomach (Fig. 17F) simple,

curved, located half a whorl posterior to kidney,

tittmersed in digestive gland. Ducr to digestive

gland single, located in middle, ventral région of

stomach. Digestive gland of about 5.5 whoris pos¬

terior to stomach, beige. Intestine hroad, with

thin walls, inner surface almost smooth, runs to

left of and ventral to kidney. Other details in

Simone & Verissimo (1995: 464, fig. 7).

Génital System (Fig. 17A, D-F)

Male. Testis rather short, located in columellar

région of second whorl posterior to stomach.

Viscéral vas deferens very hroad at lev'el of half a

whorl belore kidney, with thick, convoluted walls

(Fig. 17F). In rénal level vas deferens narrows

abruptly, runs near columella and exits to palliai

cavity in its posterior righr région. Prostate a

thick-walled. hroad, short tube of about half

length of rectum, withouc détectable aperture

(Fig. 17F). Vas deferens runs along floür of right

margin of palliai cavity, in its anterior région

stays immersetl in heâd tégument; before its

entrance to pénis, vas deferens thicker. Pénis

(Fig. 17A, D) rather short - lirtle more than half

length of palliai cavity, flattened; wilh about uni-

form width along its length. Pénis dtict very nar¬

row. clearly sinuouS in its proximal half, mnning

near left pénis margin. Pénis distal end with a

concavity turnecl to right, in its centre a rather

large papilla where pénis duct opens.

Female. Viscéral oviciuct very narrow, runing

close to columella, with a small gono-pericardial

aperture; inserted in palliai gtmoduct at righr of

albumen gland. Entité palliai oviduct very

short - about hait length of rectum (Fig. 17E).

Ingesting gland elliptical, ciicular in section,

with muscular walls; receptaculutn seminis,

connecting ingesting gland to oviduct, narrow,

long and convolute, located dorsal to ovoduct.

Albumen and capsule glands connected to each

other, with a single flattened duct which opens

in right région of anterior extremity close to

columella. Terminal pouch lacking.

Terebra gemmulata Kiener, 1 839

(Figs IG, H; 3D, E; 7G; 8C; 13A; 18-20)

Synonymy — cf. Matthews et al ., 1975: 90 and

Bratcher & Cernohorsky, 1987: 148.

224

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Braailian Terebridae

Fig. 18. — Terebra gemmulata, anatomy; A t head-loot, mantle removed, introvert partially protracted; B, detail of A in région of

tentacles in a specimen with visible eye (generally they are not visible); C. Iransverse section of middle région of palliai roof; D. ante-

rior région of viscéral mass, ventral view, digestive gland near stomach partially removed, Kidney and pericardium opened, their

internai région exposed; E, palliai organs, ventral-lnternal view. Scale bars; A, E. 1 mm; B-D, 0.5 mm.

Material. EXAMINEE). — Sâo Paulo. Brazil, off Measurements. — MZSP 28392: 37.9 x 7.8 mm:

Santos Bav, 32 m„ 14.IV.1969, 2 9 9 (MZSP MZSP 19325: 32.0 x 7.7 mm.

28392).

Rio Grande do Sui. Brazil, off Mostardas, RV W.

B es nard, sut 430, 31°33'S, 51°05’W, 21 m,

5.X1.1968, 1 <3 (MZSP 19325).

Description

Shell (Figs IG, H; 13A)

Shell description found in Matthews et al. 1975;

90-91 (figs 7, 8); Bratcher & Cernohorsky 1987:

148 (fig. 166a-e).

DISTRIBUTION. — From Rio de Janeiro, Brazil to San

Marias Gulf, Argennna.

ZOOSYSTEMA • 1999 • 21 (2)

Simone L. R. L.

Head-fuot (Figs 8C; 18A, B; 19A)

Colour homogcneous yellowish cream. Hcad

weakly differentiated Front head-foot axis

(Fig. 1 SA), Tentades ver y .short, dorso-ventrally

flattened. tip roimded (Fig. I8B). F.yes lacking in

some specimens or exrreroely small, dark, located

at tenracle base (Fig. 8C). Basal, proximal intro-

vert aperture rather broad, transverse, anterior

and ventral to tenracles (Fig 19A). Foor occu-

pying almost half a whorl, without divisions; sole

with folded borders. Columellar muscle ot two

whorls, rather thick (Fig. 18A). Males with very

large pénis in central-right région posterior to

head.

Operculum (Figs 3D, E; 18A)

Large, unguiculate, brown, nucléus terminal

(Fig. 3D, Fl). Occupving entire shell aperture

(Fig. 18 A).

Mamie organs (Figs 18C, E; 20D)

Mantle border simple, nor pigmented. Siphon

well-developed, yellowish créant, with srnooth

borders (Fig. 18E). Mautlc cavity of about

2.5 whorls. Osphradium elliptical, very long

about half of gill length; with several uniform

filaments on both sides of osphradial ganglion;

right filaments similar in size to left filaments;

anterior half broader chan posterior, which tapers

gradually. Gill narrow and long, about seven

tenths of palliai cavity length. anterior end rather

far from mantle border, with onlv a short portion

of ctenidial vein présent, in form of a very small

septum; filaments begin gradually some distance

from anterior end (Fig. 18E); each filament tri-

angular, low, apex located at the right, margin

almost straight (Fig. 18C); gill posterior extremi-

ty very far from pericardium, a long portion of

ctenidial vein frec from gill filaments (Fig. 18E).

Ctenidial vein narrow and uniform ail along Us

length, cxcept fin tes broader anterior extremity.

A proportionally broad space between gill and

rectum. Hypobranchial gland thin, located in

posterior half of’ cavity, at left of rectum, pale

cream. Anal gland with a few slender, irregular

acina immersed in right margin of hypobranchial

gland in anterior half of rectum, but far from

anus (Figs 18E; 20D), purple. Palliai gonoducts

run along right margin of posterior half of palliai

cavity, attached to palliai floor. Rectum narrow,

lying ventral to and left of palliai gonoducts; in

females, in its middle région il separates from

oviduct on margin of a palliai septum (Figs 18E,

20D). Anus far from anterior extremity of palliai

oviduct in females or prostate in males; bears a

small papilla. Anterior third of palliai right mar¬

gin without internai structures.

Circula tory and excreiory Systems (Figs 18D; 20A)

Heart of small size, position similar to those of

precedmg species. Kidtiev of almost half a whorl,

flattened, located. in right région of posterior

limit ot palliai cavity. Kidney wirh a large wide

inner chamber scparacîng nvo main glandular

masses. Dorsal glandular mass rather thick and

bearing sevcral lobed folded acini. Venrral right

mass rhin, bordering and attached tu rectum,

bearing sevcral rransverse, uniform foids. Both

kidney niasses pale cream. Nephridia! gland well-

dcvclopcd, pale cream, triangular itt section, hor-

dering dorsal margin of membrane between

kidney and pericardial chambers (Fig. 18D: ng).

Nephrostome a transverse slit in middle région of

membrane between kidney and palliai cavity.

Digestive System (Figs 7G; 18A, D; 19; 20A)

Rhynchodeal introvert very long, about 1.5 time

rhynchocoel length (coiled in retracted condi¬

tion), cylindrical (Figs 18A; 19A. B. F), muscular

walls thick, separated into nvo layers cotmected

to one another hy sevcral, uniformly distributed,

transverse muscle fibres (Fig. 19D). Orner laver

(jn retracted condition) matnly of longitudinal

fibres and inner layer mainly of’ circulât fibres.

Distal aperture of rhynchodeal introvert (rhyn-

chostome) large and with a terminal muscular

sphincter: in une side a pair of longitudinal foids

(Fig. 19D). The nvo layers of introvert walls do

not glide over one another when introvert is pro-

tracted, so whole vvall is evagined. Rhynchodeal

wall very thin, transparent, covering inner surfa¬

ce of anterior hall of haemocoelas in preceding

species (Fig. 19). Several small muscle fibres

connect rhynchodeal walls wirh inner surface of

haemocoel (more concentrated in anterior extre-

mlry) and with other foregut structures; two

muscles, in particular, more developed; one pair

posterior to nerve ring, connecting oesophagus

226

ZOOSYSTEMA • 1999 - 21 (2)

Comparative morphology of Brazilian Terebridac

Fig. 19. — Terebra gemmulata, anatomy; A, head and haemocoel, ventral view, foot and columellar muscle removed; B, foregut

removed from haemocoel, lett view, oesophagus partially deflected; C, detail of B in proximal région of oesophagus, part of rhyncho-

deal wall also shown; D, rhynchodeal introvert, details ol îts internai surface (when retracted) exposed by means of longitudinal sec¬

tion, distal (upper) and proximal (down) extremîties, double walls clear at right side; E, foregut removed, rhynchodeal wall sectioned

longitudinally, introvert rellected, salivary glands seen by transparency; F, detail of posterior région ot rhynchodeal wall and vestigial

proboscis opened longitudinally, internai surface exposed- Scale bars: t mm.

with dorsal wall of haemocoel, cmbracing also

rhynchodeal wall (Fig. I9A, B: ml), and anorher

muscle in posterior extremity of rhynchococl,

connecting oesophagus to buccal mass (Fig. 19A:

m3). Proboscis very small, reduced, présent only

as muscular ring around anterior surface ol buc¬

cal mass (Fig. 19B-F). Buccal mass spherical. Pair

of retractor muscles reduced. Radular sac wan-

ting. Salivary glands divided into two separated

masses (Fig. I9B-R), one ol rliem located antc-

riorly and the other posteriori}', connected with

eaclt other by a narrow duct inscrted in oesopha¬

gus near posterior salivary gland, whcre in sonie

spécimens there is a small elliptical pouch

(Fig. 19C: ep) (some specimens lack this pouch).

The exact position of the salivary glands also

ZOOSYSTEMA • 1999 • 21 (2)

227

Simone L. R. L.

Fig. 20. — Terebra gemmulata, anatomy; A, anterior région of viscéral mass and adjacent région of palliai cavity, ventral view, peri-

cardium and kîdney partialty dissected; B. pénis, ventral view; C, pénis, transverse section in ils middle région; D. palliai oviduct and

adjacent areas, ventral view; E. pénis, detail ot its distal extremity seen as if transparent. Scale bars; 1 mm.

varies among spécimens, it can lie more anterior

or posrerior, close to onc another, etc. Venom

gland and nutscular bulb absent, A sketch of

foregut structures shown in Figure 7F : . Inner sur¬

face ol buccal mass with several longitudinal low

folds which continue into ocsophagus (Fig. 19F).

Oesophagus a long, narrow tube, running with

some coils to anterior région where ic passes

though nerve ring, twists and retu ms to posterior

(Fig. 19A, b), this position is maintained by a

small pair of muscles ml; interior glands no

developed; posterior région with internai longi¬

tudinal folds until near stomach. Stomach

simple, curved (Fig. 18D), located half whorl

posterior to kidney, immersed in digestive gland.

Ducr ro digestive gland single, located in middle,

ventral région of stomach. Digestive gland of

about 5.5 whorls posterior to stomach and half a

whorl anterior to it (Figs I8D; 20A), beige.

Intestine broad, with thin walls and an almost

smooth internai surface; runs left and ventral to

margin of kidney. Rectum and anus described

above.

Génital System (Figs 18A, D, E; 20)

Male. Testis similar to that of preceding species.

Viscéral vas deferens runs close to columclla.

Prostate thick-walled, conical, broad and short,

228

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

located in right région of kidney and nor in pal¬

liai cavity (Fig. 20A), except for a short amener

portion; viscéral vas deferens inserted in middle

of prostate posterior surface. Palliai vas deferens

runs alcing righr margin of floor of palliai cavity,

in its anterior region remaining immersed in

head tégument. Pénis very long, longer than pal¬

liai cavity length (distal region stays twisred)

(Fig. 18A), flattencd (Fig. 20B); narrowing gra-

dually; pénis duel very narrow,. nearly srraight,

near left margin of pénis (Fig. 20B, C); on its

opposite side (right) a différentiable blond sinus

(Fig. 20C). Pénis distal end with a deep concavi-

ty turned to right and with a small aperture; in

its centre a ratlier large papilla in which pénis

duct opens (Fig. 20E).

Femalc. Ovary similar in position to testis.

Viscéral oviduct very narrow (Fig. 18D), runing

close ro columella, inserted in palliai oviduct in

its right posterior extremity; with a small gono-

pericardial duct. Albumen gland short, thick-

walled, white (Fig. 2UD). Ingesting gland rather

triangular, located in left posterior region of pal¬

liai oviduct; reccptaculum seminis long. Crossing

limir between albumen and capsule glands, insert-

ing in right margin of palliai oviduct. Capsule-

gland yellow, long - about half of total palliai

oviduct length, cylindrical, with a central, flat-

tened duct which opens in a small slit at righr of

terminal pouch base in lip ufan outstanding élé¬

vation (Figs ISF.; 20D). Terminal pouch broad,

with a longitudinal furrow turned to left and

opened; left lip of this aperture much smaller

than right one; occupies almost half of total leng¬

th of palliai gonoduct (Fig. 20D)

Terebra leptapsis n. sp.

(Figs 7F; 12B, C; 13F; 14C, G; 21; 22)

Type materiai.. — Holotype: MZSP 28704.

Paratypes: I specimen MZSP 28705; 13 shells

MZSP 28700, ail frorn type local i ty.

OTHF.R MATERIAU. Rio de Janeiro. Braxil, off

Cabo Frio. RV \V" Besiiiii/J, stn 1485, 22"39’S,

4l°33’W, 52 in, 10.111.1971, 7 shells (MZSP

19404). — Stn 1483, 23°00’S, 42 U 10’W, 64 m,

10.111.1971, 3 shells (MZSP 19403). — Off Rio de

Janeiro, stn 1476, 23°06’S, 42°54’W, 59 m,

9.111.1973. 9 shells (MZSP 19402). — Ilha Grande,

otter trawl, 50 m. VI. [971, 1 shell (MNRJ 3656).

Süo Paulo. Brar.il, oIT Uhatuba, RV' Velign, stn 26,

23“40’S, 44“54'W. 42 m, 21.1V. 1986, 1 specimen

(MZSP 28710); stn 17, 23°34'S, 44°48’W, 44 m,

22.1.1986. (MZSP 28709); stn 31, 23°5.VS,

45°09 W, .38 m, 10.VII 1986, I shell (MZSP

28703). — RV W. Bestirtfd, stn 4851, 23°36'S,

44°46\V, 48 m, 16.XII.1985, 13 shells

(MZSP 28699), 7 shells (MZSP 28702); 23 0 36.2’S.

44"39.5’W, 50 m. 26.VII.1986, 1 shell (MZSP

28701). OffSanros Bay, 32 m, 14 IV.I960. 1 shell

(MZSP 28394). — 23“36'S, 44°46'W, 48 ni.

I6.XI1.1985, 1 specimen (MZSP 28395). — No loca-

lity, Canversul-I , NOAS expédition, stn 4594,

loc. 4511, 30 m, 25.VIII.1977, 1 shell (MZSP

28700 ).

T ype I OCAI.ITV. — Brar.il, Rio de Janeiro, Campos

Bay (RV Austrogamupa).

EtÿMOI.UGY. - I he spécifie epither reters to the nar¬

row reticulate sculpture of the shell, from Greek leptos

(tentions) and hupth (net),

DISTRIBUTION. — From Rio de Janeiro to Sâo Paulo,

Braril.

Habitat. — Sandy substrates; from 32 to 64 m

depth.

Mf ASUREMENTS. — In m'dlimeters, followed by num-

ber of axial and spiral ribs in penultimate wliorl: holo-

tvpe, 15.9 X 4.0, 27, 6; paratype MZSP 28705,

15.7 x 3.7, 27.5.

Remarks

The holotype and paratypes MZSP 28705,

28706 were collecred ar several points of Campos

Bay, no précisé locality or depth was given. Even

so, they are chosen tfue ta their perfect shells, in

ail other availablc lots the shells are eroded or

broken. The single preserved specimen

(MZSP 28395) had its shell broken for anato-

rnical study.

Diagnosis

Shell small, with protoconch of two whorls;

sculptured by narrow axial and spiral ribvS, ol

cqual prominence, about thirry axial ri b in

penultimate wliorl. Temacles separated from onc

another and with eyes at tip. Forcgut complété,

with broad, flattened accessory proboscis struc¬

ture. Pénis long and narrow.

ZOOSYSTEMA • 1999 * 21 (2)

229

Simone L. R. L.

Description

Shell (Fig. 12B, C; 13F; 14C, G)

Long, slender, turriform, small (about 13 rnm

high), with up to thirteen convex whorls, pale

cream. Protoconch (Fig. 14C) ol two whorls,

convex, smooth: junction with teleoconch clear,

Sculpiure ol (ïrst releoconch whorls (Figs 12B,

C; 14G) mainly nf narrow, uniform axial ribs,

each with a subsurural rtode; gradually spiral ribs

appear, six to seven on pejuilrimare whorl, wilh

abour sa me widrh as axial ribs; a small node ai

intersection of spiral and axial ribs; subsurural

nodes prominent; abour thirty axial ribs on

penultimate whorl. Aperture simple (Fig. 13F);

outer lip with cur-edge; innet lip smooth, sig-

moid. Canal short, open, curved.

Head-foot (Figs 21 B, C; 22A)

Colour homogeneous clear beige. Head weakly

differentiated from head-foot axis (Fig. 21 B).

Tentacles short, dorsoventrally flarrened, tip

rounded, Well-separatcd from each other

(Figs 21 B, C; 22A). EyCS rather large, dark, local-

ed at tentacle tip. Basal, proximal imrovert aper¬

ture narrow, transverse, anterior and ventral lo

tentacles. Foot occupving almost half a whorl,

without divisions; furrow of pedal glands ante¬

rior. Columellar muscle of about 1.5 whorls,

rather thick. Male with pénis inserted very post¬

eriori}' (Fig. 21 B).

Operculum

Large, unguiculate, pale brown, nucléus termi¬

nal. Occupying entire shell aperture (Fig. 21 B).

Mantleorgans (Fig. 21 A, E)

Mantle border notched near siphon base and

some papilla-likc digitations to right and left of it.

Siphon well-developed, long, clear beige, with

smooth edges. Mantle cavityof about two whorls.

Osphradium. ellipticaJ long, about same Iength as

gill; with several filaments; right filaments larger

and more numeroüs than left filaments; anterior

two thirds of osphradium with filaments more

widely spaced, posterior one third wilh filaments

doser. Gill narrow and rallier short, about hall

Iength ol palliai cavity; anterior end rather close

to mantle border, a portion of ctenidial vein with

Iength about one third that of remainder of gill,

without filaments; filaments begin gradually aller

this; each filament triangular, apex nearly central,

margins almost straight; filaments of anterior half

ofgill more widely spaced than those of posterior

half; gill posterior end far from pericardium.

Ctenidial vein narrow and uniform ail along 1rs

Iength, exccpt (or its broader anterior exrremitv. A

proporiionally narrow space between gill and rec¬

tum. Hypobranchial gland thin. located in poste-

rior liait ol cavity, at left ol rectum, ycllowish.

Anal gland not différentiable. Pallia! spcrmoduct

run along right margin of posterior half oi palliai

cavity. Rectum narrow, lying ventral to and left of

palliai spermoduct. Anus near anterior extremity

of palliai spermoduct; bears no papilla. Anterior

hall of palliai right margin w ithout interior struc¬

tures.

Circulaiory and excretory Systems (Fig. 21 F)

Heart of small size, similar in location to those of

preceding spccies. Kidncy similar to that of

T. cntssireticula , exccpt in haviug ventral right

mass (bordering rectum) with fransverse lold.s

rai 1er than in T. cntssireticula. Ncphridial gland

also similar ro that ol T. crassireticula.

Nepluosrome a transverse slit iu middle région of

membrane between kidney aud palliai cavity.

Digestive systetn (Figs 7F; 2ID, F; 22A, B)

Rhynchodeal introvert occupies about one third

of remainder of foregut Iength, cvlindrical, thick-

muscular. walls double (Fig. 22A). Distal aper¬

ture of rhynchodeal introvert (rhynchostome)

wide. Internai rhynchodeal wall very rhin, trans¬

parent, covers internai surface of anterior half of

haemocoel as in preceding species (Fig. 22A, B).

Accessorv proboscis structure long (abour one

third Iength of total foregut Structures), clear

brown, flatrened, base broader, gradually tapers

into a rather rounded tip; connected to rhyncho¬

deal wall in middle région ol its left side

(Fig. 22A, B); no ligament connecting it with

internai surface of haemocoel; internai région

muscular, outer région glatidular. Proboscis coni-

cal, ol about same Iength as rhvnchocoel; its base

connectcd by small muscle fibres to rhynchodeal

wall and lo .several pairs of retractor muscles.

Buccal mass long (Fig. 22B), with internai and

external organization similar to that described for

230

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology ot Brazilian Terebridae

Fig. 21. — Terebra leptapsis, n. sp. t anatomy; A, palliai organs, ventral-internal view; B, head-foot of male, mande removed;

C, same. detail of tentacles région; D, radular tooth; E, palliai cavity roof, transverse section in ils middle région: F, last whorls ol vis¬

céral mass, ventral view, kidney and pericardium opened, adjacent région of palliai cavity also shown. Scale bars: 0.5 mm.

T. cinerea. Radula similar to that of T. Crassireti-

cula but with clearly longer and more slcndcr

teeth (Fig. 21 D). Salivary glands amorphous,

their ducts similar to thosc of preccding species.

Venom gland very long, narrow and convolute,

about one third anterior and two thirds posterior

to nerve ring. Muscular bulb elliptical, long

(Fig. 22A), section almost rriangular. Internai

surface of buccal mass similar to that of preced-

ing species. A sketch of foregut structures shown

in Figure 7F. Oesophagus long and narrow, llat-

tened, with smooth internai surface. Stomach

ZOOSYSTEMA • 1999 • 21 (2)

231

Simone L. R. L.

Fig. 22 - Terebra leptapsis n. sp . anatomy; A. head and haemocoel. ventral view, foot and columellar muscle removed. rhyncho-

deal wall partially opened longitudinally, accessory proboscis structure (ap) reflected; B, toregul (posterior extremity) and buccal

mass, rhynchodeal wall, proboscis. venom gland and oesophagus only shown in their basal région, O. pénis, ventral view, part of

palliai vas deterens also shown; D. pénis, detail ol its distal extremity as It a transparent structure. Scale bars: 0.5 mm.

simple, curvcd (Fig. 21F). located half a whorl Génital System (Figs 21 A, B, F; 22C, D)

posterior to kidney, immersed in digestive gland. Only niale examined. 1 estis - similar to those of

Duct to digestive gland single, located in middle. preceding Species (Fig. 21F). Viscéral vas defe-

ventral région ol stomach. Intestine broad, vvith rens narrow, almost Sttaight, running ventral to

thin walls, with an almost smooth internai sur- kidney (Fig. 21F). Prostate thick-walled. narrow

face, running left and ventral to kidney. Rectum (Fig. 21 A). Palliai vas deferens runs along right

and anus described above. margin of floor of palliai cavity, in its anterior

232

ZOOSYSTEMA • 1399 • 21 (2)

Comparative morphology of Brazilian Terebridae

région staying immersed in head tégument; befo-

re its enrrance to pénis, vas defercns thicker.

Pénis narrow and long, about saine length as pal¬

liai caviry (Fig. 2IB); origin very posterior, far

from tentacles; about unilorm width along its

length (Fig. 22C.). Pénis duct very narrow, clearly

sinuous in its proximal half, running near left

pénis margin. Pénis distal end with a concavicy

turned to right (Fig. 22D), in its centre a rather

large papilla where pénis duct opens.

Discussion

Shell and soft parts characters of T. leptapsis are

similar to those of T. crassireticula, from which

T. leptapsis dififers in luving: (1) proroconch with

one more whorl (two rather rhan one whorl);

(2) sculpture more délicate; (3) more axial ribs

(30 on penultimate whorl in contrast with about

18 in T. crassireticula)-, (4) ceplialic tentacles far

apart; (5) notcli in mande border at base of

siphon; (6) accessory proboscis structure broader,

shorter and without ligament in its base;

(7) venom gland with two risirds of its length

posterior to nerve ring rather than a half, and

(8) pénis narrower and longer. Botli differ from

T. doellojuradoi Carcellcs (1953: 14, Fig. 21)

(Bratcher & Ccrnohorsky 1987: 164-166,

fig. 194a, b) in being longer, more slcnder, differ-

ently sculptured (scc below) and in rhe absence

of a columellar callus. Moreovcr, /. doellojuradoi

has about 23 axial ribs on rhe last whorl (accor-

ding to Carcelles 1953 and Bratcher &

Ccrnohorsky 1987), while Y crassireticula Iras

fewer than twenty and the axial and spiral ribs

are broader; 7. leptapsis, in contrast, has alniost

thirtry axial ribs and both axial and spiral ribs are

narrower than those of the other two species.

The occurrence of I doellojuradoi on rhe

Brazilian coast is still obscure, in Sâo Paulo and

Rio de Janeiro coasts (the northern lirnit of its

range according to Rios- 1994) no specimen of

this species was found, nlthougb scveral spéci¬

mens of Y. crassireticula and Y. leptapsis were

identifted as Y doellojuradoi. I nc type locality of

Y. doellojuradoi is Uruguay and it probably

occurs on the Southern Brazilian coast, but a

more précisé définition of its géographie distri¬

bution dépends on further studies.

Terebra taurina (Lightfoot, 1786)

(Figs 6F, G; 7C; 8G; 13D; 14A; 23; 24)

Synonymy - cf. Matthcws et al. (1975: 87) and

Bratcher & Ccrnohorsky (1987; 128).

Otlters: Terebra taurina - Rios 1985: 130 (Fig. 589). -

1994: 180 (fig. 840).

MAThRIAI EXAMINE!). — Bahia. Brazil, Salvador. Baia

de Todos os Santos, 2 m. 11.1997, B. Albuquerque

coll„ i d (MZSI’2871 i).

Rio de Janeiro. Brazil, Baixos de Sâo Tomé, RV

Muriaé March , IX.1994, 2 3 â , 2 9 ? (IBU-

FRJ 8765).

Distribution. — From Florida, USA, to Santa

Catarina, Brazil.

HABITAT. — Intratidal, sandy or sandy mud sub¬

strates.

Description

Shell (Fig. 14A)

Adéquate description found in Matthews et al.

1975: 87-90 (Figs 1-6); Bratcher & Cernohorsky

1987: 128 (fig. 142a-d; pl. B, fig. 8). It is notable

for the présence of a well-developed columellar

fold (Fig. 13D) at die level of dre inner lip.

Head-foot (Figs 23A, B, C; 24A)

Colour homogeneotis clear brown. Head immers¬

ed (Fig. 23A). Tentacles moderately developed,

dorso-veturally flattened, tip rounded, broad

(Figs 23B, C; 24A). Fyes dark, located at tip of

tentacles, Basal, proximal mtrovert aperture a

broad slit just ventral to tentacles. Foot long, a

little more than half a whorl. Furrow of pedal

gland anterior, Columellar muscle of about two

whorls, thick, with an obvious cenrral-running

furrow due to shcll columellar fold.

Operculum (Fig, 23AJ

Unguiculate, pale brown, rather thick, nucléus

terminal. Occupying entire shell aperture

(Fig. 23A).

Matttle orgarts (Fig. 23D-F)

Manrle border simple, not pigmented. Siphon

well-developed, pale cream, borders smooth.

Manrle caviry very deep, extending about

2.5 whorls. Osphradium bipeccinate, elliptical,

long, about one third of gill length; with several,

ZOOSYSTEMA • 1999 • 21 (2)

233

Simone L. R. L.

Fig. 23. — Terebra lautina. anatomy; A >■! head-toot mamie removed B, same. detail ot head. dorsal view, foot partially removed;

C, same, ventral view: D, paillai cavity roof, transverse section of its middle région: E. paillai organs, ventral-mternal view, wlth two

details ot transverse section ot indîcated level ot prostate F, a whori posterlor and a whorl anterior to posterior extremivy ol palliai

cavity. ventral vtew, digestive gland adjacent to stomacb removed, distal région of oesophagus opened longitudinally wilh its internai

surface (wilh septa) exposed, kidney and pericardium partially opened: G, ventral région of stomacfi opened longitudinally. Scale

bars: A-D, 1 mm, E-G, 2 mm.

uniform filaments on both sides of osphradial

ganglion; right filaments obviottsly larger titan

left filaments and witli an angled projection

covering ctenidial vein; each filament with smooth

surface (not scalloped). Gill narrow, long, about

two thirds of palliai cavity lengrh; anterior end

near mantle border consisting only ol ctenidial

vein, in form of a small septum; filaments begin

gradually at sonie distance front anterior end;

each filament triangular, apex nearly central; gill

234

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 24. — Terebra taurina, anatomy; A, head and haemocoel. ventral view, foot and columellar muscle removed; B, foregut remo-

ved, rhynchodeal wall opened longitudinally; C, detail of posterior extremity ot foregut and buccal mass, base of proboscis and rhyn-

chodeal wall opened longitudinally; D, pénis, dorsal view. Scale bars' A, B, 2 mm; C, D, 1 mm.

posterior end very far from pericardium, with rectum. Anal gland absent. Palliai gonoducts run

about one third of posterior région of palliai along right margin of posterior half of palliai

cavity free from gill. Ctenidial vein narrow and cavityv partly connected to palliai floor. Rectum

uniform ail along ils length, except for its broad- narrow, lying at left of palliai gonoducts, its anre-

er anterior extremity; a very long portion rior région séparaied from gonoduct, attached to

without gill filaments in its posterior third. A mande (without septum). Anus locared posterior

proportionally broad space between gill and rec- to anterior third ol palliai cavity. Anal papilla

tum. Hypobranchial gland thin, small, transpa- présent. Anterior third of palliai right margin

rent, located in posterior hall ol cavity, at left of without interior structures.

ZOOSYSTEMA • 1999 • 2t (2)

Simone L. R. L.

CircuLnory and exmtory Systems (Fig. 23F)

Heart similur to rhose oF prcceding species.

Kidney of more than half a whorl, flattened,

located in right région of posterior limit of palliai

cavity. Kidney with rwo thin glandular masses,

separated by broad chambcr. Dorsal glandular

mass thin, vvhîte, boa ring a mosaie of irregular

acini, clearlv ihicker in anterior région. Ventral

glandular mass very thin, bordering rectum,

semi-transparent, bearing several, uniform trans¬

verse folds. Ncphridial gland narrow, white, rri-

angulâr in section, transversely folded, borders

dorsal margin of membrane between kidney and

pericardial chambers. Nephrostomc a transverse

slit in middle-right région of membrane between

kidney and palliai cavity.

Digestive System (Figs 6F, G; 8G; 23F, G; 24A-C)

Rhynchodeal introvert narrow and rather short

(Fig. 24A, B). Distal aperture of rhynchodeal

introvert (rhynchostome) a small transverse slit,

preceded by a thin sphincter. Internai rhyncho-

deal wall very thin. transparent, covers internai

surface ol anterior hall of haemocoel (Fig. 24A,

C), connected to ir by small muscle libres mainly

located in area from near rhynchostome to

middlc région. Proboscis conical, obviously

shorter than rhynchocoel, its base connected by

small muscle fibres and several pairs of small

retractor muscles to rhynchodeal wall. Buccal

mass spherical, with a long, broad buccal tube

(Fig. 24C), with organization similar to that of

H. cinerea , including gripping of tooth. Retractor

muscles also insert in latéral surface of buccal

mass. Radular sac short, curved, with several

radular teeth, opens sub-terminally in ventral-

rniddle région of buccal mass. Each

radular tooth (Fig 6F, G) curved, hollow, length

about 900 pm; base barbed, sinuous; basal région

narrower, concave; middle région broad, neatiy

cylindrical, section circul.tr (but nor altogether

fused); apical région sharply tapered, with a

sinuous barb edging a long, narrow aperture.

Salivary glands a pair of amorphous, white

masses connected to each other, rheir ducts

contour the oesophagus and insert in anterior

and posterior région of base of radular sac aper¬

ture. Venom gland long and convolute, about

two thirds anterior and one third posterior to

nerve ring (Fig. 24B); inscris in buccal mass close

to and at left of radular sac aperture (Fig. 24C);

no changes of glandular tissue along its length.

Pair of acccssory salivary glands long, flattened,

inserted in buccal mass close to each other in

région just anterior to insertion of radular sac

(Fig. 24C: as); rheir ducts fuse aftet a short dis¬

tance and run iu outer buccal lube wall, opening

in its anterior région. Muscular bull) rather long,

elliptical, with broader proximal région and

sharp distal exrremity (Fig. 24A, B); rwo layers of

muscular tissue, internai la)er about half thick-

ness of outer layer (Fig. 8G). Internai surface of

buccal mass similar to that of H. cinerea.

Oesophagus a long, narrow' tube with smooth

internai surface; in distal région, prcceding sto-

mach, six to seven transverse, sh.illow septa

(Fig. 23F). Stomach similar to those of prcceding

spccics (Fig. 23F); internai surface with longitu¬

dinal folds in proximal région, and a.single longi¬

tudinal fold in distal région, having several

secondary transverse folds inserted along side,

uniformly spaced (Fig. 23G). Duct to digestive

gland single, located in middle, ventral région of

stomach. Digestive gland of about 4.5 whorls

posterior to stomach, beige. Intestine narrow,

with thin wall s, internai surface altnosr smooth;

runs left and ventral to margin of kidney

(Fig. 23F). Rectum and anus describcd above.

Génital System (Figs 23A, E, F; 24A, D)

Male. Testis in columellar side of viscéral mass,

narrows gradually anterior io stomach (Fig, 23F).

Viscéral spermoduct a continuation of testis, very

narrow, runs venually, close to columella, bet-

w'ccn rectum and kidney. Palliai spermoduct runs

along right margin of palliai cavity floor. Prostate

only developed at somc distance from posterior

limit of palliai cavity, spermoduct is inserted in ir

subterrninally (Fig. 23F, F). Pénis long (Figs23A;

24A), dorso-ventrally flattened, tw'isted; pénis

duct coiled in proximal région, very narrow, run-

ning near left margin of pénis (Fig. 24D). Pénis

distal end rounded, simple, without papilla nor

chambers; pénis duct opens in rip,

Female. No fentale examined had well-preserved

posterior génital structures; it was possible to

note only a well-developed terminal pouch just

posterior to anus.

236

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

(MORG 39001), 1 shell (MOKG 39002). — Off

Itaipti, Niterôi, 18 ni (nn sand): 1 shell (MORG

39000), 1 slull (MORG 22001). — Off Rio de

laneiro, 33 ni (on sand), I slicll (MORG 39003),

'40-30 ni, I shell (IBUPRJ 8918). — Off Juatinga,

40-30 ni. 2 shells (MORG 39003). ~ Ilha Grande,

RV W. Bcsuard, stn 330, 111.1909, 1 shell (MZSP

2321 2). —Enseada Lapes Mondes, RV Ttni/ta,

stn 262-263, 20.VIi.1966. 1 shell (MZSP 23200).

Angr.i elos Reis, RV' W. Bonard coll., stn 332. 19 ni,

16.111.1969. 1 â. 1 ? (MZSP 23211); stn 349,

111.1969, 1 â (MZSP 25213). — Sanie loeahty. RV

Rrntlia , stn 24S, 29 ni. 17.V1I.1966. I shell

(MZSP 23203); stn 142c, 10 tn. I9.VI.1967. 1 shell

(MZSP 25198); stn 266c, 22.5 ni. 17.VI.1967,

1 shell (MZSP 28708); stn 172, 8.VI 1.1966, I shell

(MZSP 25202); stn 173, 20 5 ni, 22.VII.1966, I shell

(MZSP 25201); stn. 266. 22.5 m, 20.VI1.1966,

1 shell (MZSP 25207); stn 292, 25 m, 30.VI1.1966,

I shell (MZSP 25209); stn 289, 42 m, 30.V1I.I966,

Terebra spirosulcata

Simone &c P. M. Costa, n. sp.

(Figs 1E, F; 3H; 6H, 1; 7E; 8D, E; 13C; 25; 26)

Terebra doelkjitradoi (?) — Matthews et ai 1975: 97

(figs 25-28) (notCarceiles 1953).

terebra brasiliensii — Bratcher & Cernohorsky 1987:

160 (figs 186c, d) (part.) (not Smith 1873).

TYPES MATERIAU — Rio de Janeiro. Holotype:

MNRJ 3657 (Iront type localiry).

Paratvpes: otf Cabo Frio, 60 ni, X.1993, I shell

(MORG 32755). — Off Cabo de Sào Famé, RV

VP. Besnard , stn IV, 22'06'S. 4I°04'W, 16 m,

II.II.1969, I shell (MZSP 19596). — Stn III.

22" 10.5’S, 4J"59'0”W, 30 ni, 2 shells (MZSP

25210). — TVpc loeality I shell. (MNR| 7570). —

Off Cagattas Island, 30-40 m, XI.1971, ! shell

(MNRJ 3655), 2 shells (MNRJ 7571), 1 shell

Fig. 25. — Terebra spirosulcata n. sp., anatomy; A, 8 headfoot, mantle removed; B, palliai organs, ventral-lnternal view; C, detail of

A in tentacle région; D, palliai cavity roof, transverse section of ifs middle région; E, lasl whorl of viscéral mass and posterior part of

palliai cavity, ventral view, part of digestive gland adjacent to stomach removed, kidney and pericardium opened, internai structures

exposed. Scale bars: A, 1 mm; B-E, 0.5 mm.

ZOOSYSTEMA • 1999 • 21 (2)

...

\JjVA

ac H

Simone L. R. L.

1 Shell (MZSP 25208); stn 176.30.4 m, 26.VII.1966.

2 shells (MZSP 25203); stn 236, 16.2 m,

12.VII.1966. 1 shell (MZSP 25204).

Sâo Paulo. Paratvpes: Ubaruba, Lagoa Beach; 5 shells

(MZSP 28707).

Santa Catarina. Paraiypes: Campeche Island, in

beach, 11.1985, 1 shell (MORG 39004).

Type LOCAÜTY. — Brazil, Rio de Janeiro, offllha do

Pai, 30-40 m depth (sandy).

Distribution. — 1 rom Rio de Janeiro to Santa

Catarina, Brazil.

Habitai'. — From 10 to 60 m depth, sandy sub¬

strates.

Mf.A.SUREMENTS. — In millimeters. Holotype

MNRI 3657: 26.2 x 6.4. Paratypes MNRJ 7570,

24.6 x 5.7; MNRJ 3655, 26.9 x 6.6; MNRJ 7571.

29.4 x 7.0, 18.8 x 4.6; MORG 39000. 26.8 X 6.0;

MORG 39003, 23.1 x 5.8; MORG 22001,

20.7 X 5.4; MORG 32755, 24.0 X 6.5;

MORG 39001, 11.1 x .3.0; IBUFRJ 8918,

23.0 X 5.9.

Remarks

This species was simultaneously discovered by

Paulo Marcio Costa, Universidade Fédéral do

Rio de Janeiro. For this reason, he is included as

one of tlte authors herein.

Diagnoses

SE Brazilian deep-water species with medium-

sized shell. Sculptured with low axial undulations

and narrow spiral furrows. Larger specimens with

last whorl outstandingly large. Foregut with

complété venom apparatus and a very long intro-

vert.

Description

Shell (Fig. 1 F, F)

Long, slender, turriform, of medium size (about

40 mm high), up to fifteen almost fiat to convex

whorls, white créant, some specimens with

brown axial blotches. Proroconch of 2.5 whorls,

convex, stnooth. TeleOconch glossy, sculprured

mainly by very shallow axial ribs, almost undula¬

tions (Fig. 13C), uniformly spaced, about 23 on

penultimate whorl, more prominetu near uppér

suture; about eigln spiral turrows on anterior

(lower) two thirds of each whorl, rather equally

spaced, except rhe upper ones, which may be

doser to one anorher than the lower ones. In lar¬

ger specimens last whorl very large in relation to

preceding whorls (Fig. 1E, F). Aperrure simple,

large; miter lip with eut edge; inner Lip smooth,

sigmoid. Canal short, open, curved.

/ It-ad'fooi (Figs 25A, C; 26A)

Colour homogeneous cream. Head weakly diffe-

remiared from head-foot axis (Fig. 25A).

Tentacles moderately devdoped. dorso-venrrally

dattened, tip rounded, broad (Figs 25A, C;

26A). Eyes dark, lacated at tip of tentacles. Basal,

proximal introvert aperturc tather broad, trans¬

verse. anterior and ventral to tentacles. Foot of

about half a whorl, withouc elcar divisions; sole

with folded borders. Furrow of pedal gland ante¬

rior. Columellar muscle of about 1.5 whorl,

thick.

Operculum (Figs 3H; 25A)

Unguiculate, pale brown, nucléus terminal

(Fig. 3H). Occupying entire shell aperture

(Fig. 25A).

Mantle orgam (Fig. 25B, D)

Mande border simple, not pigmented. Siphon

well-dcveloped, pale cream in colour, borders

smooth. Mantle cavity of about two whorls.

Osphradium fùpectinate, clllptical, long, about

balfof gill Icngtlt; with severai uniform filaments

on both sides of ospbradial ganglion; right fila¬

ments obviously larger titan left filaments; each

filament with smooth surface (not scalloped).

Gill narrow and long, about eight tendis of pal¬

liai cavity lcngtb; anterior end rather far from

mande border; with onlv ctenidial vein présent,

in from of a small septum; filaments begin gra-

duallv at some distance front anterior end; each

filament triangular, apex located at right; gill

posterior end far from pericardiuni. Ctenidial

vein narrow and uniform ail along its length,

except for its broader anterior extremity. A pro-

portionaJly broad space between gill and rectum.

Hypobranchial gland thin, small, located in post¬

erior half of cavity, at left of rectum, pale cream.

Anal gland with severai slender, irrcgular acini

immersed in right margin of hypobranchial

gland close to rectum (Fig. 25B), purpie; post-

238

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

Fig. 26. — Terebra spirosulcata n. sp„ anatomy; A, head and haemocoel, ventral view, toot and columellar muscle removed, rhyn-

chodeal wall parlially opened with introvert reflected outside; B, detail ûl poslerior extremity ot rhynchodeum and buccal mass, ven¬

tral view; C, pénis, dorsal view; D. sarne. detail ot its distal extremity as a transparent structure; E, palliai oviduct (posterior extremity

damaged) and adjacent région of palliai cavlty, ventral view. Scale bars: 0.5 mm.

erior acini longitudinally disposed. Palliai gono- in female gonoduct séparâtes from palliai oviduct

ducts run along right margin of posterior half of after capsule gland, runs in tip of a small septum

palliai cavity, partly connected to palliai fluor. (Fig. 26E: re), far front terminal pouch. Anus far

Rectum narrow, lying to left of palliai gonoducts; from anterior extremity of palliai oviduct in

ZOOSYSTEMA • 1999 • 21 (2)

239

Simone L. R. L.

females or prostate in males; sessile, without

papilla; located in about middle région of palliai

cavity. Anterior half of palliai right margin

without interior structures.

CircuLuory and excretory Systems (Fig. 25E)

Heart similar to those of preceding species.

Kidney of almost half a whorl, flattened, located

in right région of posterior limit of palliai cavity.

Kidney with two thin glandular masses, sépara-

ted by broad, high chamber. Dorsal glandular

mass thin, whire, beating a mosaic ol irregular

acini, obviously rhicker in anterior région.

Ventral glandular mass very thin, bordering rec¬

tum, semi-transparent, bears several, unilorm

transverse folds. Nephridial gland narrow, white,

triangular in section, bordering dorsal margin of

membrane betwcen kidney and pericardial

chambers. Nephrostorne a transverse slit in mid-

right région ol membrane betwcen kidney and

palliai cavity.

Digestive System

(Figs 6H, I; 7E; 8D. E; 25E; 26A, B)

Rhynchodeal introvert narrow and very long,

about double length of rhynchodeal cavity

length, coiled within rhynchodeal cavity, cylin-

drical (Fig. 26A), with thick muscular walls.

Distal aperrure of rhynchodeal introvert (rhyn-

chostome) a small transverse sJit; a thin sphincter

preceding it. Internai rhynchodeal wall very' thin,

transparent, covering internai surface ot anterior

half of haemococl (Fig. 26A, B), conncctcd to it

by small muscle fibres mainly located near rhyn-

chostome. Proboscis conical, narrow distallv, of

about sanie length as rhy nchocoel, its base

connected by small muscle fibres in rhynchodeal

wall and by retractor muscle; proboscis wall

rather thin. Buccal mass spherical (Fig. 26B),

with a long and broad buccal tube, double-

walled. with organization similar ro that of

H. cinerea. Pair of retractor muscles also inscrts

in buccal mass latéral surface. Radular sac rather

short, curved, with several radular teeth, opens

sub-terminally in mid-ventral région ol buccal

mass. Radula of about twenty teeth, each tooth

(Fig. 6H, 1) hollow, broad, about 450 pm in

length, section almost circular (but not alto-

gether fused), base barbed, surface irregular (with

rather uniform, scale-like proximal half), nar-

rows at rip; tip harpoon-like, with a n.irrOw aper-

ture. Salivaiy glands a pair of amorphous, white

masses connected to each other; their ducts

contour oesophagus and insert in anterior and

posterior side of base of radular sac aperture.

Venom gland very long and convolute

(Fig. 26A), about half anterior and half posterior

to nerve ring; proximal half of venom gland

(Fig. 8D) broader and more flaccid than distal

hall, with thin glandular walls; distal hall has

thick glandular walls and a very narrow duct

(Fig. 8E); inserts in buccal mass close to and to

left of radular sac aperture. Muscular bulb rather

long, clliptical, with broader proximal région;

two muscular lavers, internai layer thinner than

hall thickness of outer layer. A sketch ol foregut

structures shown in figure 7E. Internai surface ol

buccal mass similar to that ol H, cinerea.

Oesophagus long and narrow, with smooth inter¬

nai surface. Stcimach similar to those ol prçce-

ding species (Fig. 25E). Duct to digestive gland

single, located in mid-ventral région of stomach.

Digestive gland of about 4.5 whorls posterior to

stomach, beige. Intestine narrow, with thin walls,

internai surface almost smooth; runs left and

ventral to margin of kidney (Fig. 25E). Rectum

and anus described above.

Génital System (Figs 25A, E; 26C-E)

Male. Testis in columellar side of viscéral mass,

sinuous, narrowing gradually anterior to stomach

(Fig. 25E), iiidcscent white in this région.

Viscéral spermoduct appears as continuation of

testis, very narrow, runs ventrally between rec¬

tum and kidney (Fig 25E). Palliai spermoduct

runs along right margin of palliai cavity floor

(Fig. 25A), without development of clear pros¬

tate tissue. Pénis very long, cylindrical, coiled

(Fig. 25A); pénis duct simple, very narrow, runs

near left margin of pénis (Fig. 26C). Pénis distal

end with a concavity turned to right (Fig. 26D),

in its centre a large, broad papilla in which pénis

duct opens. Pénis papilla about in centre of pénis

tip concavity.

Female. Posterior région of palliai oviduct not

seen in detail in Figure 26E. Albumen gland

short, ol same width as capsule gland, frorn

which it is difficult to separate. Capsule gland

240

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

long, broad, almost cyljndrical, its glandular ris-

sue suddenly finis h es, Icaving a small anterior

hollow chatrlber; a narrow duct runs from tHis

chamber anteriorly. Capsule gland aperture

small, in posterior région of terminal pouch, tur-

ned to columella. Terminal pouch about half of

capsule gland length and separated trom gland

by a narrow région; middle région broad, with a

broad, longitudinal, shallow aperture.

Discussion

The shell of /. spirosulcata spccimens resemble

that of T. brasiliensis , differing mainly by its lar-

ger slze and by its narrow spiral furrows. These

différences were noted by Bratcher &

Cernoholsky (1987) who regarded them only as

variation and grow th effett. On the basis of soft

part charâcters, however, no doubts persist of

their spécifie séparation. /.' spirosulcata difïers

from T. brasiliensis mainly in having; (1) cephalic

tentacles closer; (2) a complété proboscis (while

in T. brasiliensis it is vestigial); (3) a complété

venom a p para tus (while 7. brasiliensis lias lost it);

(4) absence of accessory proboscis structure;

(5) pénis longer, and (6) terminal pouch of

females widely open. Matthews étal. (1975) also

commented on and figured this species, calling it

a probable variation of 7. doellojuradoi.

Terebra sterigma n, sp.

(Figs 12D; 13G; 14D, H)

Type MATERIAL. — Holotype: MNHN. Paratypes:

1 shell, MNHN; 1 shell, MZSP 28727. Ail these from

type locality.

TYPE LOCALITY. — Brazil, lîspirito Santo, off

Regència, 19 n 34’S, 38 D 55’W. 340-360 m (RV

Marion-Dufresne , MD55, stn CB92. V. I98 7 ).

EtymolocIY. The spécifie epithet refers to the shell

sculpture, which is mainly axial ribs, from Greek ste¬

rigma (pôles).

Distribution. — Known only from the type locality.

Measurements. — In millimeters, follovved by num-

ber of axial ribs and spiral nodes in penultimate

whorl; holotype: 16.5 X 3.9, 25, 8; paratype; 16.7

(broken) X 4.1, 25, 8.

Diagnosis

SE Brazilian deep-water species with medium-

sized shell. Sculptured with many prominent

axial ridges

Remarks

Shell (Fig. 12D) long, slcnder, turriform, of

medium size (about 20 mm high), up to 14

whorls, homogeneous clear brown. Protoconch

(Fig 14D) of two whorls, convex, smooth; junc-

tion with tc-leoconch obvious. Teleoconch sculp¬

ture lirate (Figs 12D; 1 4H), of many axial,

onhocline ribs, uniformly spaced and close to

one another, about 25 on penultimate whorl;

spiral sculpture absent or only indicated by suc¬

cessive small, shallow nodes on axial ribs (about

eight on penultimate whorl); a shallow subsutu¬

ra! furrow. Aperture simple (Fig. 13G); outer lîp

with cut-edge, internai lip sigmoîd, smooth.

Canal short, open, curved.

Discussion

'This species was First discovcrcd by Dr. Bouchet

(MNF1N) and the specimens were kindly sent to

be included in this study. T. sterigma differs from

T. doellojuradoi, T. crossireticula and /.’ leptapsis

by the absence of well-developed spiral ribs, by

the broader shell, and, particularly from T. crassi-

reticula, by the number of protoconch whorls

(two rarher than one).

DISCUSSION

The présent discussion includes comparions of

the morphological characters of the species des-

cribed here and also comparions with other

closer (conoideans) and more remote gastropods.

The arguments presented are based on literature

data and also on self-observed species [e.g.,

Cerithioidea, Littorina fl a va, Hydrobioidea

(Simone & Moracchioli 1994; Simone 1995a),

tonnids (Simone 1995b), sevcral muricoideans

such as Bilccinanops sp. (Simone 1996), Thala

crassa (Simone 1995c), Austromitra sp. (Turner &

Simone 1998), some papers still in press]. Other

self-observed conoideans are the turnd

Cochlespira sp. (papers in press) and the conid

Conus bertarollae Costa & Simone, 1997. This

ZOOSYSTEMA • 1999 • 21 (2)

241

Simone L. R. L.

discussion excludes those made by Taylor,

Kantor & Sysoev (1993), except il there is any

interesting comment. TIku paper is recommend-

ed as a complément to this discussion.

Shell

The elongated, mu Inspirai forrn of the terehrid

shell is a long-known character of the family.

7. brasiliens is, however, présents a form of adult

shell similar to other species when they are

immature, i.e., with proportionally fevv whorls

and a iarge protoconch, combined with mature

génital organs. Thus the terni “paedomorphic”

(i.e., young form) appears appropiate to define

this character of T. brasiliensis.

Unlike the other terebrids, T. taurina possesses a

well-dcvcloped, tall told running down the entire

middle région of the columella (Fig. 13D), évi¬

dent unril near the internai lip.

The numher of protoconch whorls appears to be

in general one whorf In the présent analysis the

only species seen with more than one whorl was

T. crassireticu/a.

Head-foot

The foot of H. cinerea combined interestingly a

large foot si/e (more than half a whorl in length),

with a reduced operculum. The larger size of the

H. cinerea toot is possibly an adaptation to the

high-energy envitonment it livcs in,

Ail terebrids hâve a considérable réduction of the

temacles, In some species they can only be seen

by means of a microscope. The eve position, on

the other hand, appears to divide the terebrids

into rwo groups: the Hastula species which Irave

the eyes located in rhe renracle bases, and the

Terebra species which hâve the eyes in the rips

of the tentacles (except for T. gemmulata with

reduced eves). 77. bacillus (cf. Taylor & Miller

1990) also has basally placed eyes.

Pallial organs

As in most other gastropods, the siphon of the

terebrids has smooth margins, but //. cinerea has

a sériés of papillae on the siphon border. Of the

examined species, H. cinerea is the only one

constantly exposed to intertidal waves. The

papillae on the siphon border may be an adapta¬

tion to avoid sand pollution in the palliai cavity

(Fig, 211). This adaptation is unnecessary in the

other dccpcr water species. On the other hand,

77. hast a ta has a pair of basal projections on the

siphon (Fig. 9B), resembling those of the

Volutidae (Muricoidea).

The osphradium is always well-developed in

these piedatory animais. In some species, it is

even larger (in area) than the gill (e.g., T, nassire-

ticula, T. leptapsis) or présents scalloped leaflets

(e.g., 77 cinerea, 77, hastata). Réduction of the

left filarnenrs, found in / crassireticu/a, T leptap¬

sis and T brasiliensis, is normally associated with

size réduction as, e.g., in Columbellidae (Muri¬

coidea) (pers. obs.). These terebrid species are of

small size.

The gil! anterior extremiry of the examined tere¬

brids (except 7.' crassireticula ) is raised, seprum-

like, and the gill filaments begin after a short

distance, The presence of this small septum pre-

ceding the gill filaments appears to be a modifi¬

cation of the ctenidial vein, but more deraîled

study is necessary to confirm that. However, the

absence of this character in other families exa-

rnined (even conoideans) indicates it may be

another character liniited to terebrids.

The long, digidform glandular acini, of conspi-

cuous purple colour, found in the right région of

the pallial cavity is hert callcd the anal gland, fol-

lowing the terminology ol Marcus & Marcus

(1960). Further research, however, appears neces¬

sary to détermine its hornology with the anal

gland of thi Muricidae The terebrid anal gland

appears to be a specialized région of the hvpo-

branchial gland, rather than a structure associated

with the anus, from which it is distantly separa-

ted in some species. Also, it is lacking in T. crassi¬

reticula, T. leptapsis, T brasiliensis and T. taurina.

In females of T. crassireticula and T. spirosulcata,

rhe end région of the rectum lies on an enigmatic

septum, instead of directly connected to the

mande surface. This character is absent in males

of the sanie species, and deserves investigation in

immature females.

Kl ONLY

Except for T. brasiliensis, ail examined terebrids

hâve a kidney comprising a broad chamber limi-

ted by rwo lobes, one lobe septate, attached to

the rectum and the other flattened and dorsal.

242

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

An aJmosr solid kidney (withouc large internai

hollow chamber) appears to be more primitive,

however sonie cloubts exist with respect to the

typical kidney of conoideans.

DIGESTIVE SYSTEM

l'he rhynchodeal svall in conoideans can be mus-

cular or membranous. In the turrid Cochlespim

and some Crassispinnae (Kantor et al. 1997) it is

of the firsi type. Muscle libres présent in the

rhynchodeal wall indicate some degree of

contractile function. This State is not found

among tercbrids, of which the internai rhyncho¬

deal wall is practically only a membrane, howe¬

ver, several other conoideans show this condition

(Taylor et al. 1993). The terebrtd rhynchodeal

wall still has a thin layer ol muscular fibres in the

région near the retractor muscle ot the proboscis

(Fig. 8A).

The proboscis of the conoideans, called an

intraembolic proboscis, has been regarded as an

independent acquisition from tho.se ol ol ber gas-

tropods, parrictilarly from other caenogastropods

(Kantor 1990). On the other hand, lecem papers

on gastropod phylogeny show the conoideans as

the last clade of rhe caenogastropods, with rela-

tionships with at least iwo clades ot pleurembolic

proboscidiferous animais such as muricoideans

and canccllarioideans (Pondcr & Lindberg 1996,

1997; Kantor 1996). With rhese data in mind, it

is already possible to construct a relarionship bet-

vveen the pleurembolic proboscis and the conoi

dean intraembolic one.

Furthermorc, observing tire structures ot a com¬

plété foregut cxtractcd front the examinée! spe-

cies, more similarities with the pleurembolic

proboscis appear. J he rhynchodeal wall is a dis¬

tinct membrane, connected with the internai

surface of the haemocoel by several sntall muscle

fibres, which vary in number according to spccies

and to spécimens; these sntall muscle fibres are in

general more concentrated anreriorly, close to rhe

rhynchostomc. These muscles are easily eut in

dissections and then the rhynchodeal wall

becomes a tree structure connected only by rhe

rhynchostomc. l'he rhynchodeal wall is conti¬

nuons with the proboscis, i.e., the proboscis is

not inserted in the haemocoel internai wall. The

transition front the rhynchodeal wall to the pro¬

boscis is noted by a sudden or sometimes graduai

increase uf rhe wall muscular rissue. l’he sntall

muscle fibres which connect the rhynchodeal

vvall to the internai surface of the haemocoel are

also generally concentrated at the transition from

rhynchodeal wall to proboscis, forming a muscu¬

lar ring as a base for these structures lo work.

From the internai surface of the proboscis, gene¬

rally a pair of proboscis retractor muscles inserts,

runs towards rhe posrerior, crosses berween the

rhynchodeal wall-proboscis transition and the

buccal mass, and connects in the latéral and ven¬

tral régions of the internai surface of the haemo¬

coel just po.sterior to the buccal mass level.

The group of structures that compose the

intraembolic proboscis (rhynchodeal wall, pro¬

boscis and its pair of retractor muscles) closcly

resembles the whole pleurembolic proboscis,

from which it diifers only by lacking muscular

tissue in its outer wall (rhynchodeal wall) and by

the short retractor muscles. With rhese argu¬

ments it is possible to offer another scénario for

the évolution of the conoidean proboscis (sce

also comments on buccal mass and tube below):

the conoidean ancestor was a pleurembolic pro¬

boscidiferous neogastropod; it lost the muscular

tissue ol the outer surface of its proboscis and

shnncned its retractor muscles, becoming the

main prorractible structure of the buccal mass

pari of the proboscis, and also acquiring a per¬

manent rhynchodeal cavity where the prey was

partly or wholly brought for initial digestion. In

other words, rhe conoidean proboscis may be

homologous to the buccal mass part of the pleu¬

rembolic proboscis (protracrible) and the conoi¬

dean rhynchodeal Wall is homologous to

remaining régions of die pleurembolic proboscis

(no longer protractible). These arguments are

also corroborated by finds of muscular tissue in

the rhynchodeal wall, muinly in its anrerior

extremiry (close to rhe rhynchostome) and in its

po.sterior extremiry (close to retractor muscles) of

examined tercbrids. Moreover, Cochlespim still

has the rhynchodeal wall entirely muscular

(Simone in press-a), as well as some Crnssi-

spirinae (Kantor et al. 1997).

Parallel to conoidean proboscis évolution was

that ol the buccal mass and buccal tube. The

buccal mass of examined species and most conoi-

ZOOSYSTEMA • 1999 • 21 (2)

243

Simone L. R. L.

deans (Taylor et al. 1993) is rctained in the pro¬

boscis base even du ring protracrion. This çapaci-

ty is possible due the élongation of the buccal

tube, which connccts the buccal mass at the base

of the proboscis vvith its dp. In observed tere-

brids and in Conus bertarolhte , the buccal tube is

not only long, but also double-walled. Its ourer

wall is in general thicker and muscular. Its inter¬

nai wall is in general thin and with a gland-ular

mucous epithelium, continuons with that ol the

buccal mass. It is possible that both walls of the

buccal tube are connectcd to each other, and

their séparation is an artifact of the fixation; no

living specinten was, however, dissected for

confirmation.

A scarcely muscular rhynchodeal wall, the pro¬

boscis (or buccal mass part of the proboscis) as

the main protractible structure, and the buccal

mass situated in the proboscis base, connected

with its tip by a long buccal tube, are not churac-

ters exclusive to the conoideans. At least, rwo

muricoidean taxa developed convergendy a very

similar proboscis-buceal mass complex, such as

the olivid Arnalda (Kanror 1991) and the costel-

lariid Amtromitra (Turner &c Simone in press).

The rhynchodeal introvert is a specialized tube of

the anterior extremity of the foregut, inserted in

the rhynchostomal région of the rhynchodeum,

and can be rétractée! within it complété])'.

Although double-walled. one wall does not glide

over the other dtiring pratracdon, instead the

entire wall is evagined and protracted, as a

wholc. This is observed in several spécimens pre-

served with the introvert only partiaily protrac¬

ted. This structure appears to be another terebrid

charactcr. The introvert is very long, i.e., as long

as, or longer than the remainder of the rhyncho¬

deum, in some species (T. spirosulcam. T. gemula-

ta, T. brasi/lensis). In other species, such as

T. gemmulata , onlv the introvert is présent, and

most other foregut structures, including the pro¬

boscis and venom apparatus, hâve been lost.

Howevet, the large inctease in the introvert is not

necessarily accompatiied by loss ol the proboscis,

as shown in /. spirosulcata. Other comments on

the introvert are found in Taylor et al. (1993:

128).

A terminologtcal problem was noted in the intro¬

vert, as both its extremities, the distal and the

proximal ends, could be ealled a rhynchostome.

When it is extended, the distal aperture Works as

the rhynchostome, whereas when it is in the

retracted position, the basal, proximal aperture

pertorms this function (as observed in some spé¬

cimens, the proboscis can protract without

necessarily the protracrion of the introvert). In

the présent paper, the recommendation of

Dr Taylor (pers. comnt.) is lollowed, using the

terni rhynchostome lor the distal, apical introvert

aperture.

Figure 7A shows a schematic synopsis of terebrid

foregut structures, in ventral view, based on the

Brazilian species examines!, including the acces-

sory proboscis structure (ap). This picrate was

obtained after detailed analysis of the examinée!

species front dissections and serial sections,

results shown in several figures through this

paper and suntniarized in Figure 7B-G (these in

dorsal view). These figures were tnspired by those

in the cunent lirerature (e.g., Taylor et al. 1993:

figs 25, 26), The supposition of a close relation-

ship berween the intraembolic conoidean pro¬

boscis and the pleurembolic proboscis of its

relatives (e.g., Muricoidea and Cancellarioidea) is

summarized in Figure 27. Note, again. that the

conoidean proboscis appears to be hontologous

to the buccal mass part of the pleurembolic pro¬

boscis, and the rhynchodeal wall of conoideans

appears to be homologous to the remaining

régions of the pleurembolic proboscis. The poly-

embohe proboscis, found in some conoideans

such as Terebra brasilicnsis and 7. gemmulata

represents, as noted Miller (1989) and Taylor et

al. (1993) a grear development of the introvert

accompanied by réduction of the proboscis and

other structures of the buccal mass.

A pair of very long, well-developed accessory sali-

vary glands is présent in H. hastata and /.’ tauri-

na. Howevet, in H. /tas ta ta one ts far front the

other, inserted on the opposite side of the pro¬

boscis base (each one close to a retractor muscle),

while chose of /. taurina are inserted close to

each other in the buccal mass. Fhe homology of

these accessory salivary glands with those ol the

other neogasrropods is a matter for further studies.

As is normal for proboscidiferous prosobranchs,

most terebrids possess a pair of strong retractor

muscles at the base of the proboscis. However,

244

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology ot Brazilian Terebridae

Fig. 27. — Schematic repiesentatlori ol transveise (frontal) section ot head, not to scale nor proportions, showing the simïlarlties bet-

ween the conoidean proboscis fashion (A) with the retracted pleurembolic proboscis of, e.g., a muricoidean (B). Note that the homo-

logy of each structure can be suggested. The rhynchodeal wall (rw) of conoideans is scarcely rnuscular (excepl in the turrid

Cochlespirà) and may be homotogous the outer wall of the pleurembolic proboscis (indlcated by dotted line). The conoidean buccal

mass is basal, while it is apical in muricoideans. but both groups hâve exceptions to this stalo. Soo loxt for more explanations.

T. crassiretiçula, T. le put pus, T. brasiliensis and

T. taurina posses several small pairs, distributed

around the basal région of the proboscis, connec¬

ting it with the adjacent région of the internai

surface of haemocoel. Sonic fibres of the retract-

or muscles also are inserted into the rhynchodeal

wall and buccal mass.

The accessorv proboscis structure is another

enigmatic character of terebrids, well-commented

on by Taylor et al. (1993: 129) and regarded by

them as a fanaily synapomorphy (shared with its

sister-taxon Pervicaciinae). Due to its presenee in

only sonie terebrids (e.g., the examined T. crassi-

reticula, Y leptapsis and T. brasiliensis), while

others (and other conoideans) hâve no vestiges of

this accessorv structure, three main conclusions

are possible: (1 ) it appeared in the terebrid ances-

tor and is secondarily lost in several members of

the grottp; (2) it appeared only once in terebrids

and is a synapomorphy of a subgroup; (3) it

appeared several tintes independently in terebrid

specîes. That thelast possibility is the most likely

is corrnborared by rhe high degree of variation of

this organ, which in some species is a simple pro¬

jection, while in others it is very complot and

branched. On the other hand, it always appears

itt rhe sanie location (left side of rhynchodeal

wall) and lias a similat structure (rnuscular in rhe

centre, glandulnr at the periphery with sensory

receptor.s). This suggests homology rather than

convergence, l hi,s question may only be resolved

after furf her studies.

Most terebrids, like other groups, possess female

palliai gonoducts closely auached to the rectum,

but in 7' penniniLna and T, taurina the rectum is

deflected in the middleor posterior région ofthe

oviduct, so rhe pallia! gonoduct no longer lies

near the rectum.

The terebrids (exccpt H. bastata .) show an inter-

esting tendency for a posteriorly located anus, far

ZOOSYSTEMA • 1999 ■ 21 (2)

245

Simone L. R. L.

from the mande border. This kecps a relatively

long région at the right of the palliai cavity free

of visible structures and may be another terebrid

character. The anal papilla is also found in

Cochlespira , and its présente in terebrids may be

plesiomorphic. However, the anal papilla appears

to be lost in 77 brasiliensis, 77 spirosulcata, and

77 gemmulata.

GEN1TAI SYSTEM

The tesris-seminal vesicle complet aad the ovary

of the terebrids taper gradually until thcy unité

with the gonoduct. I his condition differs from

mher groups in which thcre is normally a sudden

constriction separating the iwo régions.

However, this character is siill obscure among

other conoideans.

The prostate location in most terebrids is in the

palliai cavity as in most caenogastropods.

However, two species are exceptions. In the case

of 77 gemmulata, the prostate is located in the

anterior région of the viscéral mass, by the side of

(and compressing) the kidney. In the case of

77 spirosulcata , no differentiated prostate is pré¬

sent, ir is possibly disrribuied as a thin glandular

tissue along the palliai spermoduct.

Some muscles of the pénis base, protruding into

the haemocoel, are clearly visible in scveral cae¬

nogastropods, but in the case of II. hastata, rhese

muscles are much more cotnplex and cnlarged,

and when fully developed, displacing the foregut

structures to the lcfi. Most of the terebrids hâve a

broad papilla in the pénis tip, but it is absent in

77 taurina. Somcthing similar to this broad papil¬

la is also found ut the turrid Cochlespira and it

may be a basal character ol the conoideans

The ingesting gland of the palliai oviduct is nor¬

mally located berween rhe albumen and capsule

glands in rhe neogasrropods. This condition is

also found in rhe terebrids. However, H. hastata,

77 crassircticulit and 77 brasiliensis hâve the inges-

ting gland posterior to the albumen gland.

A terminal pouch with a large apelture is lound

in the examined terebrids (except 7. cmssiretieula)

and also in the turrid Cochlespira. It may be ano¬

ther character of the superfaniily. 7. crassirericula

has probable lost this structure. I brasiliensis, on

the other hand, possesses a very narrow anterior

aperture of the terminal pouch, with a deep,

blind sac duct, This condition resembles the

bursa copulatrix of several Muricoidea and of

Conus bcrtarollae Costa & Simone, 1997, from

which the terminal pouch may has evolved; if so,

rhe 77 brasiliensis condition could hâve assumed

the plesiomorphic condition, which would be

another indication of paedomorphosis.

Sl’ECIHS GROUPS

The terebrids, in the same way as several other

raolluscan groups, hâve revealed that small diffé¬

rences in shell characters, which might be regar-

ded as variation, may indicate deeper différences

in internai anatomy, for example, in 77 brasilien¬

sis and 7. spirosulcata. It is interesting to note,

however, that thcre are several 'species groups”

among Rra/.ilian terebrids, i.e., species with

greatly similar shell characters. 1 his is the case

of, e.g., the H. cinerea “group”, which includes

intertidal species such as H. salleana (Dcshayes,

1859) and 77 imiranix Auftenberg N Lee, 1988;

the 77 docllojuradoi "group", which includes deep

water, rcticulate-sculptured species such as

7. crassirericula , 7. leptapsis and 77 sterigma; and

the 7. protexta (Conrad, 1846) "group”, which

includes spinally groovcd species such as 7. t/osi

Bratcher & Ccrnohorsky, 1985 and 77 spirosulca¬

ta. Perhaps in the future, when more species are

known in more anatomical detail (mainly rhe

internai anaromy), these similaritics could be

bercer analysed.

Acknowl ed gem e n ts

Spécial thanks to Dr Philippe Bouchet, Muséum

national d’Hisroire naturelle, France, for provid-

ittg Marion-Dufresne Expédition deep-water tere-

brids and référencés. To Dr José Luiz M. Leme,

M/.SP for orientation. To Dr Cuido Pastorino,

Argentina, for référencés and persona! comments

on the manuscripr. To Cinria Miyagi and

Dr Atrron S. Tataran. IOUSP, for specimens of

rhe Bacia de Catnpos project. To Dr Ana Maria

Vanin, IOUSP, for specimens of the integrated

project. To Dr John laylor, The Naturel History

Muséum, London, for suggestions on the rermi-

nology. Pot helping in SEM examination ! thank

Enio Mattos, 1BUSP. Dr Sergio Vanin, IBUSP

for criticism of the text. Paulino Souza Jr.,

246

ZOOSYSTEMA • 1999 • 21 (2)

Comparative morphology of Brazilian Terebridae

MZSP, for commcncs un the tcxt. Dr Beu for the

important révision of chc manuscript. This study

was supportée! for research grain b y F'APESP

(Fundaçâo de Amparo a Pesquisa do Estado de

Sâo Paulo) Proc. # 96-6756-2.

REFERENCES

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(Gastropoda: Terebridae). Journal of Zoology 220: rnaculosa , a new species of Costellariidae (Gastro-

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accepted on 20 February 1999.

248

ZOOSYSTEMA • 1999 • 21 (2)

Pisidium chilense (d’Orbigny, 1846)

and new species of Pisidium C. Pfeiffer, 1821

from Southern Chile (Bivalvia, Sphaeriidae)

Cristiân Federico ITUARTE

Departamento Invertebrados. Museo de La Plata,

1900 La Plata, Buenos Aires (Argentina)

cituarte@museo.fcnym.unlp.edu.ar

Ituarte C. F. 1999. — Pisidium chilense (d'Orbigny, 1846) and new species of Pisidium

C. Pfeiffer, 1821 from Southern Chile (Bivalvia, Sphaeriidae). Zoosystema 21 (2) : 249-257.

KEY WORDS

Bivalvia,

Sphaeriidae,

Pisidium ,

new species,

South America,

Chile.

ABSTRACT

Pisidium chilense (d’Orbigny, 1846), a species only known from a rather poor

original description, is hcre fully redescribed and figured. Pisidium huillichum

n, sp, and Pisidium Ihwqilihuense n, sp. respectively from Llanquihue Lake,

CJiiloc Island and Yclcho Lake, Southern Chile, are described. Both new spe¬

cies share as common features the présence of only one demibranch, one

siphonal aperture and brood potiches developing from lhe upper and poste-

rior part ot inner demibranchs. As it was described for other Patagonian spe¬

cies, the ligament in both new species is internai, however, in Pisidium

llanquihuense n. sp. the ligament is slightly visible from the exterior.

MOTS CLÉS

Bivalves,

Sphaeriidae,

Pisidium,

nouvelles espèces,

Amérique du Sud,

Chili.

RÉSUMÉ

Pisidium chilense (d’Orbigny, 1846) et nouvelles espèces du genre Pisidium

C. Pfeiffer, 1821 du sud du Chili (Bivalvia, Sphaeriidae).

Pisidium chilense (d'Orbigny, 1846), espèce connue seulement par une des¬

cription originale brève et peu précise, est ici décrite et illustrée. Deux nou¬

velles espèces de bivalves de la famille Sphaeriidae. Pisidium huillichum n. sp.

et Pisidium llanquihuense n. sp., provenant respectivement du lac

Llanquihue, ile de Chiloé et du lac Yelcho, au sud du Chili, sont décrites. Les

nouvelles espèces ont comme caractères communs la présence d’une seule

paire de lames branchiales, interne (dans lesquelles les poches incubatrices se

développent vers le haut et en arrière) et une seule ouverture siphonale. De

plus, comme il a déjà été décrit pour d'autres espèces de Patagonie, le liga¬

ment est interne, bien que, dans le cas de Pisidium llanquihuense n. sp„ il soit

faiblement visible du dehors.

ZOOSYSTEMA • 1999 • 21 (2)

249

Ituarte C. F.

INTRODUCTION

There are a tew contributions dealing with

Pisidium C. Pfeiffer, 1821 species from Chile.

D’Orbigny (1846) described Pisidium chilense

(d’Orbigny, 1 846) from Concepcion, Mabille

(1884) described Pisidium lebruni (Mabille,

1884) from Punta Arenas and long afterwards,

Fig. 1. — Location map, collecting sites are indicated with an

asterisk.

Ituarte (1995) reported (or the first time

Pisidium observatimiis I’ilsbry, 1911 from Cabeza

de Mar, 48 km nortli Punta Arenas, Southern

Chile. The remaining reports on Sphaeriidae dis-

rributed in Chile correspond to the gcnus

Sphaerium Scopoli, 1 77 7- Sphaerium Lmrienchae

(Philippi, 1869) was reported from Ojos de

Ascman, Antofagasta Province (Kuiper N' Hinz

1984) and Sphaerium jbrbeii (Philippe 1869) has

been reported from Cota Corani (Ituarte 1995),

both records being from northern Chile.

Knowledge on other geographieally close sphae-

rjid (aimas from Southern South America east

Andes mountain range (mainly from Argentine

Patagonia) cornes (rom the work of Pilsbry

(1911), Ituarte &C Gordillo (1991) and Ituarte

(1996).

In the présent paper, Pisidium chilense is redescrib-

ed and properly figured. Moreovcr Pisidium

huillichum n, sp. and Pisidium llanquihuense

n. sp. from Southern Chile, are described.

MATERIALS AND METHODS

■Samples were collected by the author during a

field trip to Southern Chile, berween Dccember

1996 and Jamtary 1997 (Fig. I ). After collection,

the specimens were immediacely relaxed and

killed by immersion in warm water (50 °C

approx.) for a few minutes and then fi.xed and

prçserved in éthanol 80 n . Length measurements

[shell length (SL), shell height (SH), shcll widch

(SW) and pre-siphonal suture (PSS)l, shape

indices and morphometrlc ratios [heighc index

(1 SH/SL), convexirv index (Ci - SW/SH), ratio

hinge length (distance heeween cusps ôf left ante-

rior latéral and left posterior latéral): shell length

(HiL/SL)], were calculatcd according to the cri-

teria followed by Ituarte (1996).

For comparative purposes, 15 svntypes of

Pisidium lebruni Mabille, 1884, lodged at

Muséum national d'Histoire naturelle (MNHN),

Paris, and photographs of two synrypes of

Pisidium chilense (d’Orbigny, 1846) lodged ai the

Natüral History Muséum (NHM), London,

were also usçd.

Voucher specimens were deposited in the mala-

cological collections of the Museo de La Plata

250

ZOOSYSTEMA • 1999 ■ 21 (2)

Pisidium species from Southern Chile

Fig. 2. — Pisidium chilense (d'Orbigny, 1846) (MLP 5345);

A, right external view: B, posterior view of a Shell. Soale bars:

2 mm.

(MLP), Museo Argentine) de Ciencias Naturales,

Buenos Aires (MACN) and MNHN, Paris.

SYSTEMATICS

Pisidium chilense (d’Orbigny, 1846)

(Figs 2-4)

Cyclas chilense d’Orbigny, 1846: 568, pl. 83, figs 11-

13 .

Pisidium chiliense- Pilsbry 1911: 604.

Type kx.aü'IY. — According to ihe original descrip¬

tion by d’Orbigny (1846), the type localiry of

Pisidium chilense is Conception (36°50’S. 72°50’W),

Chile. However, soine confusion exists due to the fact

that the original label of the two syntypes deposited in

the NHM, London reads: “Valdivta, Chile” (39“49’S;

73°16’W) (a place approxiinately 450 km south of

Conception), as the origin of those specimens

(Fig. 1). The registration book of the NHM, l.ondon

malacological collection and Cray s (1855) lisr are

coïncident and report Valdivia as the type localiry for

P. chilense.

MafLKiAL CXAMINED. — Southern Chile. (Fig. 1)

Région de los Lagos, Yelcho l.ake, Puerto Cardenas

46 km south of’Chaitén (MLP 5345). — Outskirts of

Chaiién, unnamed brook on the Carretera Austral,

[Austral Road] (MLP 5381).

Redescription

(BASED UPON EXAMINEE) MATERIAL)

Shell ovate somewhat liigh (1 SH/SL about 76-

80%). not inflated [convexity index (Ci) is less

than 50%]. Maximum ohserved SL = 5.1 mm.

Anterior end modcrately protruded, posterior

end evenly curved. Dorsal margin evenly arcuate,

however, depressed immediately below beaks

(Fig. 3A-C); ventral margin evenly arcuate. The

point of jonction ol dorsal and anterior margin

faintlly tn.ukcd by an angle. Beaks vvide, low, not

exeeeding dorsal shell margin, slighdy visible

above dorsal margin, somewhat displaced post¬

eriori)' (located at about 60% of shell lengili).

Surface rather smooth, sulcated by low and line

concentrical ribs and very slightly marked radial

st ripes (Fig. 2B), periostracum straw-yellmvish,

dull shining.

Hinge plate rather solid, hinge line representing

about 64% of shell length. Hinge: in right valve

one club-shaped cardinal tnofh (Fig. 3A, B), the

surface of enlarged posterior end sculpuired by

multiple small shallow pirs (Fig. 4A), Latéral teeth

well-developed, anterior inner latéral tooth (A,)

long with nearly central cusp, posterior inner laté¬

ral tooth (P,) shorter with posteriorly displaced

cusp. Anterior and posterior outer teeth shorter

with distal cusps. In left valve, two cardinal teeth,

the outer (CJ longer, slender and evenly arcuate,

the inner (C,) wider and shorter, almost comple¬

tel y overlapped by C.,. Anterior latéral tooth (A 2 )

robust, with nearly central cusp, posterior latéral

tooth (P,) shorrer, straight with distal cusp,

Ligament-pit deep, with S-shaped ventral mar¬

gin, slender at anrerior end, broadened and mar-

kedly arcuate at the posterior two thirds

(Figs 3A-D; 4A). Escutcheon moderately mark-

ZOOSYSTEMA • 1999 • 21 (2)

251

Ituarte C. F.

Fig. 3. — Pisidium chilense (d'Orbigny, 1846) (MLP 5345);

A, hinge ol the right valve. B, detail of rigtit cardinal tooth. liga¬

ment and right latéral leeth; C, hinge ol the tett valve; D, detail of

the left cardinal teelh. ligament and posterlor latéral tooth. Scale

bars: 1 mm.

ed. Ligament robust, internai, however slightly

visible front outside as a slcnder stripc, never pro-

truded (Fig. 4B). Ligament length represents

about 21% ofshell length.

Five pairs of muscle scars, corresponding to the

inner radial mande muscles, almost coalescent

vvith the mantle line, except for the anterior pair

which is slightly apart.

An A ) GM Y

Only inner dentibranch présent; ascending lamella

shorrer, representing less rhan 50% of the descen-

ding one. Dorsal loop of the nephridia clefr. Brood

sacs developing in rhe uppei part of the pasterior

half ot the descendent lamellae of each inner demi-

branch. Anal aperture présent onlyt pre-siphonal

suture short (representing about 15% of SL), as

long or slightly shorrer titan siphonal diameter.

Remarks

The solid hinge plate of Pisidium chilense is a

characteristic shared with the remaining known

spccies front soutbetn Chile and Patagonia, such

as Pisidium magclltnicum (Dali, 1908), Pisidium

patngûmcum Pilsbry, 1911, Pisidium ohservationis

Pilsbrv, 1911, Pisidium p/enilunium (Melvill &

Sranden, 19071, Pisidium inucuyali Ituarte, 1996

and Pisidium lebrunl Mabille, 1884 (Ituarte

1995, 1996). The ligament could be considered

neither strictly external nor internai, rending

ver)' slightly to lie external but never protruding.

It can be seen front outside as a narrow line bet-

ween valves (Fig. 4B) as ir has been described by

Meier- Brook (1967) for Pisidium foreuse. Meier-

Brook, 1967 front Minas fierais, Brazil, and also

observed in tvvo new Pisidium species front

Corneilles and Misiones provinces, northeastern

Argc-ntlna, still utipublidted (Ituarte in press).

The spécimens ol Pisidium chilense front Yelcho

Lakc, Itère studied, show slightly lower shells and

beaks faindy more centrally located when com¬

parée! to the photograps of syntypes ai MNH,

London, When Pisidium chilense is cnmpared

with the syncypes ot Pisidium lebruni (MNHN,

Paris) from Puma Are nas, Chile, rhe latter pro-

ved to hâve higlier and more inflated shells, with

posterior end shorter and more abruptly trunca-

ted, and the anterior margin sloping markedly.

Kuipcr & Hinz (1984) hâve wrongly included

(with a question mark) P. chilense in the sytiony-

myc list ot Pisidium casertanum (Poli, 1791) (a

species curremlv known as cosTitopol irait) which

shows siriking dilferences such as nvo siphonal

apertures, both, inner and ourer, demibranchs,

and ligament decidedly internai.

252

ZOOSYSTEMA • 1999 • 21 (2)

Pisidium species from Southern Chile

Fig. 4. — Pisidium chilense (d'Orbigny, 1846) (MLP 5345):

A, detail of a latéral view of the ligament: B, dorsal view showing

the narrow portion ot the ligament exposed (arrow). Scale bar:

100 pm.

Fig. 5. — Pisidium huillichum n. sp. (MLP 5358); A. right extern

al view; B. posterior view ot a Shell. Scale bar: 1 mm.

Pisidium huillichum n. sp.

(Figs 5-7)

MATER!AL EXAMINE!). -— Southern Chile. (Fig. 1)

Région de los Lagos, Yelcho Lake, Puerto Cardenas

70 ni altitude, 46 km sont 11 ofChaitén, 1.1.1997, col.

Ituarte, holorvpe (MLP 5358), 2 paratypes

(MLP 5358, MNHN, Paris), 2 paratypes (MACN,

Buenos Aires). — l.lanquihue Lake, between 40°38'S,

72'-'3 1 ’W and 4 1 "20 S, 73"O’W, Puerto Üctay,

26.XII.1996, col. Ituarte, paratypes (MLP 5350).

Ltymoloi.y. — Lite spécifie narric rclers to Huil-

liclies (Indian word ihar meâns “souiliern peuple”), a

iribe of the Mapuchcs Indian people, who were the

ancient inhabitants of Southern lands in Chile.

Description

.Shell small (maximum SL = 2.45 mm), solid,

rounded oval outline, high (mean 1 SH/SL =

83 ± 2), inflated (average Ci = 73 ± 6). Beaks

wide, inflated, well visible above dorsal margin,

posteriorly displaced (beak position: 58-63% of

SL). Anterior end prorrttded in a sharp curve,

posterior end minette, widely arcuate. Anterior

half of dorsal margin laintly eurved, posterior

hall forming a markedly descendant curve.

Ventral margin evenly and strongly arcuate. Shell

transluceni. Surface glossy, straw-

yellowish, sculprured wirh well-marked, evenly

spaced, fine striae (Fig. 5A, B).

Flinge plare somewhat solid, hinge line rather

short, represenring about 50% of SL. Hinge: in

right valve one cardinal tooth (C-) slightly eurved,

slender at anterior half, broadened at posterior

end, forming a rounded sulcated small Itead.

Latéral teeth very strong, short, wirh markedly

rugose inner surfaces. Inner anterior latéral tooth

(A,J very large, wirh inner margin maikedly

eurved. Outer anterior latéral (A-,) minuscule

ZOOSYSTEMA • 1999 • 21 (2)

253

Fig. 6. — Pisidiun huillichum n sp, (MLP 5350); A. internai view

of a right valve; B. detail oi the hinge ot right valve; C, internai

view ot a lett valve; D detail of the hinge of left valve. Scale

bars: 1 mm.

(Fig. 6A, B). In right valve, two short cardinal

teeth, the otiter (C 4 ) a slender lamellae, overlap-

ping C, at posterior half or a little more. Latéral

Fig. 7. — Pisidium huillichum n. sp. (A. MLP 5350: B. MLP

5358); A, detail of right cardinal tooth, ligament and ligament-pit;

B. dorsal view showing the included ligamental pit. Scale bars:

A. 1 mm; B, 100 gm.

teeth very robust, short, high, with pointed

cusps, almost central in A,, displaced backwards

in P, (Fig. 6C, D).

I igamenr-pit encloscd, deep, with ventral margin

evenly curved (Fig. 7A). Escutcheon well-marked

(Fig. 7B). Ligament internai, relatively short,

representing about 19% of SL (Fig. 7).

Amatomy

Only inner demibranch présent; short ascending

lamellae not mttch longer than one fourth to one

rhird oflength of descending one. Brood sacs

developing tn the ttpper part ol the posterior half

of the descendent lamellae of inner demibranchs,

occupying more than three quarters of the demi¬

branch length, maximum nuinber of incubated

embryos found; 12 shelled larvae, measuring

abour 0.5 x 0.7 mm (minimum and maximum

diameters). (.7nly anal opening présent.

Presiphonal suture short to somewhat long (ave¬

rage PSS 18 ± 2% of SL), representing one to

254

ZOOSYSTEMA • 1999 • 21 (2)

Pisidium species from Southern Chile

one and a hall fold the diameter of the anal

opening. Dorsal loop of the nephridia, clefr.

Posterior adductor muscle s cars small, posterior

foot retractors powerful. Inner radial mantle

muscles weak, five to six nor well-deFined mus-

cular bundles (exccpr for the siphonal one),

attached almost onto the mantle line.

Remarks

Pisidium huillichum n. sp. can be easily distin-

guished from the remaining Chilean known spe¬

cies by the small-sized, relatively high and

inflated shell. The general shell shape and the

strongly ntarked surface striation are also quite

distinctive characters. The high number of

embryos in the brood sacs is also a peculiar fea-

ture. The presence of the internai ligament and

strong surface sculpture chtefly disringuishes

P. huillichum n. sp. from orher similar small-sized

South American Pisidium species such as

Pisidium vile Pilsbry, 1897 and Pisidium puncti-

ferum (Guppy, 1867). The latter also differs in

having higher and less inflated shell, and smaller,

lower and more centrally Jocated beaks.

Pisidium llanquihtiense n. sp.

(Figs 8-10)

MateriAI. EXAMINE». — Southern Chile. (Fig. 1)

Puerto Octay, north of Llanquihue Lake, 70 m altitu¬

de. between 40"38'S, 72°31'W and 4l°20'S, 73°0’\V,

26.XII.1996, col. Ituarte. holotype (MLP 5349),

2 paratypes (MI-P 5349, MNHN, Paris), 2 paratypes

(MACN, Buenos Aires). — Chiloé island, a small

water course at Notuco Bridge, on the national route

No. 5, imniediately befbre the branch road to Cucao

(on the Pacific coast), 26.XI1.I996, col. Ituarte, para-

types (MLP 5344),

EtyMOI.OGY. — The name refers to Llanquihue

Lake, the type locality.

Description

Shell of medium size (maximum observed size:

5.2 mm), ovate, tending to be sub-quadrangular,

rather liigb (mean I SI i/SL = 85 ± 1), somewhat

compressed (mean Ci = 62 ± .3), nearly équilatéral.

Beaks slightly displaced backwards (located at

about 56% ol the shell letigth), wide, low, tainrly

visible above dorsal rnargin (Figs 8; 9). Surlace

finely striated, white-yellowisb, silky-glossy.

Fig. 8. — Pisidium llanquihuense n. sp. (MLP 5349); A, right

external view; B, posterior view of a shell. Scale bars; 2 mm.

Hinge line evenly arcuate, sunken at the médian

point, just below central point of beaks, relatively

short (representing about 57-60% of SL), rather

solid. Hinge: in right valve an Y-shapcd cardinal

tootb (Cj) witb very short anteriot end, curvcd

and enlarged at posterior end lorming a deeply

sulcate triangular head; anterior and postetior

latéral leeth relatively short and robust, matkedly

rugosc (Figs 9A, B; J OA). IriJeft valve, outer car¬

dinal tootb long, oblique, almost straigfht, over-

lapping C, at posterior hall or slightly more,

inner cardinal tooth short, curve al anterior end,

diverging strikinglv from (Fig. 9C, D).

Anterior and posterior latéral teeth (A, and P 2 )

robust, cusps distal.

Ligament-pit deep, its ventral rnargin open S-

ZOOSYSTEMA • 1999 • 21 (2)

255

Ituarte C. F.

Fig. 9. — Pisidium llanquihuense n. sp. (MLP 5344); A, hinge of

the right valve; B. detail of right cardinal tooth, ligament and

posterior latéral teeth; C, hinge of the left valve, D, detail of left

cardinal teeth, ligament and posterior latéral tooth. Scale bars;

shaped, its dorsal wall is a thin calcareous laver

which in dried specimens can fracture in several

small plates (Fig. 1 OC). Ligament internai, how-

ever it is slightly exposed ro the exterior through

a narrow slit (Fig. 10B-D). In larger specimens,

the ligament might be protruding, but it is

ahvays covered by the calcic plates which form

the roof of the ligament-pit (Fig. 10D).

Rd. 10. — Pisidium llanquihuense n. sp (A B, MLP 5344; C, D,

MLP 5349): A, detail ot the nghl cardinal tooth, B, dorsal view

showing a narrow portion of the ligament visible (arrowl; C, dor¬

sal view of a Shell showing me small plates which form the dor¬

sal wall of the ligamental ph (arrow.); D, dorso latéral view of the

escutcheon. showing a strong ligament elevatlng the dorsal wall

ot the llgament-pir farrow). Scale bars: A, iû0 pm: B, 200 pm;

C, 0,5 mm; D, 1 mm.

256

ZOOSYSTEMA • 1999 • 21 (2)

Pisidium species from Southern Chile

Anatomy

Only the anal siphon présent. Pre-siphonal suture

short (representing about 13% of shell length),

approximatcly as long as the siphonal diameter.

Dorsal loop of the nephridia deeply cleft. Only

one (the inner) pair of demibranchs présent,

ascending lamellae shorter, representing about

one-third of the descencling one. Bood sacs deve-

loping postero-dorsally, maximum number of

incubated embryos found: eight shelled larvae

(1.2 mm length) in an adult of 5.2 mm of shell

length. Multiple (12-13) weak inner radial man¬

de muscles, besides a powerful siphonal retractor,

inserring not aparr Iront the mande line (howe-

ver, in para types ol the lot MI.P 5344, inner

radial muscles are well-dcveloped and muscle

scars very well-markcd, but always lying jList

above or in close contact vvith the mande line).

Remarks

Pisidium llanquibuense n. sp. may be easilv dis-

tinguished from other pisidia from Southern

Chile and Patagonia by irs high ovate, tending to

sub-quadrangular, shell shape. The type of liga¬

ment, internai but very siighrly exposed and the

presence of one detnibranch and one siphonal

aperture, place thiç species in the vicinity of

those from the Para n,4, Uruguay and Rio de La

Plata drainages such as Pisidium foreuse Meier-

Brook, 1967 and rwo vet unpublished new spe¬

cies from Comentes and Misiones provinces

(northeasrern Argcntina). Howevcr, the solidness

of shell and particularly that of the hinge plate

agréés with the pattern whtch is characteristic for

Patagonian species. I he short prc-siphonal suture

and the reduced litlcr size, are also distinctive.

The type of nephridia, the siphonal arrangement,

the number of demibranchs and position of

brood sacs, reminds one of those species inclu-

ded in the subgenus Afropisidiunu however, the

characteristics of the hgament-pit and the pecu-

liar position of the ligament arc discordant cha-

racters.

Acknowledgements

Dr P. Bouchet and Dr F. Naggs kindly contri-

buted by sending syntype specimens and photo-

graps of type materials housed at MNHN, Paris

and NHM, London, respectively. The fine work

of Lie. R. Urréjola from MLP scanning électron

microscopy unit is also appreciaicd. The author

is a researcher of the Consejo Nacional de

Investigaciones Cientificas y Tecnicas (CONI-

CET), Argentina.

REFERENCES

Gray J. E. 1855. — List of the shelh of South America

in die collection of the British Musetim, collected and

described by M. Alcide d'Orbigny. in the “Voyage

dam l'Amérique Méridionale'' , [Préfacé 1 net.

1854). I aylnr and Francis. London, 89 p.

Ituarte C. F, 1995. — Nuevo.s registres de Pisidium

Pfeiffer, 1821 y Sphaenum Scopoh, 1777 (Bivalvia:

Sphacriidae) en Chile, Bolivia y Noroeste argenti¬

ne. Nfotropica 41 ( 105-106): 31-41.

Ituarte C. F. 1996. — Argentine species of Pisidium

Pfeiffer, 1821, and Mutculium l.ink, 1807

(Bivalvia: Sphacriidae). The Veliger 39 (3):

189-203.

Ituarte C. F. in press. — Pisidium laragnyense and

Pisidium pi poème- new species boni northeastern

Argentina (Bivalvia: Sphaeriidae), The Veliger 42

(4).

Ituarte Ci. F. Ht Gordillo S. 1991. — Nuevas citas de

peledpodos dulciacuCcolas de lsla Gable, Ticrra del

Fuego, Argentina. Neotrnpica M (97): 29-30.

Kuiper J. G. J, & Hinz W. 1984. - Zur Fauna der

Kleinmuscheln in den Anden (Bivalvia: Sphaeri-

idae). Archiv fUr Mollusken kunde 114 (4-6):

137-156.

MabilleJ. 1884. —Notices m.il.icologiques. Bulletin

de la Société philomatique de Paris, série 7. 8: 39-49.

Meicr-îirook G. 1967. — Pisidium foreuse , a new spe¬

cies boni Brar.il (Mollusca: F.ufnniellihranchiata;

Sphaeriidae), Archiv für Hydrobiologic 64 ( I ):

63-68.

Orbigny A. D. d’ 1846. — Voyage dam l'Amérique

Méridionale exécuté pendant les années 1826-1833,

volume 5, part. 3 : Mollusques. P. Bertrand. Paris,

758 p.

Ptlsbry 11. A. 1897. — New species of mollusks from

Uruguay. Proceetlings of the Aeademy of Natural

Sciences of Philadelphia May 1897: 29Ô-298, 2 pis.

Pilsbrv Fl. A. 1911. — Non-marine Mollusca of

Patagonia. Reports of the Princeton University

Expédition to Patagonia (1896-1899) 3 (5): 513-633.

Submitted on 26 Mardi 1998;

accepted on 12 October 1998.

ZOOSYSTEMA • 1999 • 21 (2)

257

Taxonomie review of the elongated cockles:

généra Trachycardium, Vasticardium and

Acrosterigma (Mollusca, Cardiidae)

Jacques VIDAL

Laboratoire de Biologie des Invertébrés marins et Malacologie,

Muséum national d’Histoire naturelle

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

malaco@mnhn.fr

Vidal J. 1999. — Taxonomie review of the elongated cockles: généra Trachy¬

cardium, Vasticardium and Acrosterigma (Mollusca, Cardiidae). Zoosystema 21 (2) : 259-

335.

ABSTRACT

The cardiids of rhe subfamily Trachycardiinae Stewart, 1930 (sensu Keen,

1969, genus Pnpyridea excluded), are reviewed, wirh spécial attention given

to the généra Trachycardium , Acrosterigma, and Vasticardium. No change is

proposed here to the relatively well-dcfmed taxonomy of Trachycardium ,

considered to be exdusively American, with six subgenera, nor to the

American Acrosterigma, In contrast, the generic taxonomy of the lndo-Pacific

Trachycardiinae, quasi-randomly distributed by auihors among the three

généra cited above, was not clear and is reevaluated. Ail of the spccies are

regrouped here into two généra Vasticardium and Acrosterigma whïch receive

clear and ttsable définitions. The American genus Trachycardium difiers

widcly front them in both hinge and rib morphology. The two généra

Vasticardium and Acrosterigma are distinguished rnainly by rib morphology.

These three généra are now groupée! in the subfamily Cardiinae. In sevetal

previous articles, I hâve analyzed in detail the genus Vasticardium, including

fifteen Recent species. The results are summarized here. The genus

Acrosterigma is represented in America by several fossil species and two

Recent species; in the lndo-Pacific, where no general study r has previously

been undertaken, it is represented by several fossil species (one new) and

twenty-five Recent species, of which nine are new; these species are divided

into six species-groups. Neotypes are proposed for Cardium magnum Linné,

1758 and Cardium hiradiaturn Bruguière, 1789 and lectotypes for Cardium

laevigatum Linné, 1758, Cardium serratum Linné, 1758, and Cardium mar-

moreum Lamarck, 1819.

KEY WORDS

Mollusca,

Bivalvïa,

Trachycardium,

Vasticardium,

Acrosterigma.

ZOOSYSTEMA • 1999 • 21 (2)

259

Vidal J.

MOTS CLÉS

Mollusca,

Bivalvia,

Trachycardium,

Vasticardium ,

Acrosterigma.

RÉSUMÉ

Revue taxonomique des « coques allongées » : genres Trachycardium,

Vasticardium, Acrosterigma (Mollusca, Cardiidae).

Cette étude est une révision de la sous-famille des Trachycardiinae Stewart,

1930 (au sens de Keen, 1969, à l’exclusion du genre Papyridea). Une atten¬

tion spéciale est donnée aux genres Trachycardium., Vasticardium et

Acrosterigma. Aucun changement ne sera proposé à la taxonomie de

Trachycardium , considéré comme exclusivement américain et qui comprend

six sous-genres, ni aux Acrosterigma américains. Par contre, la taxonomie

générique des Trachycardiinae de l'Indo-Pacifique, répartis à peu près au

hasard par les auteurs dans les trois genres cités ci-dessus, a été reconsidérée.

Toutes ces espèces sont regroupées ici dans les deux genres Vasticardium et

Acrosterigma qui reçoivent des définirions claires er facilement utilisables. Le

genre américain Trachycardium diffère nettement d'eux par la charnière et

par la morphologie des côtes. Les deux genres Vasticardium er Acrosterigma

sont séparés essentiellement par la morphologie des côtes. Les trois genres

sont placés maintenant dans la sous-famille des Cardiinae. Dans plusieurs

articles, j'ai analysé en détail le genre Vasticardium qui comprend quinze

espèces vivantes ; les résultats de ces études sont sommairement indiqués. Le

genre Acrosterigma est représenté en Amérique par plusieurs espèces fossiles

et deux espèces actuelles ; dans l’Indo-Pacifique, où aucune étude générale

détaillée n’a encore été entreprise, le genre est représenté par plusieurs

espèces fossiles (dont une nouvelle) et vîngt-cînq espèces actuelles, dont

neuf sont nouvelles, et qui sont réparties dans six groupes-espèces. Cette

revue propose des néotypes pour Cardium magnum Linné, 1738 et

Cardium biradiatum Bruguière, 1789 a nsi que des lectotypes pour Cardium

laevigatum Linné, 1758, Cardium serratum Linné, 1758 et Cardium mar-

moreum Lamarck, 1819.

CONTENTS

Introduction .261

Material and methods .262

Genus Trachycardium .264

Genus Vasticardium .268

Genus Acrosterigma .271

Species-group of Acrosterigma

dalli .272

Acrosterigma pristipleura .272

Acrosterigma magnum .274

Acrosterigma bure hardi .276

Species-group of Acrosterigma

cygnorum ..277

Acrosterigma cygnorum .279

Acrosterigna rarement . 280

Acrosterigna kersLikae .281

Acrosterigma marielae .283

Acrosterigma abrolhense n. sp.284

260

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Species-group of Acrosterigma

variegatum .285

Acrosterigma variegatum .285

Acrosterigma oxygonum .288

Acrosterigma selene n. sp.289

Acrosterigma disais n. sp.290

Acrosterigma mauritianum .291

Species-group of Acrosterigma uniorna-

tum .293

Acrosterigma uniornatmn n. sp.293

Acrosterigmaprofundum n. sp.296

Acrosterigma arnirante n. sp.297

Acrosterigma su!iianam n. sp.298

Acrosterigma paulayi n. sp.299

Species-group of Acrosterigma

maculosurn .299

Acrosterigma maculosurn .299

Acrosterigma maculosurn howense .. 303

Acrosterigma itupolitum .305

Acrosterigma transcendent .308

Acrosterigma seurati n. sp.312

Acrosterigma dianthinum .313

Acrosterigma punctolineatum .314

Acrosterigma hobbsae n. sp.315

Acrosterigma simplex .318

Species-group of Acrosterigma

biradiatum .322

Acrosterigma biradiatum .323

Acrosterigma attenuatum .327

Acknowledgements .329

Index of spécifie and subspecific

taxa .329

References .331

INTRODUCTION

The species discussed here can be grouped under

the vernacular name “elongated cockles", because

most of them hâve height appréciable greater

rhan length with an ovoid more or less oblique

shape, characters that are exceptionally présent

together in other Cardiidae, Recent or fossil,

except for several species in the subfamily

Laeviçardiinae (Keen, 1951), which still awair

révision and cannot be treated in this paper. For

many authors, Trachycardimn , Vastiçardinm and

Acrosterigma constitute the basis of the subfamily

Trachycârdiinac. lt will be shown here that this

subfamily should no longer be utilized, as already

donc by sortie authors (Popov 1977; Kafanov &

Popov 1977; Schneider 1992, 1995).

When Stewart (1930; 271) defiiied the

Trachycârdiinae, thé given critcria of séparation

frorn the other subfamilies were limiied to the

si/.e of the cardinal teeth: “Cardinals verv un-

equal, the right anterior and left posrerior being

obscure’. If The two généra Dinocardia and

Cerastodertna (which Stewart considered as “pro-

bably dose to the Cardiinae”) are exduded, the

original composition ot rhe sublamily is reduced

to two généra: Irachycdrdium and Acrosterigma.

The genus l'rachycardium , represemed by nunte-

rous well-studicd American fossil and Recent

species, was divided by Stewart imo five sub-

genera, taxa still valid and in current use. On the

other hand, Stewart hesirated to give generic sta¬

tus to Acrosterigma (cOmprisilig-, at that time.

some fossils and two living species, ail

American), because he could not see any diffé¬

rence in che hinges, comparative to

Trachycardium. As far as lndo-Pacillc species are

concerned, Stewart admitted being unable to

place rhem in any genus, and left this question

open. h must be rccognizcd that, at that time,

Indo-Pacific species were far less well known

than the American species.

Iredale (1927: 75) had already created rhe genus

Vastiçardinm lor the Indo-Pacific species, bascd

mainly on geographical considérations; this

action did not bring anything new to rhe prci-

blem, nor did the new genus Regozttra introdu-

ced by ltim in 1936 (p. 275).

Keen (1969) at first grouped the Indo-Pacific

Trachycardiinac in the genus Acrosterigma (sub-

genera Regozam and Vastiçardinm), then (1980)

in two généra Acrosterigma and Vastiçardinm.

However, her crireria taken into account for the

définition ol rhese taxa were unclear and uneon-

vincing, and few authors followed rhese propo¬

sais. The majority of subséquent authors grouped

ZOOSYSTEMA • 1999 • 21 (2)

261

Vidal J.

the Indo-Pacific species, sometimes seentingly

more or less randomlÿ, in the genus-groups

Trachycardiunu Acrosterigmd \ Vcistiçardium and

Regozara , without accürate criteria of définition.

This situation lias neccntly heen sumniarized bv

Oliver & Chcsney (1997: 65), who stated: “The

generic subdivisions of the Trachycardiinae bave

always been debâtcd and there is little consisten-

cy in their use ’. Several other authors hâve also

admitted that the generic taxonomy for this

group is unsatisfacrory, for exarnple Maxwell

(1978: 20) who wrote “Interrelaiionships of the

various genus-group taxa that hâve been propo-

sed in the Trachycardiinae are far Irom cleai ".

Wilson & Stevenson (1977: 74) noted that “the

shell characrers on which the current classifica¬

tion of the subfamily Trachycardiinae is based,

are unSatisfactory”. They, however, rnade a (un-

damenta! observation: that Indo-Pacific

Trachycardiinae as well as the American

Acrosterigmd hâve cardinal teeth separated in the

right valve (Fig. 1 D, F), while these teeth are

fused together in the American Trachyciirdiutn

species (Fig. IA). They, rherefore, recognize two

généra in the Trachycardiinae: Trachyeardiutn s.s.

(santé as Stewart) and Acrosttftgtnu including the

original American species and ail the Indo-Pacific

Trachycardiinae. Wilson &C Stevenson treated the

genus Vdsticdrdium as a synonym of Acrosterigmd,

although they make another imcresting observa¬

tion (1977r 77): “We hâve observed otie sculptur¬

al character of possible generic significance: in

the smaller Australian species [six species cited],

the South-Fast Asian species art'ni col uni and Lhe

Japanese species bunhdrdi , the posterior ribs are

divided down their centres [...while...] in other

species the ribs are simple'. Thus, these authors

demonstrated one of the main criteria for separat-

ing Acrosterigmd front Vasticürdittm in the

Indo-Pacific.

The former group is also disringuishable by other

characters. as already noted by Powell (1958:

76): “These shells [ urcnicola-cygnorum group],

plus several other Indo-Pacific species, differ

from the massive 7 ra chycürdiurn and

Vdsticdrdium in their ligltter builcl, smaller si/.e

and actite flattened beaks, but they are Itéré

retained in Vasticardium , pending a better

understanditig of the tropical Pacific species”.

This “better understanding” required a general

study as detaîled as possible of the

Trachycardiinae sensu Keen, particularly of these

from the Indo-Pacific. I hâve undertaken, in

several préviens papers and in this article, to pré¬

sent this Study, confirming thaL Lhe Recent

Indo-Pacific Trachycardiinae can be distributed

between two generic groups: (1) rite group al¬

ready separated by Powell, and Wilson &

Stevenson, which has numerous affinitïes with

the American Acrosterigmd; (2) the generic group

Vdsticdrdium , having different characters.

MATERIAI. AND METHODS

The materia

muséums:

1 contes from the following

AMS

Australian Muséum, Sydney;

ANSP

Acadcmy of Natitral Sciences,

Philadelphia;

AIM

Auckland Inscitute and Muséum,

Auckland;

BPBM

Bernice P. Bishop Muséum,

Honolulu;

BMNH

l he Natural History Muséum,

London;

1RSNB

Institut royal des Sciences natu¬

relles de Belgique, Bruxelles;

LACM

Los Angeles County Muséum of

Natural 1 listory, Los Angeles;

Natal Muséum

Pietermaritzburg;

MHNG

Muséum d'Histoire naturelle de

Genève, Gencva;

MNHN

Muséum national d’FIistoire

naturelle, Paris;

MNZ

Muséum of New Zçalarid Te

Papa longarewa, Wellington;

NHMW

Naturhistorisches Muséum,

Vienna;

Queensland Muséum, Brisbane;

RMN H

Na normal Natuurhistor isch

Muséum, Leiden;

UGML

Unjvetsity of Guam Marine

Labo ra tory, Mangilao, Guam;

UMZ

üniversity Muséum of Zoology,

Cambridge;

USNM

National Muséum of Nacural

History, Washington D.C;

262

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

UUZM University Zoological Muséum,

Uppsala;

WAM Western Australian Muséum,

Perth;

ZMA Zoologisch Muséum, Amsterdam;

ZMUC Zoologisk Muséum, Copenhagen;

American fossil material has been examined in

ANSP and USNM (about sixty specimens of

twenty-four nominal species and subspecies,

including nineteen holotypes). Data on the

Australian and New Zcaland fossils were mainly

derived from the literature. Unfortunately inves¬

tigation of the other Indo-Pacific fossils was

more difficull, because of the poorer quality of

material and descriptions, and the difficulty or

impossibility of examining specimens.

Accordingly, only a few hypothetical suggestions

hâve been made about this field for the présent.

As far as methods of identification of taxa, spéci¬

fie and even generic, are concerned, detailed rib

morphology plays a prominent part in the analy¬

sis of the concerned gênera. I hâve already explain-

ed the elaborate character and the importance of

this rib morphology in several articles (notably

Vidal 1997a).

Measurements of shells concern;

H heighr, measured along the médian

rib;

L lengtli, measured parallel to the

hinge;

W width [when only one valve is avail-

able, the indicatcd width is extra-

pollated, and placed in brackets

01 ;

Ratio D measures the asymmetty of the

hinge; it is determined by dividing

the length of the line from the tip

of the utrtbo to the tip of the post-

erior latéral by the corresponding

distance from the umbo to the tip

of the anterior latéral;

Angle A is formed by two Unes joining the

laterals to the main cardinal in the

right valve, measured by taking a

print of the hinge on modelling

clay.

In the measurements tables, ~ means circa:

For description of rib morphology and its varia¬

tions, shells are divided into four “quarters”:

PQ posterior quarter;

MPQ medio-posterior quarter;

MAQ medio-anterior quarter;

AQ anterior quarter;

Longitudinally, shells arc divided schematically

into two parts, a ' juvénile" (or umbonal) part

and an “adule” (or marginal) part.

To avoid excessive!y long descriptions of rib

ornamentation some préfixes are used:

rccro-

concerning posterior flank or top

margin (e.g. retro-ruberculated or

rerro- cre n ulated) ;

pro-

concerning anterior flank or rop

margin (e.g. pro-ridged or pro-

festooned);

bi-

concerning both flanks or top mar¬

gins, but not the top zone irself

(e.g. bi-crenulated);

top-

concerning the lop zone of ribs

péri-

(e.g. top-ridged);

concerning ail the rib, top and

flanks (e.g. peri-ridged);

Some terms are used witli parricular meanings:

crenulated or festOoned are applied to the edges,

the margins of the top zone ot the ribs, vvhile

ridged or tuberculated describe the flanks or the

top zone ol the ribs;

interstices notchcd means thaï tliey are characteri-

zed by successive, regularly disposed small holes,

grooves, or norches, while striated encans sculptu-

red with very fine parallel concentric striae.

To simplify descriptions, “ncologisms” are utilized

also for some often-used descriptive éléments:

Crestal fold a longitudinal rib, more or less

individualized, at the apex of

some triangular ribs;

Pseudo-interstice the posterior scaled part of PQ

ribs can disappear on some

Acrosterigma species, tnerging

into the interstice wbich

bccomcs enlarged; the rib

becomes reduced to the scale-

less anterior part;

Sterigma “support’ internai médial rib

in the umbonal cavity, more or

less high ayd developed

(Fig. 2C);

Sublunule smooth ftrst part of AQ beside

ZOOSYSTEMA • 1999 • 21 (2)

263

Vidal J.

the lunule, without internai

marginal “ribbing”.

SYSTEMATICS

Family CARDIIDAE Lamarck, 1809

Subfamily CARDIINAE Lamarck, 1809

Genus Trachycardium Môrch, 1853

Trachycardium Môrch, 1853: 34

(introduced as a subgenus of Cardium).

Type species. — Cardium isocardia Linné, 1758, by

subsequenr désignation (Von Marrons, 1870: 586).

Distribution. — East and west coasrs of tropical

America, from Eocene to Recent.

iNCLUDll) SliBGENIIîA.- — Six subgenera, based on rib

morphology:

1) Subgenus Tracbycardium Môrch, 1853, with four

Iiving species [in ihe Pacific: T. comors (Sowerby in

Broderip & Sowerby, 1833) : in the Atlantic: T. iso-

cardia (Linné, 1758), 71 egmontianuni (Shuttleworth,

1856) and /', manueli Prado, 1993], and at least tour

fossil nominal species.

2) Subgenus Agnocardia Stewart, 1930, with at least

six fossil nominal species.

3) Subgenus l'hlogncardia Stewart, 1930, with one

Iiving species | /. belcbcri ( Broderip & Sowerby, 1829)

in the Pacific, Fig. 1H |. and at least four fossil nomi¬

nal species.

4) Subgenus Mcxicardia Stewart, 1930, with two

(probable synonym) Pacific Iiving species [71 proce-

rum (Sowerby in Broderip üc Sowerby, 1833) and

T. panamense (Sow r erby in Broderip & Sowerby,

1833)], and at least two fossil nominal species.

5) Subgenus Conilocardium Vokes, 1977, with one

fossil species.

6) Subgenus Dallocardia Stewart, 1930, with three

Iiving species 1 in the Pacific, 71 senticosum (Sowerby

in Broderip &c Sowerby, 1833) and T. quadragena-

rium (Conrad, 1837) ; in the Atlantic: 71 muricaturn

(Linné, 1758), Fig. Il], and at least nîne fossil nomi¬

nal species.

Remarks

Some authors bave rreated sortie of the above

subgenera as généra.

Most of Tracbycardium species hâve an elongared

ovoid shape. Llowever, some individuals in cer¬

tain species, particularly in tbe subgenus

Dallocardia, lose this characteristic, acquiring a

length équivalent to or even slightly greater than

the height (for example, Tracbycardium qitadrage-

narium can hâve L/H up to 1.10),

The genus Tracbycardium , as defined above, is

considered here as exclusively American. As far as

fossils are concerned, ! tbmk thaï tins genus was

erroneously utilized for groups of species from

outside America, for example in the Upper

Crctaceous of India, the Paleocene and Lower

Locene of Europe, the Neogene of the Pacific.

On the otlier hand some species front outside

America, placed or not in the genus

Tracbycardium, sharc numerous characters wdth

some species of this genus, particularly of the

subgenus Dallocardia:

1) That is the case of the West African species

Cardium ser ru latum Deshayes, 1855 and

Cardium eu parti Nicklès, 1955.

2) The European Neogene Cardium mitlticosta-

tutn Brocchi, 1843 is quasi identical to the

Miocène subspecics Tracbycardium ( Dallocardia)

dominicense hadranintm Woodring, 1982 (type

séries USNM 647469 and 647470). Brocchi

(1843: 313) wfutc: " Tanta c la conformait cbe ha

questo cardio /C. multicostatum/ col muricaturn

di l.innco, cbe io fui da principio tentait) di ris-

guardarlu conte nna sitnplice varie th di esso". Some

authors liave placed Brocchi’s species and several

related taxa in Tracbycardium . e.g. Sacco (1899:

41) who cited Trachycardium multicostatum

(Brocchi), Cossmann & Peyrot (1912: 473) who

cited Cardium ( Tracbycardium) multicostatum

(Brocchi); Rossi Ronchetti (1952: 70) vvlto cited

Laevicardium ( Tracbycardium) multicostatum

(Brocchi). Popov (1977) placed C. multicostatum,

together with threc others, in a nesv subgenus

Europicardiutn Popov, 197' 7 . Von Cosel (pers.

conim.) considers that this taxon is worrhy ol

genus status, plaeing in it the two above West

Africân species, Eu ropicardium sertit latum and

E. caparti.

3) The Indo-Pacific Vepricardium species also

share several characterisncs of some Dallocardia

(and Ettropicardium) species, including shape,

lunula, hrnge, rib ornamentation, présence of

pustules, etc. Conrad (1837: 230) described

C. qtutdragenarium as being “allied to C. as'tati-

CunT\ a commun Indo-Pacific Vepricardium spe¬

cies, and Sacco (1899: 41 ) stated that

T. multicostatum has affinities with T. muricaturn

264

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Fig, 1 . — A-C, Trachycardlum isocardia, right valve from Martinique. MNHN; A, right hinge; B. partial view ot PO and MPQ; C, detail

of AQ ornamentation: '‘horseshoe" scales, D, E, Acrosterigma magnum, right valve from Martinique, MNHN; D, right hinge: E, View

of PQ and three ribs of MPQ (on right): F. Vasticardium elongalum, hinge of a right valve Irom Phuket, MNHN; G, Vasticardium assi¬

mile, a left valve trom Al Flntas, Kuwait, Persian Gult, MNHN; view ot PQ and three ribs of MPQ (on lett); H, Trachycardium beicheri,

a right valve from Coiba Island, Panama. MNHN; partial view of PQ and MPQ; I, Trachycardium muricatum, a right valve from

Antilles. MNHN: partial view of PQ and MPQ. Scale bars: 10 mm.

ZOOSYSTEMA • 1999 • 21 (2)

265

Vidal J.

Table 1. — Comparative diagnosis of the genus Trachycardium with both généra Vasticardium and Acrosterigma.

Genus Trachycardium

Shell higher than long, subovate to elllptical, attenuate towards

umbones; often asymmetrical with anterior part dorsally raised

and posterior receding but obliquely more expanded and with

more or less marked truncation. Ribs otten curved backwards in

projection, with weak or absent posterior angulation.

Hinge shon, usually less than hait the length ol the Shell.

Cardinal area wide and moderately curved (Angle A between

140= and 160 ).

Teeth strong, although cardlnals are very unequal, right anterior

and left posterior bemg obscure. Cardinals in right valve connec-

ted by a high dorsal saddle (Fig. IA).

Ribs always high and strong, essentially ornated on posterior

flank.

PQ not or weakly contrasting with rest of Shell (Fig. 1 B, H, I).

On médian part ol shell. ribs ornated with coma-like scales to

laminae, pinned against posterior llank (Fig. IB, H, I); rib rarely

ornamented on anterior part (Fig. 1 B); top margins never serra-

ted. On anterior part, ribs often ornated with “horseshoe-like'’

scales (Fig. IC), never with transverse ridges.

Généra Vasticardium, Acrosterigma

Same diagnosis

Hinge generally short.

Cardinal area of variable width curved (Angle A between 85°

and 140').

Teelh otten strong; cardinals unequal, right anterior and left pos-

lerior being smaller. Cardinals in right valve sépara parated or

merely touching al their base, no dorsal saddle (Fig. 1D, F)

Ribs variable, sometimes low and weakly marked, mainly ornat¬

ed on top or top margins.

PQ contrasting with rest (Figs 1È, G 5F M) with tubercular

(Figs 1E 5F, 10E) or elongated lop scales (Figs IG. 2J).

On médian pari ol Shell, ribs ornated wilh top scales or

tubercles. top margins sériations (Fig. 14L) and flank ridges or

lubercles. never wilh posterior lamiriae.

On anterior part, nbs almosr always ftnely transversally ridged.

never with “horseshoe-like" scales.

Table 2. — Comparative diagnoses of Vasticardium and Acrosterigma.

Vasticardium

1. Shell large to medium, rarely small (small = H below

25 mm); one exception

2. Shell not dorsally tapered: angle A always above 125° (one

exception), up ta 140=

3. No internai umbonal ridge (sterigma); two exceptions.

4. No internai umbonal V-shaped coloured rays (three excep¬

tions); ventral margln often coloured.

5. Mean rib number low to medium (28.4-43.5)

6. Ribs generally high and well ornamented.

7. On PQ ribs high, squared. not longitudmally divided. inter¬

stices relatively wide (Figs IG, 2J).

Interstices relatively wide with respect to the ribs.

8. On juvénile médian and anterior parts, ribs enlarge quickly

following small smooth vary early shell, becoming high, well

ornamented and overhangmg. interstices in umbonal area. Ribs

rather more ornamented on juvénile than on adult parts.

9. Interstices sometimes striated, never hollowed or notched on

médian and anterior parts.

Acrosterigma

1. Shell medium to small, rarely large (large = H over 80 mm);

one exception.

2. Shell often dorsally tapered; angle A often below 125°, down

to 85 : (rarely reaching 135'’).

3. Several species with an umbonal ridge (sterigma).

4 Umbonal V-shaped coloured rays always présent: ventral

margin exceptîonally coloured.

5. Mean rib number sometimes medium (36.2-40.0), but more

otten high (44.2-62.8).

6. Ribs generally low and weakly ornamented.

7. On PQ ribs low. divided so lhat each has a smooth anterior

part and a scale or nodule bearing posterior pari, sepa rated by

a furrow wich can encroach into the posterior part between the

ornaments (Figs 1E: 2K 10E). Interstices thin, sometimes

contused with above funows.

8. On juvénile médian and anterior paris, ribs appear very gra-

dually following small, smooth very early shell, remaining low.

simple and smooth in umbonal area of adults. Ribs much less

ornamented on juvénile lhan on adult parts

9. Interstices sometimes hollowed or notched, never striated, on

médian and anterior parts.

266

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 3. — Statistical data (mean values).

Species

L/H

W/L

Ribs

Genus Vasticardium

elongalum

0.79

0.77

1.18

125°

37.9

fidele

0.77

0.84

0.95

125“

30.7

papuanum

0.84

0.74

=1.0

130°

36.9

godanii (Fos.)

0.78

0.83

1.15

135“

29.8

orbila

0.82

1.31

130“

42.2

luteomarginalum

0.82

0.80

1.21

135°

32.5

assimile

0.77

0.87

1.20

130°

33.4

rubicundum

0.81

0.84

1.10

135“

36.0

rhegminum

0.83

0.84

0.95

110"

42.2

thomassini

0.82

0.84

= 1.0

135°

43.5

flavum

0.90

0.75

=1.0

135“

29.0

pectiniforme

0.92

0.71

1.14

130”

30.6

vertebfatum

0.83

0.84

1.19

130“

28.4

ornatum

0.88

0.76

1.28

135°

29.0

angulatum

0.86

0.66

=1.0

125°

32.1

sewelli

0.89

0.78

=1.0

135”

38.2

Genus Acrosterigma

dalli (Fos.)

0.75

0.68

1.14

105°

35.0

prlstipleura

0.77

0.76

1.08

119”

35.0

magnum

0.79

0.76

1.16

123“

33.8

burchardi

0.81

0.67

0.86

106”

42 2

cygnorum

0.90

0.65

1.03

125°

42.6

sorenseni

0.94

0.56

0.91

121“

51.2

kerslakae

0.91

0.69

1.02

124”

40.9

marielae

0.90

0.62

1 13

128'

62.6

abrolbense n. sp.

0.99

0.56

1.00

130”

44.5

variegalum

0.91

0.67

0.84

128“

40.2

oxygonum

0.85

0.70

0.85

117°

38.9

selene n. sp.

0.94

0.66

122”

36.6

discus n. sp.

0.93

0.60

0.90

120”

56.3

mauritianum

0.89

0.71

0.95

118”

44.1

uniornatum n. sp.

0.83

0.82

1.00

128“

38.2

profundum n. sp.

0.82

0.79

1.03

122”

56.9

amiranten sp.

0.84

0.82

0.87

130”

53.5

suluanum n. sp.

0.83

0.80

0.95

130°

43.8

paulayin. sp. (Fos.)

0.86

0.77

1.27

130”

48.7

maculosum

0.84

0.71

1.02

115“

51.7

impolitum

0.85

0.70

1.04

118“

39.5

transcendens

0.87

0.73

0.98

123“

64.4

seurali n. sp.

0.89

0.72

1.03

127 e

62.8

dianthinum

0.89

0.69

1.00

125”

46.1

punctolineatum

0.86

0.73

1.04

122“

48.9

hobbsae n. sp.

0.88

0.72

0.86

128

61.9

simplex

0.85

0.80

1.07

119°

47.6

biradiatum

0.80

0.67

0.98

110°

50.9

attenuatum

0.66

0.68

1.03

88”

56.9

and T. fimbriatum (Wood, 1815), another cora-

mon Indo-Pacific Recent Vepricardium species.

4) The Atlantic South American species

Cardium delicatuLum Smith, 1915, with length

markedly superior to height (L/H = 1.12 in the

holotype), is also to be compared to Dallocardia

ZOOSYSTEMA • 1999 • 21 (2)

267

Vidal J.

and Vepricardium. Powell (1960: 182) and

Castellanos (1970: 231) placed it in the genus

Trachyeardium.

The conclusion of ail these remaries is that a pro¬

gressive variation txisrs from Trachycdrdium to

Vepricardium, with intermediate forms represent-

ed by Europicardium species. Consequently a

subfamily break cannot be placed beeween

Trachycdrdium and Vepricardium, as in Kecn's

classification. Accordingly Tracbvcardiinae is

interpreted as a synonym of Cardiinae, a conclu¬

sion reached by Katanov & Popov who placed

Trachycdrdium in the Cardiinae (1977: 31 1).

This papet will show that there is a more signifi-

cant break, withour any intermediate form, bet-

ween Trachycdrdium on the one hand, and

Vasticardium and Acrosterigma on the other.

These two lutter généra will, however, also be

placed in the Cardiinae.

Genus Vasticardium Iredale, 1927

Vasticardium Tredale, 1927: 75.

Type SPECIES. — Cardium elongatum Bruguière, 1789

(by original désignation).

Diagnoses. - - See Table 2.

Subdivisions. — l'he genus Vasticardium is divided

here into six species-groups. Further research, includ-

ing fossil species, might justify treating thèse

species-groups as formai subgenera.

Most species of this genus hâve already been revis-

ed (Vidal, 1991, 1992, 1993, 1996, 1997a,

1997b, 1998). Only a summary is given here,

with a short diagnosis of the different

species-groups. The important additional lots

and localities, observed after the issue of the

above papers, are mentioned.

Group of Vasticardium elongatum

IncîLÜDKD Recent species. — (sec Vidal 1992, 1993.

1996 for details):

1) Vasticardium elongatum (Bruguière, 1789)

[Synonyms: Cardium mode Sowerby, 1841a; Cardium

serricostatum Melvill & Standen, 1899; irachy-

edrdium okinaiodensc Kuroda, 1960; Trachycdrdium

wilsoni Voskuil & Onverwagt, 1991].

Six subspecies: elongatum: enodtr, wilsoni ; indioceanum

(Vidal, 1993): cipangense (Vidal, 1993); coralense

(Vidal, 1993).

2) Vasticardium fidele (Vidal, 1992).

3) Vasticardiumpapuanum Vidal, 1996.

Fos.xil SPECIES. — Vasticardium gortanii (Nardini,

1937), Pleistocenc, Red Sea.

Additional lus i kibu i ion data. — V. elongatum on

a beach of F Komodo, Indonesia (MNHN Vidal

1998). — V, fidele in Tuticorin, India (MNHN), in

Sri Lanka (AMS 047163). in Pliukct (Roussy private

cnil., MNHN Vidal), in far north ot Zululand, South

Africa (Natal Muséum), in Singapore (MNHN

Vidal), in the Philippines ÇLACM 90013, USNM

230318) and in Wallis and Futuna Tcrritory area

(MNHN, MÜSORSTOM 7 c-ampaign 1992). -

V, papuanum tn Sibuko Bay, Bornéo (USNM

239129).

DlAGNOSIS. — Shclls large to very large, markedly

elongated, sometimes asymmetrical and expanded

baekwards; lunule small. Foundation of anterior teeth

receding (not “hooked”). On PQ. ribi top-scaled with

straiglit to sltghtly twisted main scales, and with

seenndary sériations or seales on botlt edges of top;

elsewhere, ribs high and square-sided, overhanging

interstices, tops srnooth and bi-crenulated. except on

AQ whcre ribs are top-rîdged; interstices srnooth (one

exception).

Group of Vasticardium orbita

INCLUDKD RECENT SPECIES. — (see Vidal, 1997a for

details):

1) Vasticardium orbita (Broderip & Sowerby, 1833)

[Synonyms: Cardium mendanaense Sowerby, 1897;

Cardium philipptnense Hedley, 1890; Cardium pseu-

doanguialum Lnilow, 1905; Trachyeardium hawaiensls

Dali, Bartsch & Rehder, 1938], Four subspecies: orbi¬

ta ; mendanaense: ph/lippinense: huioaiemis.

2) Vasticardium Inteoinatginaruiu (Voskuil &

Onverwagt, 1991) (Synonym: Trachyeardium oiareru-

brttm Voskuil & Onverwagt, 1991 ). Three subspecies:

luteornargmatum ; marerubrum ; insu/are (Vidal,

1997a).

AtUtl LIGNAI DISTRIBUTION DATA. — V, orbita philip-

pinense on beaches of NW Lombok, N Sumbawa,

F Komodo, Tndonesia (MNHN Vidal 1998). —

K lutcomarglnatum insulure in Southern Mozambique

and northern Zululand (Natal Muséum). —

V. papuanum in Palau (UGMF).

DlAGNOSIS. — Shclls large to verv large, moderately

elongated, olten poscertorly expanded and "evinged”;

lunule rather large, with a raised margin. Foundation

268

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

of anterior latéral tecth "liooked . On PQ ribs squared,

with twisted ro conical main seules and possibly

secondary lamellar thin seules ar their anterior margin.

Elsewhere, ribs squared to slighllv rounded and bi-

crenulated, wirli or wirhout tbe équivalent of PQ top

main seules, ridged and somenmes “herringboned” on

anterior part ofshell. Interstices strongly strîated.

Group of Vasticardium assimile

INCLUDED RECENT SPECIES. — (see Vidal 1998 for

details);

1) Vasticardium assimile (Reeve, 1844) (Synonym:

Cardium lacunasum Reevc, 1844), I wo subspecies:

assimile and iacuwsum.

2) Vasticardium rubicundum (Reevc, 1844)

[Synonyms: Cardium mindaneme Reeve, 1844;

Vasticardium compunctum K ira, 1959; Acrosterigma

kmgdluorum Voskuil & Onverwagt, 1992],

3) Vasticardium rhegminum (Oliver & Chesney,

1997),

4) Vasticardium ihnmasüni Vidal, 1998.

ADDITIONAI DISTRIBUTION DATA. — V. rubicundum

on beaches of NW Lombok, Indonesia (MNHN

Vidal 1989). — V. thomassini abundant off far nortb

coast of Zululand, Sourh Africa, 45-78 m (Natal

Museurn).

Diagnoses. — Shells medium, rarely large, about

équilatéral and variably elongated, Foundation ol

latéral teetli almost “hooked . On PQ, ribs low, with

shurp anterior edges; iherc are top rubrreleo or seules

but no secondary latéral seules or serrations.

Elsewhere. ribs high and square-sided, sliglitly

top-scaled or ridged, sometimes hernngboned, with

serrated edges and often beaded or ridged flanks.

Shells rend to be brightly coloured, but lusterless.

Remarks

This species-group is the closest to Acrosterigma,

in characters such as: possible sterigma, double

umbonal coloured ray, curved hinge, flattish and

rarely dividcd ribs on PQ, with no secondary

marginal ornamentation.

Group of Vasticardium flavum

Regozara Iredale, 1936

INCLUDED RECENT SPECIES. — (see Vidal 1997b for

details):

I) Vasticardium flavum (Linné, 1758) [Synonyms:

Cardium fucatum Spengler, 1799; Cardium subrugo-

sum Sowerby*, 1838; Cardium dupuchense Reeve,

1845; Cardium gratiosum Deshayes, 1855; Cardium

tumidum Deshayes, I855|, Three subspecies: flavum',

subrugmwm dupuchense.

2) Vasticardiumpeetiniforme (Born, 1780) [Synonyms:

Cardium régulare Bruguière, 1789; Cardium rugosum

Lamarek. 1819; Tracbycardium peregrinum

Jousseaume, 1888; Vasticardium mgropunctatum Habe

& Kosugc. 19661.

3) Vasticardium vertebrarum (Jouas, 1844) ISyno

nyms; Cardium reeveanttm Dunker, 1852; Reposant

olivifer Iredale, 1936],

4) Vasticardium orruitum (Sowerby, 1877) [Synonyms;

Cardium fvltoni Sowerbv, 1916; Acrosterigtna sower-

byurum Voskuil & Onverwagt, 1992[.

AddiTIONAE DISTRIBUTION DMA. — K vertebratum

in java; 5°41,5’S, I05°37’E (ZMUC).

Diagnoses. - Shells medium-sized, often symmetri-

cal, sometimes sliglitly expandcd posreriorly, but littlc

elongated: lunule variable in widtb and depth. On

PQ, ribs squared with major slightlv oblique top

scales and secondary small scales or serrations on botit

edges (see Fig. 2J). Elsewhere, ribs rounded to trape-

z.oidal: they may be peri-ridged. with top ridges that

may hâve a critshed appearance, oi tidges may be limic-

ed ro sides of ribs. Interstices striated.

Group ol Vasticardium angulatum

INCLUDED RECENT SPECIES. — (see Vidal 1991 for

details):

1) Vasticardium angulatum (I.autarcie, 1819)

[Synonym: Cardium alternation Sowerby, 1840], This

species is distinguishable front ail other Recent

Vasticaidium and Acrosterigma species by ils gaping

valves; triaogular ribs are also rallier uncommon iil

l ’asticardium species.

Diagnosis. — Shells large, oblique]y ovoid, very

asymmetrical, and gaping aiucriorly and posteriorly;

lunule very small. ( )n PQ, ribs rriangqlar; top-seul es a

little twisted and almost radially disposed, with nr,

secondary scales. On MPQ, ribs rriangular and

retro-ridged, bccommg rounded and top-ritlged on

anterior part ol shell. Interstices smooth with a round¬

ed régulât riblet.

Group of Vasticardium sewelli

Inci DDKl) Rf.CE.NT SPECIES. — (see Ter Poorren 1997

for details),

I ) Vastirardium sewelli (Prashad, 1932) [Synonym:

Cardium laddt Abjatd, 1946, an Upper .Mioetne fossil

from Epi Island (Vanuatu)]. The single species of this

group bas a very distinctive rib morphology; tbe top

curved scales resemble those of Trachycurdium tsoedr-

dia (but not fornting laminae pinned against posterior

flank of ribs), and tbe top and latéral ornanients of the

ZOOSYSTEMA • 1999 • 21 (2)

269

Vidal J.

Table 4. — Comparative diagnoses of the species-groups of Acrosterigma datli, A. cygnorum and A. variegatum.

A. dalli species-group

A. cygnorum

species-group

A. variegatum

species-group

Dimensions shape

Medium to large.

Somewhat elongated, point-

ed. ovoid and équilatéral

(one exception).

Small to medium, Little

elongated: moderately

pointed, slightly inequl-

latéral (often bi-straighten-

ed posleriorly).

Medium: slightly elongated

and pointed; symmetry

variable.

Lunule

Little marked.

Well-marked, slightly

depressed on both sides.

Well marked, often depres¬

sed in one or both sides.

Hinge and interior

Hinge variably asymmetri¬

cal. markedly angled (A

105 -123").

A sterigma in one species

In addition to génotype.

Hinge symmetrical: mode¬

rately angled (A 121-130).

A sterigma présent in two

species

Hinge asymmetrical (D

0.66-0.95): moderately

angled (A 117 -128").

A sterigma in one species.

PQ rib morphology

Anterior part of ribs wide,

posterior very reduced;

scales small. irregular to

nearly triangular, almost

longiludinally disposed,

somewhat connected-

Both parts of ribs of équiva¬

lent width; scales elongated

fo "herringboned", slightly

irregular, slightly connected

fogether.

Both parts of ribs of

variabie width; oblique

scales with regular elonga¬

ted ovoid shape. not

connected logether.

Last ribs of MPQ

Flatly rounded, smooth to

slightly retro-tuberculated

Rounded to subtriangular,

retro-ridged.

Triangular with a crestal

fold, smooth to retro-ridged.

Médian part of Shell

Ribs low, fiat, bicrenulated,

overhanging thin inter¬

stices.

Ribs variable: flat-rounded

to squared. bl-tuberculated.

becoming top-ridged.

Ribs variably triangular to

rounded. often finely retro-

ridged (one exception).

AQ rib morphology

Same fiat ribs as above,

may or may not become

top-ridged.

Same ribs as above,

becoming top-ridged.

Same ribs as above,

becoming top-ridged.

First ribs of AQ near

lunule

First one or two ribs widen,

swell, loosing ornaments.

No change of ribs: clearly

deflned limit with lunule.

No change of ribs (one

exception).

ribs are typical of Vasticardium (Fig. 14L), as is the

hinge.

Additionai. DISTRIBUTION DATA. — Phuket (coll.

Roussy). — Guam, Hapra Harbour (USNM 849694),

Oca Point (USNM 851281). — Santo, Vanuatu

(USNM 769249).

Diagnosis. — Shells medium-sized, little elongated,

and almOst equilateral. Foundation ol latéral teeth

receding. Ribs high, rounded ro square sided, and

regularly ornamented over entire shell on top of ribs

with tltin, slightly oblique, curved to U-shaped scales

longer cm the posterior side of ribs, becoming spatub-

form on anterior part nfshell. In addition to these top

ornaments, ribs in juvénile shells serrated or finely ridg-

ed on both sides. Interstices finely striated.

Other fossil species (not assigned

to species-groups)

According to literature data, severai fossil fortns

front West-Indonesia and Burma seent to be

close to Vasticardium , for example:

Cardiuin subalternatum Jenkins, 1864: 60, pl. 7,

fig. 7A, B. Miocène of Java. Comparée! by its

author with C. alternatum.

Cardinal protosuhrugosum Noetling, 1901: 179,

pl. 10, figs 10-11. Miocène of Burma. Compared

by Pannckoek (1936: 71) with spolongensc and

sedanense.

Cardium ( Trachycardium ) spolongense Martin,

270

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of rhe elongated cocldes

Table 5, — Comparative diagnoses of the secies-groups of Acrosterigma uniornatum, 4. maculosum and A biradiatum.

A. uniornatum

species-group

Dimensions Shape

Small to medium, little elon¬

gated. moderately polnted;

ovoid and équilatéral;

rather tumid

Lunule

Well marked and delineat-

ôd, sllghtly hollowed.

Hinge and interior

Rather symmetrical, mode¬

rately angled (A 122 e -

130 ); cardinals sllghtly

connected In rlght valve.

No sterigma.

PQ rib morphology

Anterior smooth part

variable; long, thin scales

regularly obliquely placed,

not connected, encroachlng

upon anterior part.

Last ribs of MPQ

Médian part of Shell

Ribs low, flatly triangular;

same scales as on PQ.

disappearing or not.

Ribs variable, becomlng

relro-rldged, and finely pro-

ridged (two species).

AQ rib morphology

Same ribs becoming peri-

ridged, sometimes herring-

boned.

First ribs of AQ near lunule

Become slightly tubercular:

clear limit with lunule.

A. maculosum A. biradiatum

species-group species-group

Medium, ra'ther elongated Medium: variably elongated

and polnted, variably and polnted; often inequiia-

inequilateral (otten bi- or tri- teral (bi- rarely tri-straighten-

straightened posteriorly) ed).

Generally small and not de Small to bardly marked;

pressed; poorly delirteated. confused with sublunule.

Hinge symmelrical, angled Nearly symmetrlcal; mar-

(A 115 ,3 -128°). A sterigma kedly angled (A 88 e -110°).

in type species A sterigma in both species

Both parts of ribs more or Bipartition of ribs with

less équivalent; scales scales only on juvéniles: in

variable, rather tubercular adults ribs smoothed but

and irregular. separated. distinct, with wide pseudo¬

interstices: two last ribs

remain prominent.

Flatly rounded to subtrian- Very low but always mark-

gular, retro-ridged ed, wider than those more

Same as above, but some Ribs very low, close-set,

what retrocrenulated. marked and retro-ridged, or

only visible by colours.

Same ribs as above, beco- one third as above, one

ming top-ridged. Concentric third with few distinct ribs

alignmentsorndges, and striae: first third as

below. Concentric ridges.

First ribs degenerate; top Smooth sublunule in about

ridges become tubercular, third of AQ: no ribs, no

concentric teatures appear. marginal Internai crenula-

tions

1916; 266, pl. 4, figs 107-109. Lower Miocene

of Java. Shape and ribbing seem rvpical of

Vosticnrdiim.

Cardium ( F.ucardium) talahabeme Martin, 1922:

484, pl. 61, ftg. 105, A, Lower Miocene of Java.

Shape and ribbing seem typical of Vasticardium.

Cardium ( Imchycardium) sedanense Pannekoek,

1936: 72, pl, 4, fig. 51, A. Lower Miocene of

Java. Elongated shape, strong rib sculpture; com-

pared with spohugenSt by its aurhor.

Cardium (Acant/jocardia) dm tiras tu la tu m Beets,

1941: 163. pl. 8, figs 319-326, Upper Miocene

of Bornéo. Compared with lacunosum , flavurn ,

elongatum, fultoni by its author.

In addition to chese cxclusively fossil species, some

living species are also cited from the Neogene, for

example, by Martin, C. elongatum and C. rugosum

(1883: 245-246) and C. dupuebnesse [r/cj (1879:

106), in the “Tertiary Schists of Java”.

Genus Acrosterigma Dali, 1900

Acrosterigma Dali, 1900b: 1073, 1090, introduced as a

section of subgenus Trachycardium, genus Cardium',

only one species is cited by this aurhor.

Type SPECIES. — Cardium dalli Heilprin, 1887 (by

original désignation: 131, fig. 70).

ZOOSYSTEMA • 1999 • 21 (2)

271

Vidal J.

FOSSIL RECXIBOS. — I Itéré are no records nf fossils of

Acrosterïgma species before the Miocène, The genus

Trachycardium is geo1ogjc.il!)' older. alre.idy rcprésen¬

té d in the Oligocène .nid possibly liocene of America.

Several fossil American Acrosterïgma specics hâve been

described (see bclovv). Sonie Australien fossil spedes

undoubtedly bclong herc (sec bclovv). Less ccnainly,

some Japanese fossil specics may also bclong herc, for

example:

Cardium tokyoensis Tokunaga, 1906: 51, pl. 3.

fig. 12A, A , Pliocène of Tokyo. Compared to

Acrosterigrna burebardi by its author, but said to be

doser tii Acrosterignut unicnior

Vasticardium ntukai Hatai & Nisiyama, 1952: 35.

Previouslv described as Cardium bttrehardi Yokoyama,

1925: 120, pl. 14, fia. 9. from the Oligocène Ôgano

Formation and citedliy Yokoyama (1926: 134, pl. 19,

figs 3; 4) from dte Miocene Kanomarazarawa Forma¬

tion.

Vasticardium kantoense K.mno. 1958: 176, pi. 2,

fig. 4A, B. From Nagura Formation (Lower

Miocène). I lescribcd as closely relatcd to V. artnicala.

Vasticardium arcnicoioides Akutsu, 1964: 284, pl. 59,

figs 6; 7. Miocene of sonth and central Japan.

Compared to arénicole1 and burebardi by its author.

Trachycardium. butchttrdi and 71 umeuhr are also citcd

as Neogene fossils from Taiwan (Gu /.hi-wei et al.

1976: 70).

In Europe, Cardium ( Trachycardium) (raternum

Mayer, 1864: 356, from the l.ovver Miocene ol South¬

west France, probably belongs to the genus Acro-

sterigma, according to the description and figures of

Cosmann & Peyrot (1912: 502, pl. 22, figs 34 and

38-42); ir is the same for C. proecedens Mayer, 1858,

Oligocène, same area.

DlAGNOSIS. — See Table 2.

Subdivisions. - The genus Acrosterigrna is subdivid-

ed here into six species-groups. Further research,

including fossil species, might jusrify treating these

species-groups as formai subgenerit.

Species-group ol Acrosterigrna dalli

(Heilprin, 1877)

Included SPRC1F.S. — 1) Living spedes: A. pristipleu-

ra (American Pacific); A. magnum (American

Atlantic); A. burebardi (Japan).

2) American fossil species: No synthetic study has ever

been done on this group of shells. Having examined

almost ali the American type material in ANSP and

USNM, 1 wi|l limit in y self tu ttientioning the publish-

ed nominal specics:

A. dalli (Heilprin, 1887), from CaloosahatchLe

Formation, Florida. Pliocène.

A. hoerieomm Vokes, 1977, from Chipola Formation,

Florida. Miocene.

A. iucouspicuum (Guppy, 1866), from Bowden,

Jatnaica. Miocene (see Woodring, 1925).

A. watlandi (Woodring, 1925), from Btnvden,

Jatnaica. Miocene.

A. liuguatigris (Maury, 1917), from Cercado and

Gnrabo Formations, Dominion Republic. Neogene

(sec Vokes, 1989).

A. déclive (Gabb, 1881), from Costa Rica. Pliocène.

DlAGNOSIS. — See Table 4.

Rémarks

Cardium dalli is by far the larges: of ail the

Acrosterigrna species, reaclting a height of

137 mm Marty specimens hâve an internai ele-

vated mcdial ri b, radiaring from the urahonal

cavity (Fig. 2C), from which the name

Acrosterigrna \- umbonal support) stems. But this

"sterigma" is rather inconstant in the présent spe-

cies and has never been found in any spedmen of

the orher American fossil species of the genus,

nor in the rvvcj living American species. For this

rcason it has been considered as of no svstematic

value by several aurhors. Neverthcless it is pré¬

sent, in rather attenuated form, in several living

lndo-Wesc Pacific Acrosterigrna species.

Acrosterïgma dalli (Fig. 2A-C) is very close to

A. pristipleura (Fig. 2D, Fl) from whîch it differs

by its larger size, more flattened posterior part,

wider First ribs beside the lunule, more rounded

rlbs, and wider interstices,

Acrosterigrna pristipleura (Dali, 1900)

(Fig. 2D, E; Table 6)

Cardium (Trachycardium)pristipleura Dali, 1900a: 389.

Cardium maculosum Sowerby in Broderip & Sowcrby,

183.3: 85; Sowerby, 1834, fig. 18. Not C. maculosum

Wood, 1815.

Cardium maeulatum Sowerby, 1841b: 4. Not

C. maeulatum Gmelin. 1791: 3255 [= Cardium rnbus-

tum Solander, 1786). Nomen novum for C. maculosum

Sowerby.

Cardium bornelli Tonilin, 1928: 194. Nomen novum

for C. maeulatum Sowerby.

Types. — Cardium pristipleura : types not rraced.

Cardium maculosum Sowerby; titrée s bel U in RMN H,

Cutning collection, front 1res Marias Islands, Mexico.

The smallest nvo fit the figure and the dimensions

given by Sowerby, and Reeve’s figure (1844: fig. 58).

1 lie largest was selectcd as Icctotvpe of Cardium

maculosum Sowerby by Vnskuil 6c Onverwagt (1991:

68, pl. 3, fig. 5) and registered BMNH 1991.043/1.

272

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Fig. 2. — A-C, Acrosterigma dalll, specimen Irom La Belle, Hendry Cty. Florida, MNHN. H. E. Vokes gift; dimensions: 109.1 x 71.7 x

55.6 mm. with 34 ribs: C. umDonal cavily witb a sterigma: D. E, Acrosterigma pristipleura , lectotype ol Cardium maculosum Sowerby;

F, G, Acrosterigma leucostoma, lectotype. neotype of Cardium magnum Linné: H. I, Acrosterigma burchardi, specimen Irom Kyushu,

Japan, MNHN J. Vasticardium pectinitorme, specimen from Nouméa, New-Caledoma, MNHN, détail of PQ and lirsl rib ol MPO: K,

Acrosterigma cygnorum, specimen (rom Revesly Island, South Auslralia; detail ol PQ and First rib o( MPQ. Scale bars: A, B, D-l,

20 mm; C, 25 mm; J, K, 5 mm

ZOOSYSTEMA • 1999

Vidal J.

Table 6, — Measurements (in mm) and rib count of Acrosterigma pristipleura (Dali. 1900).

L/H

W/L

A°

Ribs

Lectotype maculosum Sow.

70.3

53.6

44.0

0.76

0.82

1.06

120

Syntype idem

64.0

48.7

36.8

0.76

Syntype idem

63.3

51.0

39.0

0.81

0,76

MNHN, Monterey?

50.5

41.0

31.1

0.81

0.76

125

MNHN, off Coiba

52.0

41.1

33.4

0.79

0.81

0.98

120

LACM 129-34

73.0

57.0

(40.0)

0.78

0.70

120

LACM 86-29

59.0

44.2

(36.4)

0.75

0.82

1.14

LACM 50904

84.6

64.7

(46.0)

0.76

0.71

115

LACM 50904

87.3

66.3

(47.4)

0.76

0.71

—1.0

Total adult shells measured and rib counts

General mean values

0.77

0.76

1.08

119

35.0

Standard déviation

0.03

0.04

0.08

3.7

2.1

Largest specimen observed. LACM 50904 (see above).

Materiai. EXAMINER. — The folJowing lots in addi¬

tion to the above cited type materiai:

Mexico. Gull ol California (USNM 152227). —

Bahia Magdalena, 18 m (LACM 129-34). — Très

Marias Islands (LACM 38-4), — Barra de Navidad

(LACM 68-41). — Socorro Island, 26-33 m (LACM

129-34).. — Clarion Island (LACM 38-10). —

Acapulco (MN1 IN Dutaillv 1850).

Costa Rica. Colto de Papaguayo (LACM 86-29,

1986). — Cano Idand (LACM 72-63, 1072),

Panama. Al'uera Island (MNHN Vidal). — Coiba

Island (MNHN Vidal).

Ecuador. Playas de Atacarnes (Esmeraldas) (MNHN

Holfstetter 1056), — M.mta (MNHN Hoffsicrter

i 956).

Dubious and wrong localitics. Monterey, California

(MNHN du Petit Thouars 1839). — Philippines

(LACM 50904 Burch).

Distribution. — Southern portion of Culf of Cali¬

fornia to Ecuador; approximately from northern tropics

to equator, cxtcnding along about 3800 km of coasts.

Description

Shells medium to (rarely) large, ovoid to

“pear-shaped", rarely elliptical, and équilatéral

with possible weak iruncation on posterior mar-

gin. Always elongated in adult stage: mean

L/H = 0.77 (range 0.70-0.84); width rather

variable, mean W/L - 0.76 (range 0,70-0.82).

Ribs straight or slighdy curved in projection,

l.unular area rather large.

Exterior well-colouted, with orange, pink, brown

and/or purple; interior white, with posterior

margin generally orange and ventral and anterior

margins purple. Hinge line moderately arched in

adults, mean < A = 119° (range 115°-125°) and

symmetrica'l (ratio D close to 1). No stcrigma

observed.

Mean rib number35,0 (range31-4l ).

Rib morphology: on PQ anterior part o( ribs

wide, posterior reduced wirh very small scales.

On médian part of shell, ribs low, fiat, overhang-

ing thin interstices; botforn of interstices on

juvénile shells regularly notched by elongated or

crescenr-shaped notches, which continue on

lower sides of ribs; contact between interstices

and ribs marked, on both sides, by a very chin

longitudinal depressed line. As in A. magnum

(see below), the last rib of MPQ, rnainly on right

valve, can bear a longitudinal gash on its anterior

part.

Remarks

Acrosterigma pristipleura conforms with the spe-

cies-group diagnosis as far as rib morphology is

concc-rned, and is very dose ro the genotvpe

A. dalli, in chat its ribs are “flattened and pecu-

liarly approximated, interstices, deep narrow

curs'’ (Reeve 1844, Sp. 58). Acrosterigma pristi¬

pleura, when adult, is easily distinguishablc from

A, magnum , the other living représentative of the

species-group in the Americas, rnainly by its fiat

low ribs and very thin interstices, h is a rather

uncommon species, little cited in the literature.

274

ZOOSYSTEMA ■ 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 7. — Measurements (in mm) and rib count of Acrosterigma magnum (Linné. 1758).

W/L

A°

Ribs

Lectotype leucostoma

64.2

51.8

38.6

0.81

0.75

1.18

120

Lectotype marmoreum

57,6

47,0

38.0

0.82

0.81

1 29

125

Possible syntype subelongatum

73.0

55.0

44.0

0.75

0.80

1,12

125

MNHN, Lamarck coll.

73.0

61.2

47.1

0.84

0.77

1.18

125

MNHN, Calypso stn 28

62.6

48.0

34.5

0.77

0 72

1.06

120

MNHN. Guadeloupe

66.3

50.0

39.5

0.75

0.79

1.13

130

MNHN, Salvador, Brazil

61 6

46.2

35.0

0.75

0.76

1,26

120

MNHN, Panama

44,2

35.7

28.8

0.81

1.28

130

MNHN. Antilles

45.0

36.8

27.2

0.82

0.74

1.16

130

MNHN. Les Saintes

64.0

49.3

40.0

0.84

0.77

1.27

125

Total adult shells measured and rib counts

General mean values

0.79

0.76

1.16

123

33.8

Standard devialion

0.03

0.08

4.5

1.9

Largest specîmen in literature, Clench & Smith (1944: 6), 87 x 67 x 46

Acrosterigma magnum (Linné, 1758)

(Figs 1D, H; 2F, G; Table 7)

Cardium magnum Linné, 1758: 680.

Cardium kucostoma Boni. 1780: 46, pl. 3, ligs 6-7.

Cardium marmoreunt Lamarck, 1810: 9 [var [2] exclu-

ded].

CardiumsuMoiigatum Sciwcrby, 184la: 108.

Selected référencés: Cardium magnum Linné —

G nu-lin 1791: 3250 [référence Beta only|.

Wood 1815, 1: 22l.pl. 53, fig. 3.

Trachycardium (Acrosterigma) magnum (Linné) -

Clencii St Smith 1944: 5, pl. 4, ligs i-2.

Nor Cardium magnum Linné - Boni 1780 : 46, pl. 2,

fig.5. [= C. rabumim Solander, 1786] - Chemnirz

1782: I96.pl, 19, fig, 191 [ angidntum Lamarck,

1819J - Bruguière 1~89 ; 229 |= C. Ho ugu t uni

Bruguière, 1789],

Not Cardium kucostoma Boni - Reeve 1845, Sp. 47,

pl. 13, fig. 47 |= Vasticardium luteomarginmuni

(Voskuil & Onverwagt, 1991)].

Not Cardium marmnrcum var. |2] Lamarck, 1819: 9

[= Vasticardntm lutenmarginalum (Voskuil &

Onvervvagt, 1991)1-

TYPl-.s. — Cardium kucostoma Born, 1780 is the

oldest well-idenfified and uliqueslionablc name of the

species in question here. st.iliili/.ed by twn synrypes:

1) The shell ftgured by Lister (1685, pl. 331,

fig. 168), From Jamaica, which is nor rraced;

2) a shell from Queen Maria Theresia’s collection,

figured by Born 1780, pl. 3, Figs 6; 7, now in

NLIMV, Reg. No. 857a (Fig. 2F, G). Born stared

only thar rhis species "lives in America, according

Lister”; figured as lectoiypc of Cardium leucostoma

Born by Voskuil & Onverwagt (1991, pl. 3, fig. 2).

As noted below, this will also be the neotype of

C. magnum Linné.

Cardium magnum: remains an enigmatic name. Tl is

probable that the species so named belongs ro the

grottp of the elongated cockles [qualïfied as “oblong”

by Linné], but I do not share the confidence of

Clench &: Smith (1944: 7) as to iis truc identity,

based only on the given locality “Jamaica”, which

could be crroiicous. Linné left no référencé and no

type spécimen lias yet been traced. I here are only

short descriptions, frcnn which 1 will retain three pas¬

sages;

1) "magnituditie marins' ll764: 489) [. magnitude of

a hand], C. kucnsrmna never rcachcs half of this size; I

know nF only rwo species with oblong shells that

confbrm to this critc-rion: Vaukurdium rlongatum

(Bruguière) and K orbite (Brodetip Sc Sowerhy). Of

the.se, I tliinlc the less improbable-i.s the Former becali¬

se the lutter was probably vc-rv rare in collections in

the mtddleoFthe cightoenlh cennirv:

2) and 3) “sulas augulatis lateiv serratis" (l~58: 680)

h aiigular ribs laterally serrated] and “snlcis retrorsum

crenaüs ’ (1764: 489) [= ribs antenorly ridgcd|. That

description dues not seem appropriate to C. kucosiu-

rna , which bas a rat hcr smooth macroscopie aspect,

hnt it could fit V (longetutti and, even hecter,

V. angulatum [ChemnitzJ F 7 82 interprétation of

C. magnum, figured shell pl. 19, fig. 191, still stored

in 7.MUCJ which tan also be large (heighi niore tlian

100 mm).

In my opinion, Clench N Smith chose the Iess pro¬

bable possibilité in selccting a figure of C kucostoma

Born (Lister 1685, pl. 331, fig. 168) as type figure of

C. magnum Linné. However, in considération of the

existing doubt, tlteir sélection, which stabilrzcs the

name, cannol be disregardcd; therefore I adopr it,

especially since the tendency in (lie American litera-

ture lias been to use it exclusive!)' Neveribcless the

shell figured by Lister is not traced, and a neotype can

be selected; so, 1 here select the lectotype of C. leuco-

stoma as neotype of C. magnum Linné (Fig. 2F, G).

ZOOSYSTEMA • 1999 • 21 (2)

275

Vidal J.

Cardium marmominr. Lunardt dcsignatffd four figure

référencés: Lister 168S (pl. 331, (ig. 168), Boni 1780

(pl. 3, ggs 6-7). Chemnit/. 1782 (pl. 17, fig. 179),

Lamarck 1816 (pl. 297, fig. 3). Üf chese, only Born’s

specimen eau be rraced tnday (see above).

In addition, Lamarck hud three sheils, labelied in hi.s

own hand as marniumm: une in MNHN, with no

locality data; two others in MHNG, from his personal

collection.

Ail the figures and sheils refered ro by Lamarck are

A. magnum , except une in MNHG (Reg. 1085-53)

from Ceylon (Maday coll), described and labelied as

“w [2| testa majore" (which is not a syntype (art.

72b of 1CZN)]. 1 11 is shell is actually Vastïmrdiurn

luteomarginatum (Voskuil & Onvcrwagi 1991). lu

order to avoid confusion. I lierc select the other spéci¬

men of Lamarck's collection in MNHN (Reg.

1085-52) as leciotype of Cardium marmormm

Lamarck.

Cardium subelongatunr. the types of Sowerby’s taxon

are not identified in BMNH, Novorthek-ss, there are

tw'o uncatalogued spécimens from Ctiming collection

from the type locality (St Thomas lsland, West

Indies) which could be constdered as syntypes. J he

largest spécimen (73 x 55 x i4 mm) is probable the

one figured by Reeve ( 1844; fig. 57).

MateriAL EXAMINED.. —The following lots in addi¬

tion to the type nutcrial discussed above:

Cuba. (MNHN de Boury).

Lesser Antilles. (MHNG). — (NHMV), (MNHN

Vidal). — Saint Croix (USNM). — Cap Salomon,

Martinique (MNHN Lamy 1984). — Les Saintes

(MNHN Vidal 1998). — Viens Bourg, Guadeloupe

(MNHN). — Grand Cul-De-Sac, Guadeloupe

(MNHN). - Guadeloupe (MNHN Cabanis

1973). — Guadeloupe (MNHN Douarinou

1973). — Guadeloupe (MNHN Poinrier 1973). —

Marie Galante (MN1 IN Letellier 1949).

Panama. Porto Bcllo Bay (MNHN Vidal). —

Farallon Isknds (MNHN Vidai).

Brazil. Salvador (MNHN Vidal).

Calypso 1961 -62 MNHN: stn 19, 03°50’S, 32°26’W,

Fernando de Norona lsland. — Stn 7, 03°50’S,

32°26W, Atol da Rotas, 47-54 m. — Stn 1,07°29 8,

34°30’W, off Paraiba, 45 m. — Stn 22, ()8'T5’S.

34”42’W, 33 m. — Stn 27, 08°25'S, 34"48 > W.

33 m. — Stn 28, 08°27'S, 34’55’W, 27 m. -

Stn 29, 08°28’S, 34’55’W, 22-30 ni, off

Pernambuco. — Stn 33. 09 D 45'S, 35°35’W. 32 m. off

Algoas. — Stn 45, I l n 22'S. 37“10’W. 31 m, off

Sergipe. — Stn 85, 17' , 50’S, 39"07’W, 2-5 m, Siriba,

Abroîhos Archipclago.

Wrong and Unlcnown localities. Zanzibar

(MNHN). — Singapore (MNHN Denis 1945). —

(MNHN Staadr 1949). — (MNL1N Jousseaume

1921).— (BMNH), (MHNV).

DISTRIBUTION. — South Florida and Bahamas, south

through the West Indies and Gulf of Mexico to Brazil

(Abrolhos Archipelago), i.e. for about 6000 km along

t he eastern coast of the Americas.

Descripi ION

Sheils medium (very rarely large), almost perfect-

Iv ellipsoidal and équilatéral, with exceptionally

Ituncarion ol the posterior part. Ribs show pro-

nouneed hackwards curvature, but posterior part

very rarely expanded. Adult sheils ulways elougat-

ed (mcan L/H = 0.79; range 0.74-0.84); width

also relatively constant (meart L/W = 0.76; range

0.70-0.82). l.unular area very small.

Glossy exterior surface brightly coloured with

orange, whire, yellow, pink and purple. Interior

often eolored only in umbotial area and on ven¬

tral niargiti. Hingc typical of genus, (mean <

A = 123°. range 115°-130°; mean ratio D = 1.16,

range 1.04-1.29). No sterigma.

Mcan rib nnmbcr 33.8 (range 31-40).

Rib morphology conforms to species-group dia-

gnosis (Table 4), but ribs in médian and anrerior

parts are higher than usual and interstices wider;

a gash very frequcntly occurs along posterior 8th

or 9[h rib ol MPQ in right valve. Interstices

regulurlv notebed by transverse, elongated, some-

rimes erescent-shaped notchcs.

Rf.marks

Acrosterigma magnum lias often been confused

with Vasticardium luteomarginatum (Voskuil &

Onverwagt, 1991), and bas also been described

as “cxçeedingly close” (Clench & Smitlt 1944; 7)

to Vasticardium ckmgaium (Bruguière, 1789).

While it is truc that somc smooth forms of the

latter two câll hâve a superficial tesemblance,

tbey differ, without an) ambiguirv, by ail the

charàcter.s separating the two généra (see

Table 2). In addition, A magnum is distingutsh-

etl by its bright colours and glossy surface, and

above ail by the typical notching of interstices

and lowcr part of flanks of ribs, while A. hiteo-

margimyum h as finely strîuted interstices and

elongatum smooth interstices.

Acrosterigma burchardi (Dunker, 1877)

(Fig. 2H, I; Table 8)

Cardium burchardti [sic\ Dunker, 1877: 67 [dedicated

to G. Burchard].

276

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 8. — Measurements (in mm) and rib count Acrosterigma burchardi (Dunker, 1877).

L/H

W/L

A°

Ribs

Holotype burchardi

71.0

58 2

37.6

0.82

0.65

105

USNM 304262, Awaji

76.7

60.2

41.0

0.78

0.68

0.92

110

USNM 605943, Tosa Bay

62 8

49.4

34.1

0.79

0.69

0.73

105

MNHN ( Nikawa

84.0

65.0

45.0

0.77

0.69

=1.0

105

Idem

79.0

61.3

42.7

0.78

0.70

105

MNHN, Kyushu

62,5

52.7

34.2

0.84

0.65

0.79

105

MNHN, Kyushu

60.8

50.5

33.6

0.83

0.67

0.87

110

LACM 23134, Kyushu

75.2

63.5

40.5

0.84

0.64

BPBM 11060, Matsuka Ki

84.2

71.0

45.3

0.84

0.64

Total measured adult shells and rib counts

General mean values

0.81

0.67

0.86

106

42.2

Standard déviation

0.03

0.02

0.09

2.3

1.7

Largest specimen observed, BPBM 11060 (see above).

Cardium burchardi Dunker, 1882: 210, pl. 15,

figs 4-6.

HOLOTYPE. — The shell described and figured by

Dunker, 71 mm high, witb 40-42 ribs; not rraced.

Ma i priai iXAVlINFD. — The following lois:

Japan. (I ACM 52862), — (BPBM 11060). —

Nikawa, Aicbi pref. (MNHN Vidal). — Kyushu

(MNHN Petit 1972); (MNHN Vidal); (LACM

48715); 48 m (LACM 50911, 23134, 13434). —

Boshi (NMW MeJvill). — Kii (LACM 13444); 18 m

(LACM 13452). — Marsuka Ki (BPBM 10233). —

Saganii Bav (BPBM 204080, 204081). — Pükura,

Awaji (USNM 304262), — T osa Ray (US NM

605943).

DISTRIBUTION, — The living species occurs exclusive-

ly in Japan, Honshu (Boso Peninsula, east of Tokyo,

as northern limit), Shikoku and Kyushu, Japan.

Description

Shells medium to large, very asymmetrical; antc-

rior margin raised and rounded; posterior margin

receding and almost srraiglit due to llattcning of

PQ, whlçh forms a rounded angle witli rest of

shell. MPQ margin may also be somewhnt

straightened. Shell moderately elongated (L/H

range: 0.77-0.86) and apprcciably depressed

(W/L range: 0.64-0.70).

Lunule narrotv, more devclopcd on right valve, a

little hollowed and bounded by a widc, high,

longitudinal fold, wbich bcars fine oblique

ridges, corresponding to the AQ first rib.

External colour yellowish, sometimes with pink-

ish shades; PQ often purplish. Interior vvhite.

Hinge line asymmetrical (ratio D range

0.73-1.00), and strongly angled (< A range

105°-110"). Umbona! eavity filled with a brown

callo.sity that often terminâtes ventrally in a short

sterigma.

Mean rib numbet 42.2 (range 40-45).

Rib morphology conforms to spepies-group dia-

gnosis (Table 4); on PQ, posterior thin tubercles

of ribs can be joined rogether by a iliîn ridge; on

otber parts of shell ribs flat-topped, slightly over-

banging interstices, teiro-crenulated on MPQ,

progressively bi-crcnulated, and tben very sightly

top-ridged. On most of the adult zone of large

specimens, ribs becotne rounded and interstices

wider on MPQ.

Species-group of Acrosterigma cygriorum

(Deshayes, 1855)

INCLUDED species. — 1) Recent: A. cygriorum

(Deshayes, 1 855); -A. sarenseni (Powell, 1958);

A. maridae Wilson tk Stevenson, 1977; A. kenlakae

Healy & Lamprell, 1992; A. abrolhenm n, sp,

2) Fossil: According lu the literauire data, the tollo-

wingspecies very prob.lhly helong to this gnemp:

Vusticardium [Resazara) praecygnontm Ludbrook,

1955, from Dry Creek Sands (Pliocène), Adelaide,

South Auscralia This shell, with 48 ribs, vvas compared

by Ludbrook (1955: 61) with Acrosterigma kcrslakae

Healy & Lamprell, 1992 (see below).

Trachycardium ( Regozara ) delectabile Maxwell, 1978,

ZOOSYSTEMA • 1999 • 21 (2)

277

Fig. 3. — A, B, Acrosterigma cygnorum, specimen from Adelaide, South Australia: C, D, E, Acrosterigma sorenseni, holotype;

E, view of PQ and MPQ; F, G, Acrosterigma kerslakae, paratype AMS 80144, from Collaroy Beach. Sydney; H, Acrosterigma marie-

lae, a left valve from Cheyne Bay. Western Australia, AMS 310554; I. Acrosterigma marielae, same specimen, detail of the médian

zone; J, Acrosterigma abrolhensis, holotype; K, Acrosterigma abrolhensis, holotype; detail of the médian zone. Scale bars: A-E, H,

J, 10 mm; F, G, 20 mm; I, K, 2 mm.

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Fig. 4. — Distribution ot the species of the species-group of Acrosterigma cygnorum (A. sorenseni excluded: see Fig. 11).

from Clifdenian (Middle Miocene). Waiau River,

South Island, New Zealand. This shell, with 57 ribs,

was contpared by Maxwell (1978: 22) with

Acrosterigma sorenseni (Powell, 1958) and A. cygnorum

(Deshayes, 1855) (seebelow).

Distribution. — (l igs 4; il) AU the spedes, living

and fossil, arc rcstricted to Southern part of Australia

and/or ro New Zealand,

DlAGNOSIS. — See Table 4.

Acrosterigma cygnorum (Deshayes, 1855)

(Figs 2K; 3A, B; Table 9)

Cardium cygnorum Deshayes, 1855: 331.

? Cardiutn joveolaturn Sowerby, 1841a: 111.

Table 9. — Measurements (in mm) and rib count of Acrosterigma cygnorum (Deshayes, 1855).

L/H

W/L

A°

Ribs

Syntype cygnorum No. 1

52.5

45.0

30.0

0 86

0.67

Idem No. 2

44.2

40.5

25.6

0.91

0.63

Idem No. 3

32.7

30.5

21.4

0.93

0.71

USNM 253444

50.5

47.0

28.8

0.93

0.61

1.10

125

USNM 160215

42.6

36.6

26.6

0.86

0.73

1.12

125

MNHN Western Australia

55.4

50.2

29.1

0.91

0.58

=1.0

125

Idem

46.0

42.3

26.4

0.92

0.62

1.05

125

MNHN, Aaelaide

51.7

47.5

29.5

0.92

0.62

0.95

125

MNHN, Brlghton

51.1

46.5

29 4

0.91

0.63

=1.0

125

LACM 23132

55.7

46.8

32.2

0.84

0.69

Total adult shells measured and rib counts

General mean values

0.90

0.65

1.03

125

42.6

Standard déviation

0.03

0.04

0.05

2.2

Largest specimen observed, LACM 23132 (see above)

ZOOSYSTEMA • 1999 • 21 (2)

279

Vidal J.

Laeviatrdium ( I rticbycilrdium) gaillardi Fischer-Pierre,

1977: 58, pl. 6, figs 1-2.

Types. — Car rhum cygnomnn three syntypcs in

BMNH, rcg. 1971-23, Cuming collection, from

Swan River, Western Australie. Cardium fovenltUum:

from Swan River also, not traced; Wilson &

Stevenson {19"-: 91) werc eonvinced of the synony¬

me with C. cygntmtm and suspcctcd tlt.it a type spcci-

men could Lie One of rhe syntypcs of C. tygnorum.

Laevicardium (Tntchyairdiitmi gnUbirdr. holotype in

MNHN, collected by Quoy & Gaymard during the

voyage of /, 'Astrolabe (1826-1829), labelled as coming

from New-Zealand. However, C cygnorum has never

been recorded Irom tins countr)', and die spécimen

probably uriginates Iront Australia wItéré the Astrolabe

called at several tintes during the sanie voyage.

MaTeriAI eSaMINI.d. — The following lots in addi¬

tion to the type tmtcrîal:

Australia. il At ..VI 13356), (MNHN Vidal).

Western Australia. Géographe Bay (Hobbs). —

Freemantle (ANSP 263242).

South Australia. (I 5NM 253444, 160215). —

(ANSP 72345). Adelaide (MNHN Vidal). —

Revesly Island, Banks grotip (MNHN Vidal,

2 lots). — Rrigliton, Adelaide (MNHN Vidal). —

Norntanville (L.ACM23 I 32). — Hardwicke B a v

(LACM 13342). — Hirdwickc Bay (USNM

321673). — Fort Granville il AC VI 28314). - St

Vincent Gulf (LACM 13369, 508 Vi). — Wallaroo

(LACM 50890, 50858). Sémaphore Bav (ANSP

186763).

Victoria. (USNM 203909). — Port Bav (LACM

28212). — Melbourne (ANSP 98954).

Tasmania. (MNHN Stanley).

Distribution. — (Fig- 4) According to Wils*on &c

Stevenson (1977: 92) ''Southern Australia, the most

northern records being Fremantle in the west coast,

and Momagu Island on the east coast’’. It is also “fai r-

ly common along the north coast” of Tasmania (May

1958: 13).

Description

Shell medium-sized, generally symmerrical with

anterior margin rounded and raised, and poste-

rior slighdy receding, with a small oblique tron¬

cation. PQ a iiftle flartened, fbrming a rounded

obtuse angle with rest of shell. Not elongated

(mean L/H = 0.90; range 0.84-0.95) and mode-

rately compresser! (mean W/F = 0.65; range

0.58-0.73).

Lunule narrow, identical on both valves, and

slightly hollowed, with posterior margin well-deli-

mited. External colour cream to yellowish, with

tnore or less developed brownish splashes; interior

white, Hinge approximatelv symmetric-al, and

moderately angled (< A about 125°). A low

Sterigma occurs in sonie shell.s about 40% of lots.

Mean rib nnmber 42.6 (range 36-47).

Rib morphology: on PQ (Fig, 2K), posterior rib

scales sometimes prolonged aboyé the anterior

sntooth part, fnrming herringboties. On MPQ,

ribs radier high, variably rounded to subuiangu-

lar in section, sometimes with a rhin top-crest;

always retro-ridged on adult part, becoming

bi-tuberculatetl. Interstices thin on juvénile shell,

one third to haJf the width of ribs in adult. On

adult part, anterior lialf of ribs becomes progres-

sively more square-sided, bi-tuberculatetl or

bi-ridged, then top-ridged. Ridges always thin

and tree (not imbricated throughout), sometimes

herringboned. Interstices similar to chose on

MPQ, smooth and slightly overhung by ribs.

Acrosterigma sorenseni (Powcll, 1958)

(Fig. 3C-L; Table 10)

Trachycardium ( Vasticardium ) sorenseni Povvell, 1958:

76, pf. 11, figs 6; 7.

Types. — Holotype: a rigbt valve in Auckland

Muséum AK 701236 Iront 14 en liant Ray, Raoul

Island. Kermadec Islands, 29H6 S, 178"03’E, 0 ni.

Two other smaller valves in Aucldand Muséum from

Cuilatheu 1952, stn 674 off Raoul island, 29*T5’S,

177°67'L, 75-85ni, were clted but not qualifîed para-

types by rhe aurhor.

MATERIAI EXAMINED. —- The holoty r pe and the fol¬

lowing lors:

New Zealand. Raoul Island, Kermadec Islands, in

MNZ (M.202885 Bollons). — (M.213883 Oliver

1908). — (M.202889).

AdkrOit 1975: stn 75443, off Meyer Islet, 29°14,7’S,

177-52,7'W, 22-27 m (M.225794). — Stn 75436,

S F of d’Arcv Point, 29'M8.5'S, 177“54.5'W, 44 m

(M.225784).'

A (héron 1976: stn 765”3, between Davrel and

Chanter lslets, 29°I5.00'S, 17^50.90’W 31-45 m

(M.226972). — Srn 76567, Lasr Anchorage,

29 o 16.00’S, I77 I ’51.58’W, 42-47 m (M.226611).

DISTRIBUTION. — (Fig. Il) Raoul Island, Kermadec

Islands, New Zealand.

Description

Shell of medium size, almost symmetrical and

280

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of rhe elongated cockles

Table 10. — Measurements (in mm) and rib count of Acrosîerigma sorenseni (Powell, 1958).

L/H

W/L

A°

Ribs

Holotype sorenseni

42.3

40.0

(24.0)

0.95

0.60

120

Galathea 674

26.5

25.0

(15.0)

0.94

0.60

Galathea 674

20.5

20.0

(10 0)

0.98

0.50

MNZ M.202885

44.5

39.7

(23.0)

0.89

0.58

=1.0

MNZ M 213883

40.6

34.2

(21.6)

0.84

0.63

=1.0

MNZ M.225794

29.1

28.0

15.5

0.96

0.55

0.82

115

MNZ M.202889

29.4

27.5

(15.2)

0.94

0.55

120

MNZ idem

19.3

20.0

(10.7)

1.04

0.53

MNZ M.225784

28.0

27.0

(15.0)

0.96

0.56

130

Total adult shells measured and rib counts

General mean values

0.94

0.56

0.91

121

51.2

Standard déviation

0.05

0.03

0.08

5.4

1.4

Largest specimens observed, the holotype and NMZ.M202885 (see above)

rounded in ourline (mean L/H = 0.94, range

0.84-0.97, but only two fully adult shells hâve

this ratio less than 0.93, see measurements table),

rather tompressed (mean extrapolated

W/L = 0.56, range 0.53-0.63). PQ area very

slightly flattened.

Lunule non-existem, ribs practically reaching

margins. Colour ol 1res h shells whitish heavily

blotched with orange-pink; interior with external

colour pattern showing rhrough: holotype, a

faded beach shell, shows a sparsely speckled pat¬

tern on a pale ground. Hinge variably asymme¬

trical and rather moderately angled (< A range

1 15-130). No sterigma.

Mean rib number 51.2, range 48-54.

Rib morphology: PQ (Fig. 3E) conlorms to spe-

cies-group, with irregu la r, somewhat elongated

oblique tubercles on posterior liait of ribs. On

MPQ (Fig. 3F,), ribs retro-ridged, almosr

square-sided, but with rounded tops, and a slight

forward bending; interstices about one third of

ribs widrh, and loosely striared. On ancerior parr

of shell, ribs become higher, square-sided,

flat-topped and top-ridged with wide touching

ridges wdiich are sometinies imbricatcd.

Interstices become progressivcly narrower.

Remarks

Acrosterigma sorenseni is close to A. cy griot uni.

The former mainly differs in: (1) higher rib

number (48-54 against 35-47 in cygnorum); (2)

presence on PQ of a discontinuous axial furrow,

formed bv successive holes between the scales;

(3) different rib morphology on AQ, i.e. rhinner

interstices, higher ribs which are square-sided

rather thau rounded and, on the ribs, wider,

more imbricâted litiges which almost toüch (ins-

tead of the thin, free ridges of A. cygnorum).

The rwo valves in New Zealand Océanographie

lnstiture cited bv Powell (1958: 76) from E of

Philip lsland, Norfolk Island “similar to the

Kermadec species in sculpture but of oblique-

ovate outline [...) and with a rib count 48-50”

are very probable A. maculositm howense (see in

this subspeties, abondant in Norfolk Island).

Acrosterigma kei'slakae

Healy & Lamprell, 1992

(Fig. 3 F, G; Table 11 )

Acrosterigma kerslakae Healy & Lamprell, 1992: 84,

pl. 3, figs a-d.

Cardium oxygonum Sowerby, 1833 - Hedlev, 1923:

304.

Tvi'CS. Holotype: a paired specimen

AMS C31559, from Burpengary, Queensland,

27° I O’S, 152°57T,. Paratypes: one lot in AMS from

Collaroy Beach, Sydney (AMS C80164), and two lots

in QM from Caloundra (MO33056) and Southport

(MO32904).

MATERIAU EXAMINEE), — rhe following lots in addi¬

tion to thethree para type lots:

Queensland. Snellv Beach. Caloundra (MNHN

Vidal). — Caloundra (WAM 4796-68). — ZOMO’S,

1 53°07’E, Alexandra Head (AMS C315813). —

ZOOSYSTEMA • 1999 • 21 (2)

281

Vidal J.

Table 11. — Measurements (in mm) and rib count of Acrosterigma kerslakae Healy & Lamprell, 1992.

L/H

W/L

A°

Ribs

Holotype kerslakae

28.0

27.9

16.5

#1.0

0.59

120

AMS C80164, paratype

39.7

35.2

(25.6)

0.90

0.73

125

Idem, paratype

36.4

34.0

(23.0)

0 93

0.68

1.05

AMS Cl45036

34.4

30.7

(22.8)

0.89

0.74

125

AMS Cl05746

38.0

35.5

(26.0)

0.93

0.73

*1.0

AMS C62419

35.0

31 0

(22.0)

0 89

0.71

125

AMS Cl 45034

32.3

28.6

(18.2)

0.89

0.64

125

AMS Cl45088

34 2

31 0

(22.0)

0 91

0.71

120

AMS Cl 45037

37.2

34.6

(24.0)

0.93

0.69

125

AMS Cl4442

40.2

34.0

(27.2)

0.85

0.80

*1.0

Total adult shells measured and rib counts

General mean values

0.91

0.69

1.02

124

40.9

Standard déviation

0.03

0.05

0.02

2.1

2.4

Largest specimen observed, AMS Cl4442 (see above)

26°49’S, 153°10'E, Caloundra (AMS C315819,

C315820. C315821. C054640, C047785.

CO 12772). — 27“ 1 5’S, 153"15’E, Moxeton Bay

3.5-11 m (AMS C315829). — 27°26’S, 153°32’E,

Point Lookout, Stradbrokc Island, Moreton Bay

(AMS Cl45026, C315832). — 2 7 °31'S, 153°4()'li.

off Morc'ton Bay. 75-80 m (AMS 315826). —

28°05'S, 1 5 3 " 2 7 ' F, Grtat Burleigh Hcad

(AMS Cl45027). — 28 U 06'S. 15.V28T., Tallebud-

gera Crcek, Big Burleigh (AMS C145028). —

28°10’S, 153“32’, Coolangatta (AMS C315827,

C315824).

New South Wales. 28°38'S, I53*37’.S, Byron Bay

(AMS C0052I5). — 28“52'S, 153“34’E, Ballina

(AMS C005088). - 29”29’S. 153“22’P, Angourie

Point (AMS C315935. C3I5853). — 29“32'S.

153“22'E, Shclly Bcaclt. 8 km soulii ufYamba

(AMS C315851). — 29°46'S. 153 l, 18'E. Minnic

Waters (AMS C315959). — 30°0Û’S, 1S3 Ù 12’E.

Woolgoolga (AMS Cl45029). - 30°12’S, 153° 16 P.

South Soluary Lsland, oll Colfs Harbour

(AMS C108804). — 30°53 , S, 153°04’E, Trial Bay

(AMS C028435). — 32°04’S. 152“33’P, Point

Halliday, ueat Porstei (AMS C0928141 — 32°42’S,

152°05'b', Port Stephens (AMS C084363.

Cl45031). — 32"45"S. 152“l TE. Fingal Bay, Port

Stephens (AMS C315960). 33 n 04‘S, 151“36'E,

Pake Maquarie (AMS C01444-2). — 33°05’S,

151“39'E, Blacksmïths Beach, Swansea (AMS

C315961). — 33*3 l’S. I51°I9’F.. Océan Beach.

Brokcn Bay (AMS Cl 4 5030). 33 Ù 35'S. 151°19'E,

Palm Beach, Sydney (AMS C315831). - 33°36'S.

151“17'E, limer Basin, Pittwater, Sydney

(AMS Cl45032). — 33“42*S. 151°I8’E, Narrabcen

Beach, Sydney (AMS Cl 45033). — 33*44'S.

151°18‘E, Collatoy Beach, N of Sydney

(AMS C315937, C0624I9). - 33*45 S, 151“19’F',

Long Reef, Collaroy, Sydney (AMS C145034,

Cl45035). — 33°48’S, 151°16 K, North llarbour,

Port Jackson, Sydney (AMS Cl45036). — 33°48’S,

151 ,J i7 E, Manly Beach, Sydney (AMS C088850,

C105746, C315962). — 33°48'-33°50’S,

1 5 1 * 1 4’-1 51 “1 6’E, Middle Harbour. Sydney

(AMS C017776). — 33°49 S. r5l“l5'E, Baimoral

Beach, Sydney (AMS C00I882). — 33°50’S,

151 " 16’E, off.Sow and Pigs Reef, Port Jackson,

Sydney (AMS 0)55349). - 33°50’-33“52’S,

15 1° 1 2’-] 5 I * Jô’E, Port Jackson, Sydney

(AMS C145037). — 33*51% 15t°i4’E, Port

Jackson, Sydney (AMS Cl 45038). - 33*50’-

33°52’S, I 5 | * I 2’- I 51 *16’E, Sydney Harbour

(AVIS Cl45092). - 3.V’58.76’S, 15I’’13 69E, Yârra

Bay. Borany Bav (AMS C3 15938) — 33“59’S,

15‘|°12'E. Borany Bay (AMS C3I3963). —

34"0.58’S, 151" I2’38E. Kurnell. Borany Bay

(AMS Cl45040). - 34*03'.$,. 151*09*411. Cfonulla

Beach (AMS C 145034). — 34°32'S. I50°52’E,

Windang (AMS C066189). — 34*35’$. I50°52’E,

Shell Harbour (AMS C315965, 008306). —

34°49”S, 150*46’H, Sevcn Miles Beach, near

Gerringong (AMS Cl45089). — 35’24’S, 150*27’E,

Rtitril 1 ake, S of U i J a «J u 1 la (AMS C315966,

145090). — 36*13’*$, 150°08’4E, Narooma

(AMS 045091).

DISTRIBUTION. — (Fig. 4) Eastern coast of Australia

along more rhan 1000 icm of coasrEme, (rom

Caloundra, Queensland (26°49’S) south to Naruma,

New South Wales (36"13’S).

DESCRIPTION

Shell medium-sized, almost symmetrical with a

very small truncation on PQ, only slightly elon-

gated (mean L/H = 0.91; range Ô.88-1.00) and

282

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

rather compressée! (mean W/L = 0.69; range

0.59-0.80).

Lunule small, a lirtle hollowed, almost équivalent

in both valves, well-delimited posteriorly, and

always colored pink-purple. Exterior colour

cream-yellow, wirh prominent pink-brown

blotches, whicb are fused near umbones; interior

white, cxcept For occasional pale V-shaped

umbonal rays. Hinge symmctncal (ratio D about

1.0) and moderately angled (< A range

120-125°). A low sterigma présent on some

shells in 13% ol lots.

Mean rib number 40.9 (range 35-45).

Rib morphology: PQ excepted, ribs variable in

profile, generally low and slightly rounded, but

sometinies almost triangular or trapézoïdal,

flat-topped and overhanging interstices. Rib

ornaments also variable in number and size,

generally retro-ridged on posterior MPQ, smootli

to rétro- or bi-tubcreulated on central part, and

top-ridged with thin ridges on AQ, sometimes

herringboned. Interstices always narrow.

Remarks

Acrosterigrna kersldkde is very close to A. cygno¬

rum-, the former is generally smallcr, with lovver,

smoother ribs and narrower interstices, and has a

lower mean rib number (40.2 compared to 42.7

in cygnorum). I lowever, these two fbrms are per-

fectly parapatric, and the weakness of rheir diffé¬

rences cannot exclude the possibilitv of the

existence ol two subspecies. Nevcrtheless, rhey

hâve been treated as distinct species by the majo-

rity of aurhors, for example Hedley (see above),

and Ludbrook (1955; 61) who stated: “Iredale

(1936: 276) bas pointed out that NSW shells

reterred to Cantium cygnorum (typically from

western Australia) are not referablc to cygnorum

[...] and the writer is inclined to agréé with this

opinion”.

Acrosterigrna marielae

Wilson & Stevenson, 1977

(Fig. 3H, I; Table 12)

Acrosterigmn mariel/lc Wilson & Stevenson, 1977:

103, pl. 6, figs 11-15

Types. — Holotype: a specimen from CSIRO 1963,

stn 216, 31°18'S, 115“03'E, W of Cape Leschenault,

Western Australia, 91 m (WAM 215-67). Paratypes:

28 (no details in Wilson & Stevenson).

Mai l KIAI EXAMINER, —The following lots:

Western Australia. Giiscoync 1962: 33°43 S,

125”04’E, East of Rocky Point, Grcat Australian

Bight, "'7-80 m (AMS t 31 055 1). — 34°21'S,

121"I6’E. E of I lood Point. Great Australian Bight,

82 rn (AMS C310553). — 34“25’S, 12H20T., saine,

158 ni tAMS C310552). — 34°55'S. 1 Proo'R. E of

Chcvne Bay, 71-76 ni (AMS C310554).

Diamantina 1972: 28°18'S,113"58’E, W of North

Island. Hautnian Abtolhos, 108 m (WAM

233-90). — 28”43‘S, 113"5l'E, (5 un mi Island,

Eastern Group. Abtolhos, 43 ni (WAM T 85-91). —

28‘'45 S, 113®50'E, Littlc N'orth Island, LastCrn

Group, Ahrulhos, 45 m (WAM 784-91), — ,30°34'S,

1 |4°11*F, NW of Green Island. 128 m (WAM

27-94). 30"34'S, 114"56’, ' 15“06'E, W of Guil-

derron, 146 m (WAM 29-94, 30-94). — 30 U 35’S,

114"35’E, SW of Cervantes, 110 m (WAM

37-94). - 31°00'S, I I4°52'E, W of Lancelin, 150 m

(WAM 37-94). — 32°02’S, ! I5°22’i:, W ofRorrnest

Island, 110 m (WAM 2.32-90).

Sprightly 1976: 29”067S, 113°58’5E, r. 32 km W of

Dongara, 91 m (AMS C310604).

Mo res b y 1980: 31°40'6S, 115°09 6E, 100 m

(AMS C310603).

Distkibi tlON. • (Fig. 4) According ro Wilson &

Stevenson (1977: 104), “between 33 and 152 fathoms

[60,3-274.3 m| off the mid-west coast of Western

Australia”,

Description

Shell small, rliin, slightly obliquely ovare, inequi-

later.il wirh anteriur dorsal margirt rounded and

posterior margin larher srraight and not tnjnea-

ted on PQ. Not very elongated (according to

Wilson & Stevenson Î977, mean L/H - 0.91;

range 0.84-0.96) and moderately compressée!

(mean W/l. = 0.62; range 0.57-0.70).

Lunule well-defined, wider and projecling Irom

the margin in riglu valve, and somewhat hollow¬

ed. Exterior glossy, cream in colour, with small

splashcs of orange; interior white cxcept for pink

V-shaped umbonal ravs, lunule and hinge below

it orange-pink, mainly in right valve. Hinge

ncarly symmetrical (ratio D close to 1.0) and

moderately angled (< A range 120°-130°).

Mean rib number 62.6, range 56-68.

Rib morphology: on PQ, scales on posterior parts

of ribs variable; prismatic, roundly cubercular, or

ZOOSYSTEMA • 1999 • 21 (2)

283

Vidal J.

Table 12. — Measurements (in mm) and rib court of Acrosterigma marietae Wilson 8 Stevenson, 1977.

L/H

W/L

A 0

Ribs

Holotype marielae

22.6

20.0

12.8

0.88

0,64

1.25

120

WAM 232-90

22.8

20.8

(12.8)

0.91

0.62

1.21

Idem 20.7

18.6

(11.8)

0.90

0.63

125

WAM 27-94

19.2

17.1

(11.4)

0.89

0,67

130

AMS C310553

188

17.0

(10.0)

0.90

0.59

130

AMS C310554

18.4

16 8

(10 0)

0.91

0.60

1.10

WAM 30-94

23.1

20.0

(13.6)

0.87

0.68

M.O

Total adult shells measured and rib counts

General mean values

0.90

0.62

1.13

128

62.6

Standard déviation

0.03

0.04

0.08

3.6

3.0

Largest specimen observed, WAM 30-94 (see above)

sometimes elongated. On rest of shell (Fig. 31),

ribs gencrally rounded to roundly triangular,

retro-ridged to retro-Utberculated; ridges or

tubercles variable in sbape, prismatic and point-

ed to elongated and acute, placed on npper part

of sides of rib.s. Ribs become progressive!)' more

top-ridged on AQ, but on adult part of some

specimens, bi-tubcrculated ribs appear as early as

on the anterior part of MPQ and develop fur-

ther, changing into herringbone sculpture on

AQ.

Remarks

A. marielae is close to A. cygnorum and A. kersla-

kae as far as shape and colour are concerned, but

differs by its smaller size, much larger rib num

ber, and rib ornamentation (stronger and more

acute latéral ridges or lubercles),

Acrosterigma abrolhetue n. sp.

(Fig. 3J, K; Table 13)

Types. — llolotype: a paired specimen, ilitulm

1977, sut 8, 28°37.5'S, 113°51.5’E, off l.ittlc North

Island, Easter Group. Abrollios Islands, Western

Australia, 42 m (WAM 775-91). l’ararvpe 1: a left

valve, Grucoyne l%2, 34 D 55’S, 119°00’E, east of

Cheyne Bay, Western Australia, T 1 1-76 m

(AMS C310554). Paratvpe 2: a right valve, Rumphius

1 1973, stn Li 2, 03“13’S, 128M4 E, east of Piru Bay,

Ceram, lndonesia (WAM 43-94).

EtYMOLOGY. — I rom tire Abrolhos Islands, Western

Australia.

MATERIAL EXAMINED. — Additionally to the type

material, a righr valve. Western Australia. Sprightly

1976. 29" 10.<)'S, I 14”43'H, 24 km NW of Dongara,

40 m (WAM 777-91).

Dis I RIHUTION. — (Fig. 4) Western Australia, in the

vieillit) ol the Abrolhos Islands and in Cheyne Bay

ncar Albany. 1 rhink irs présence in Ceram, represent-

ed by a single valve (paratvpe 2), needs confirmation.

Description

A small shell, obliquely ovate, and slightly

inequilateral. vvith posterior margin slightly trun-

cated on PQ vvhich forms a vague obtuse angle

wirh MPQ; not elongated (L/H close to 1.0) and

somewhat compressed.

Lunule small, equally dcveloped in both valves,

not sliifted front margin, and slightly hollowed.

Colour wbite, both externally and internally,

except for lunule, slightly pinkish. Hinge sym-

metrical and moderately anglcd (c. 130°).

Rib number43-46.

Rib morphology: on PQ. rcgular elongated

oblique scales occur on posterior part ot ribs; on

MPQ, ribs low, slightly rounded, and retro-

ridged or retro-tubcrculnced on rheir lower sides,

ncar interstices; in MAQ (Fig. 3K) rib.s become

ptogrcssivcly flat-topped, slightly squared, and

bi-tuberculated, side lubercles being roughly

pyramidal to slightly elongated and .tctite and

almost touching across interstice. More anterior-

ly, side tubercles leitgtben and become slutrp

latéral oblique ridges wliich rend to join on top,

forming herringbone sculpture; on AQ, ribs

become at once laterally tuberculated and

284

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 13. — Measurements (in mm) and rib count of Acrosterigma abrolhense n. sp.

L/H

W/L

A 0

Ribs

Holotype abrolhense

13.2

12.3

7.3

0.93

0.54

*1.0

130

Paratype No. 1

12.6

13.0

(7.0)

1.03

0.54

=1.0

Paratype No. 2

10.0

(5.6)

1.00

0.56

130

WAM 777-91

10.0

10.3

(6.0)

1.03

0.58

top-ridged with thin free ridges. Interstices

always very narrow. Tops of ribs covered by a

multitude of very thin holes, sometimes concen-

trically aligned on juvénile part, probably corres-

ponding to lost pustules.

Remarks

The rib ornamentation cil A. abrolhense is com¬

parable to that of A. nuirielae, but the rib mor-

phology is different (compare I'ig, 31 and K), as

is the lunule, the number ol ribs and the colour.

A. abrolhense d i fiers from A. cyguoruni and

A. kerslakae by ils smaller si/e, colours and rib

ornamentation. I bis species is further charaçreriz-

ed by the very numerous pustular holes on the

ribs, présent but never as abundant in several

other Acrosterigma species.

Species-group o( Acrosterigma variegatum

(Sowerby, 1840)

INCLUDED SPECIES. — Recent: A. variegatum

(Sowerby, 1840); A. oxygonum (Sowerby, 1834);

A. selene n. sp.; A. disais n. sp.; A. mauritianum

(Dcshayes, 1855).

Description

See Table 4.

Remarks

This species-group is the closest to the genus

Vasticardium.

Acrosterigma variegatum (Sowerby, 1841)

(Fig. 5A-C; Table 14)

Cardium vanegatütn Sowerby, 1841a: 107.

TYPES. — lhree shells considered as syntypes in

BMNH, not catalogued, Cuming collection, labelled

from Ticao (Philippines) and another location (label

illcgihle). The dimensions given by Sowerby lûr bis

figltred spécimen cotlkl fit thé smallcst spécimen, but

the colour pattern, the rib number (48 given), and the

locality (Leyti given) do not match. The specimen

figured bere (Fig. 5A-C), better lits the figure, des¬

cription and locàlitv given by Reevc ( 1845: Sp 75).

Mater lai. examined. — The following lots in addi¬

tion to the syntypes:

Philippines. (MNHN). — (MNHM Vidal). —

Magellan Bay (MNHN Vidal). — Burîas Islattd

(USNM 237011). — Zamboanga. Mindanao

(LACM 5D834, USNM 248371), — Mindanao

(USNM 23721 T ). — Sa Cruz Island. Mindanao

(USNM 248371), — Davao, Mindanao (USNM

248301). — Prie 1064: Doe Can Island, Sulu

Arihipel.tgo, 1-3 m (WAM 656-66). — Jolo Island,

Sulu Archipelago (USNM 235598, LACM

90046). — Siasi Island, Sulu Arcbipelago (USNM

612432, TR’ RM 203569. MNHN Vidal). — Sulu

Arcbipelago (.VINHN Vidal).

Indoncsia. Moluccas (ZVIA de Serriere). — Flores

(ZMA Winckelswcep).

Papua New Guinea. Weleluku, Kiveco

(AMS C.3164). — S.i ma rai (QM MO20733).

Queensland. Murray Island, Torres Strait

(AMS C30279). Port Douglas (MNHN Vidal).

Wallis and Futuna. MUSORSTOM 7 1992

(MNHN): sut HW 529, 12°31’S. IHi'HO’W,

Waivrwitdi Batik, 500 ni. — Stn DW 538, 12‘'3I S,

176'-’40'W, W.uerwitcl) Bank, 275-295 m. — Stn

DW 588 12”17'S, 174'45’W, Field Bank,

490-500 m.

Distribution. — (Fig. 6) Tropical western Pacific as

far east as Wallis and Futuna; very abundant in the

Philippines.

Description

Shell medium-sized, ovoid, almost équilatéral

(anterioi dorsal margin slightly angled,), not post¬

eriori}' truncared. nor elongated (mean

L/H - 0.91: range 0.87-0.95) and moderately

compressed (mcan W/L = 0.67; range

0.58-0.78).

ZOOSYSTEMA • 1999 • 21 (2)

285

Fig. 5. — A. B, Acrosterigma variegatum, syntype: C. Acrosterigma variegatum. specimen from Magellan Bay. Philippines, MNHN:

detail of the particular lunule; D, E, Acrosterigma oxygonum, syntype; F. Acrosterigma oxygonum. specimen from Rameswaran,

India, MNFIN; detail of PQ and MPO; G, H, Acrosterigma selene, holotype: I. Acrosterigma setene , holotype; view of the deep lunule;

J, Acrosterigma selene, paratype 1 ; K, L, Acrosterigma discus, holotype; M, Acrosterigma discus, holotype; detail of PQ and MPQ.

Scale bars: 10 mm.

ZOOSYSTEMA - 1999

Taxonomie review of the elongated cockles

Fig. 6. — Distribution of the species of the species-group of Acrosterigma variegatum.

Lunule wide on right valve, projecting from the

margin against left valve, and composed of two

parts: a strongly hollowed part near margin,

fornting an elongated Socket, and an elevated

part behind thar, forming a large rib-Iike ridge,

both parts being concentricallv finely ridged

(Fig. 5C); practically no lunule on left valve

(Fig. 5C). Exterior beige variegated with large

irregular patches ot orange, brown, and purple,

which are visible interiorly. Hingc markedly

asymmetrical (average ratio D = 0.84; range

0.78-0.88) and moderately angled (average

< A = 128°; range 125 a -130°).

Mean rib number 40.2, range 36-43 (never as

nvany as 48, as indicated by Sowerby).

Rib morphology; on PQ, anterior part of ribs

Table 14. — Measurements (in mm) and rib count of Acrosterigma variegatum (Sowerby, 1840).

W/L

A°

Ribs

Syntype Variegatum No. 1

55.6

48.6

33.4

0.87

0.69

0.83

120

Idem No. 2

53.0

50.6

29.6

0.95

0.58

0.80

125

Idem No. 3

47.7

43.1

340

0.90

0.78

MNHN, Sulu

43.7

40.0

27.3

0.92

0.68

0.87

130

MNHN, Philippines

34.0

32.0

21.0

0.94

0.66

0.88

125

MNHN, Philippines

46.3

40.4

28.0

0.87

0.69

0.80

125

MNHN

40.8

37.3

24.7

0.91

0.66

0.82

130

QM MO20733

48.3

42.3

29.7

0.88

0.70

0.88

130

AMS C30279

58.1

52.6

(37.0)

0.91

0.70

Total adult shells measured and rib counts

General mean values

0.91

0.67

0.84

128

40.2

Standard déviation

0.03

0.04

3.0

2.2

Largest specimen observed, AMS C30279 (see above)

ZOOSYSTEMA • 1999 • 21 (2)

Vidal J.

Table 15. — Measurements (in mm) and rib count of Acrosterigma oxygonum (Sowerby. 1833).

L/H

W/L

A°

Ribs

Syntype oxygonum No. 1

41.2

34.0

25.0

0.81

0.74

0.82

115

Idem No. 2

37 5

32.7

23.0

0.87

0.70

0.81

110

MNHN, China

36.8

31.8

23.0

0.86

0.72

0,95

120

MNHN, India

37 5

32.3

22.2

0.86

0.69

0.86

120

Idem

36.0

307

20.3

0.85

0.66

0.83

115

MNHN, Phuket

36.9

31.6

23.0

0.86

0.73

0.82

120

MNHN

42 8

37 0

26.0

0.86

0.70

0.80

110

Total adult shells rneasured and rib counts

General main values

0.85

0.70

0.85

117

38.9

Standard déviation

0.02

0.05

4.1

1.4

Largest specimen observed. MNHN (see above)

much wider, witli oblique seules straddling a very

thin inter médiate furrow. Ribs ol M PQ triangul-

ar with a slightly concave posterior side, and a

small, entirely smooth crestal fold. Anteriorly

ribs become progressively rounded, first a little

retro-ridged rhen top-ridged on anterior half.

Interstices always rounded (PQ excepted), and

slightly narrower than ribs.

Remarks

A. vttriegatum is very constant in characters. lt

cannot be confused wirh any other species, parti-

cularly because of its unique lunule.

Acrosterigma oxygonum (Sowerby, 1833)

(Fig. 5D-F; Table 15)

Cardium oxygmum .Sowerby, 1833:11g. 9; 1841 a: 107,

Types. — Two shells considered as syntypes in

BMNH, Cuming collecrion, labelled “Philippines”.

The dimensions and rib number giveu by Sowerby

could fit che specimen figured here (Fig. 5D, E); but

the locality given by Sowerby (China Sea) does not

agréé.

MaTERTAT. EXAMINER. — The tollowing lots in addi¬

tion to syntypes:

Zanzibar. (IRSN'B Daurzenberg).

India. Gulfof Munaar (BMNH Winckworth). —

Tuticorin (BMNH Wfnckwonh). — Ranteswaram

(M-NHN Vidal). — Vishakhapatman (USNM

622130).

Sri Lanka. (MNHN Staadt). - (1RSNB

Dautzenbcrg). (NHMW 839). — (AMS C03408

- (BMNH 1875-4-8-2 Holdworth - (USNM

149929).

Burina. S ol Akvab, Çiulf of Bengal (ANSP 239957,

293956). 10°37’N, 97M4T, Twin lsland,

Andaman Sca (ANSP 291908).

Thailand. H ai Nan Beacli. Phukec (MNHN

Vidal). - Nai Yang Reef, Phukei (MNHN

Vidal). — Kant Bette h (ZMUC). — Southern

Thailand (LACM 13488). — 9”11 N, 98°13’E, Gulf

ul l'hailynd (ZMUC).

Malaysia, l.àngkawi lsland, Andaman Sea

(ZMUC). — 6 Q 27’N, 99 D 50'E, Langkawi lsland

(AMS Loch).

Philippines. (NHMW). — (AMS). — Ticao lsland

(MHM\V).

China. (MNHN Denis 1945). — (LACM 50836). —

(MHMW).

Unidentified and unknown localities. Toujong Rhu

(AMS CI92747), — Three lors (MNHN).

DiSTRIfil. TloN, — (Fig. 6) Excepr for a lot said ro be

Iront Zanzibar (to he confirmée!}, exdusively in nor-

thern hémisphère: Gulfof Mannar, Andaman Sea and

Malaeea Str.tit, Gulf of J hailand. (South?) China,

Philippines. Published records l’rom Australia (South

Queensland and New South Wales) arc erroneous and

refer to A. kerdukttr.

Description

Shell medium-sized; ovoid, and almost équilatér¬

al, sometimes with posterior dorsal rnargin a

little straightened; hardly elongated (mean

l./H = 0.85: range 0.81-0.89) and not apprécia-

bly compressed (mean W/L = 0.70; range

0.66-0.74). Ribssciaight in projection.

Lunule about équivalent on both valves, well-

delineated, and appreciably hollowed on both

stdes. External colour beige, variegated with

rather dark brown to purple in large irregular

288

ZOOSYSTEMA ■ 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 16 . — Measurements (in mm) and rib count of Acrosterigma selene n. sp.

L/H

W/L

A°

Ribs

Holotype selene

27.3

26.4

17.7

0.97

0.67

0.68

120

Paratype No. 1

35.3

32.9

(23.6)

0.93

0.72

0.67

Idem No. 2

30.2

27.8

(20.0)

0.92

0.72

0.69

Idem No. 3

23.2

22.6

(16.0)

0.97

0.71

125

Idem No. 4

25.1

23.8

(16.0)

0.95

0.67

0.63

Idem No. 5

20.0

18.5

12.3

0.93

0.66

0.65

120

Total adult shells measured and rib counts

General mean values

0.94

0.66

122

36.6

Standard déviation

0.02

2.4

0.8

Largest specimen observed, paratype No. 1

(see above)

patches; interior white, excepr for iimbonal rays.

Hinge appreciably asymmetrical (avefage ratio

D = 0.85; range 6.80-0.95) and angled (average

< A = 117°; range 110“-1 20°). A long, thin, weak

umbonal sterigma on some siiclls in about 20%

of lots.

Mean rib numbet 38.9, range 36-41.

Rib morphology: on PQ (Fig. 5F), both parts of

équivalent vvidth, with a weakly marked médian

furrow and scales tbat are sometimes irregtdar;

interstices well-marked- Flsewhere, rjbs alvvays

triangular, except for first ones of AQ, whicb

become rounded; on MPQ (Fig. 5F), ribs first

retro-ridged, becoming retro-tuberculated ante-

riorly, with numerous stnall tubercules situated

on lower part of flank; anterior flank smooth,

and a signilicant smooth crestal fold présent. On

anterior half, anterior flanks of ribs become at

first pro-ruberculated at base, then pro-ridged,

with ridges rerminating in crestal tubercles, but

not reaching interstice; in contrast, posterior

flanks of ribs become and remain smooth throu-

ghout. Interstices triangular and smooth.

REMARK.S

A. oxygonum lias very constant characters, parti-

cularly an elaborate and distinctive rib morpho¬

logy. One lot of this species in BMNH (reg.

1910-12-13-15), from .Sri Lanka, is labellcd as a

type séries of "Cardiurn sanguineotinctn" Preston;

another lot AMS CG34080, Iront Sri Lanka, is

labelled as a cotype lot of the same taxon. 1 could

not find any vérification of this name in the lite-

rature, even in Adam (1971).

Acrostengrna selene n. sp.

(Fig. 5G-J; Table 16)

Types. — Ail shells from several dredging campaigns

in rhe vicinity of New ( aledonia, MNHN, Richer de

Forges ORSTOM collection. Holotype: a bivalved

shell (Fig. 5Q-I), 1AGON 1985, stii 416, 22°38’.S,

167°l4’ll, S zone ol Lagoon, 40-50 m. Paru type 1: a

loft valve (Fig. 5J), CORAIL 2 1988, stn CP25,

20"25’S, t61°05’F, I msdowin-Fairwav Plateau,

67-70 ni, Paratype 2: a le h valve, CHAI.CAL 2 1986,

stn DW 83, 23°20'S. 168*06'K, SK of New C'aledonia

ridge, 200 ni. T’aratvpe 3: a right valve, CHALCAL

1984, stn D39, 20 n 29'S, HfiMl’E,

Cliesterfield-Bellona Flateau, 40 m. Paratype 4: a left

valve, CORAIL 2 1988. stn DW 18, 20°44'S,

16P00’E, Lansdowne-Faîtway Plateau. 69 in.

Paratype 5: a bivalved shell, LAGON 1985, stn 379,

22“3) S 167" 11 Té, Grand Récit Sud, '’G m. Paratypes

6 to 10: six smaller valves or bivalved shells, from

Che.vterfield and Landsdowne-Fairway areas.

E rVMOLUCV. — Selene, greek name of the ntoon; an

allusion to the unique lunule of this shell ( lutiula :

smull moon in Latin).

MatERIAL EX/VMINEO. — The type sériés (eleven indi-

viduals).

Distribution. — (Fig. 6) SE ol New Calcdonia,

Chescerfteld-Bellona Plateau and Landsdowne-

Fairway bancs.

Description

Shell medium-sized, almost équilatéral, neither

elongated (L/H range 0.93-0.97) nor inflated

(W/L of holotype = 0.67) and very slightly trun-

cated on posterior margin.

Lunule well-delineated and extremely hollowed

ZOOSYSTEMA • 1999 • 21 (2)

289

Vidal J.

Table 17. — Measurements (in mm) and rib count of Acrosterigma discus n. sp.

W/L

A°

Ribs

Holotype discus

29.2

27.5

15.6

0.94

0.57

0.78

125

Paratype No. 1

26.9

25,0

15.6

0.93

0 62

0.79

120

Idem No. 2

25.3

23.6

14.7

0.93

0.62

0.82

120

Idem No. 3

24.4

22 5

13.0

0.92

0 58

0.85

120

Idem No. 4

24.3

23.0

12.8

0.95

0.56

095

120

Natal. Natal Mus. S4039

29.2

26.0

(15.8)

0.89

0.61

= 1.0

120

MNHN, Kenya

23.6

220

13.9

0.93

0.63

= 1.0

120

MNHN, Comores

24.1

22.1

13.6

0.92

0.62

0.86

115

MNHN, Madagascar

23.9

22.3

(14.0)

0.93

0.63

115

ZMA, Moluccas

25.5

23.8

14.2

0.93

0.60

0.76

115

USNM 321695

31.5

28.0

17.8

0.89

0.64

Total adult shells measured and rib counts

General mean values

0.93

0.60

0.90

120

56.3

Standard déviation

0.02

0.03

0.08

2.9

3.2

Largest specimen observed, USNM 321695 (see above)

on both sides, the righr being wider (Fig. 51, J).

Externsl colour yellowish-beige, with irregular

darker zones and pink to orange spots. Interior

whitisli with traces of external spots, and two

pink umbonal rays which are also visible exte-

riorly. Mitige moderately angled (< A about

120°), and strongly asymmetrical (ratio D range

0.63-0.69), in concrast to the shell itself, which i.s

almost perfectly équilatéral, as previously raen-

tioned.

Mean rib number 36.6 (range 35-38).

Rib morphology: on PQ, anterior smooth part of

ribs much wider than posterior part; intermé¬

diare furrow very thin and not always visible in

adult région. Scales elongated, sometimes pris-

matic, straddling the longitudinal furrow, and

often almost longitudinally disposed rather than

oblique. On MPQ, ribs triangular and slighrly

asymmetrical with posterior flank fiat, shorter,

and more abrupt, and anterior flank slightly

convex. Ribs dtemselves pracdcally smooth (rare-

ly slightly retro-ridgcd), with a cresral fold mark

ed by régulai', short undulaiions derived from

PQ scales. On anterior hall, ribs become progres-

sively less triangular, then rounded, lirst

retro-ridged or crenulated, then top-ridged.

Interstices rounded, smooth, as wide as ribs.

Remarks

Acrosterigma selene superficially resembles

A. mauritianum in shape and colour, but has

fewer ribs and its rib morphology is different

(ribs more triangular). It differs from ail Acro¬

sterigma species by its unique lunule.

Acrosterigma discus n. sp.

(Fig. 5K-M; Table 17)

Trachycardium cf. mauritianum (Deshaves). - Drivas

& Jay 1988a; 17, % 14.

Types. — Ail from Mauritius, in MNHN, H. Fischer

coll.: holorype and four paratypes.

ETYMOLUGY. — Shape close to a dise.

MATERIAL EXAM1KED. — The following lots in addi¬

tion to the type material:

South Africa. N Zululaud (in Natal Muséum):

Berween Rhanga Neck and K dm Rav. 34 ni (D9816

Herbert). — l.eadsmun Shoal. 23 ni (1.2453 Herbert).

Mririuç Nnudc 1987. SI', of Kosi River Mouth:

stn ZA'l, 26"56.9'S, )2“54.5'E, 50 m (D9219). —

Sm 7.A2. 26“56.U'S. _U°54.7’E. 40 m (D7304). —

Soi 7.A9, 26“44,6’S. 32“55-VF. 40 m (D6246). —

Sm TA 12, 26‘55.0'S, 32 u 55.S'E, 65 m (08065). —

Stn ZA29. 26 U 44..VS. 32“54.8'E, 48 rn (D8703). —

-Stn 7,330. 26°54.3 S, 32 0 55.5T, 50 m (D7932):

N MDP 1990. o fl Kosi Bay; stn ZA37, 2654.0S,

32°55.5H, 40 m (S,3973). — Sm 7A41, 26 fr 52.9’S,

32’54.3’E, 49 m (S4641). — Sm 7A48, 26 l, 53.5’S,

32’55.6T:. 51 iq (S4039). - ZR12 27°00.4’S,

32°55.3’E, 67 m (S6335).

Mauritius. (MNHN). — (BMNH). — (NHMW). —

290

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

(AMS 047231 ). — (USNM 149930, 321695), —

(ANSP 315581). — 21 “2 LS, 65°52’E, SE Rodriguez.

Island, 60 m (USNM Brunn).

Madagascar, THOMASSIN'S Survey 1962-72,

Tulear area (MNHN): stn DI 6, 23*29’36"S,

43 D 4L35"W, 13-17 m. — Stn D2I, 23“20’35”S,

43“41 ’35"W, 50 in. — Stn D50, 23°29’06”.

43°22’24’’W. 29 m. — Stn D51 23"29'00"S,

43“52'5r'W, 10 m. - Stn 230, S of Grand Récif,

21 m. — Stn 240, Grand Récif, 36 m. — Stn 261,

Grand Récif, 26 m. — Srn 615. 616, Beach N of

Fiharenana, 8 m. — Srn 619, 621, fulear Lagonn,

South Pass. — Stn 738. Tulear I.agoon, South Pass,

12 m.

Comores. 43 ni (MNHN Plaute 1975).

Kenya. Wasin Channel (off Michangani). Shimoni,

10 m (MNHN).

Sri Lanka, (BMNH 1937-7-9-31-33).

Singapore. Scivrosa Island réclamation (AMS

C310566).

Moluccas. (ZMA).

DISTRIBUTION. — {Fig. 6) Mafnly lndian Océan, very

abundant in Mauritius, whcrc it îs sympatric with

A. mauritianum and similarly collected expisce-, record-

ed in Ui Réunion only by Drivas & Jay (1988a: 17);

Known, but rare, as far east as the Moluccas.

Description

Shell medîum-sîzed to small, almost circulai'

(L/H range 0.87-0.97), almost perteerly équilaté¬

ral, sometîmes very slightly truncatcd on PQ,

and very rarelv slightly expanded backwards.

Shell also srrongly flattened (W/L range

0.53-0.65), giving it a “disk-like” appearance.

Lunule extrcmcly narrow to non-existent on

both valves, and may be absent altogether, svuh

reduced ribbing practically rcaching margin. On

right valve this margin appreciably raised near

umbo. Periostracum rhin and bomogencous.

External colour light beige, more or less motrled

with brown-purple, and umbo sometimes with

two radial purplish rays. Interior white with pur-

plish stains correspondmg to external markings.

Hinge nearly symmetrical, but with ratio D

always less titan 1.00 (range 0.76-1.0). Angle A

rather high. ranging front 115 to 125°.

Mean rib number 56.3 (range 53-63).

Rib morphologv: on PQ (Fig. 5M), anterior

smooth part of eacli rib unusually large, axial fur-

row very thin and often confused with posterior

part of rib, which is also very thin, sometimes

practically non-existent. Persistent scales inserted

in the furrow are high. elongated, ellipsoidal in

section and slightly twisted; they entroach slight¬

ly upon anterior part of rib. The.se scales placcd

ai a narrow angle to axis of shell, altnost longitn-

dinally. On MPQ, ribs asymmctrically triangular

with posterior flank sceepcr; they are general ly

smooth (rarely with ridges in continuity with

scales of PQ as in Fig. 5M). The rib top (apex of

triangle) bears a smooth, regular, longitudinal

crestal fold (Fig. 5M: sbining line). Interstices

rounded and smooth. On anterior half of shell,

ribs become progressively asymmerrically roun¬

ded, more or less rétro-ridged or festooned, then

rop-ridged; on first ribs of AQ, top scales become

tubercular and a lirtle irregular, and tend to take

on a vaguely concentric alignment.

Remarks

Acroslerigma discus resembles A. mauritianum

with which it lias been confused. Llowever, it dif¬

fère in its discoid shape, higher rib number, and

more triangular shape of the ribs on MPQ.

Acrosterigma mauritianum (Deshayes, 1855)

(Fig. 7A, B; Table 18)

Cardin >n mauritianum Deshayes, 1855: 331.

Trachycardium nebulosum (Reeve) - Drivas & Jav

1988a; 17, fig. 15; 1988b: 140, pl. 55 [Not Cardium

nebulosum Reeve, 1845 Acrosterigma simplex

(Spengler 1799)].

Fîtes. Thrce synrypes in BMNH. Cuming collec¬

tion, not catalogued, from Mauritius (stomach of a

large tish). I fie largest is figured bere (Fig. 7 A, B).

Mai CRIAI EXAMINEE). — The following lots in addi¬

tion to the ihree synrypes;

Mauritius. (MNHN Jousseaume 1921). — (MNHN

Carrié 1911). (MNHN J. de Lh.). — (MNHN

Vidal). — (AMS Cl47223. 147224, 147226,

147227, 038053).— (BMNH).

Seychelles. Amirauté Island, stn F.16 (BMNH

1910-8-31-701 Gardincr).

R eues 2 1880 MNHN: stn 4, 05°O8'S, 56“35'E,

32 ni. — Stn 18, 05°45 S, 56°35'E, 50 m. — Stn 20,

05°36‘S. 56 C, 19T. 35 m.

Sri Lanka. (BMNLr 1937-7-9-31-33)?

Philippines. Port Calera, Mindoro (BMNH

1914-6-12-52)?

Unknown locality. (MNHN).

DISTRIBUTION. — (Fig. 6) Mauritius and Seychelles,

sometimes présent well-preserved in the stomachs of

ZOOSYSTEMA • 1999 • 21 (2)

291

Fig. 7. — A. B. Acrosterigma mauritianum, syntype; C, D, Acrosterigma uniornatum, holotype; E. F. Acrostengma uniornatum, para-

types 1 and 2; G, Acrosterigma uniornatum. paratype 28; H, Acrosterigma uniornatum, paratype 29; I, J, Acrosterigma profundum,

holotype; K, L. Acrosterigma profundum, paratype 1; M, Acrostengma profundum, paratype 2; N, O, Acrosterigma amirante, holo¬

type: P, Q, Acrosterigma amirante, holotype; view of PQ and MPQ on both valves. Scale bars: 10 mm.

ZOOSYSTEMA • 1999

JE 1a

Il m

' ■ 1

i-' J

Taxonomie review of the elongated cockles

Table 18. — Measurements (in mm) and rib count of Acrosterigma mauritianum (Deshayes. 1855).

H L W L/H W/L D A° Ribs

Synt. mauritianum No. 1

33.1

29.0

20.8

0.88

0.72

=1.0

120

Idem No 2

32.8

30.0

21,0

0.91

0.70

0.98

110

Idem No. 3

29.8

27.3

20.0

0.92

0.73

0.90

115

MNHN, Mauritius

37.0

31.6

23.3

0.85

0.74

0.89

115

MNHN, Mauritius

30.6

274

19.6

0.90

0.72

0.92

120

MNHN, Seychelles

26.2

21.5

17.0

0.82

0.79

0.96

120

MNHN, Mauritius

38.5

32.8

23.7

0.85

0.72

=1.0

120

MNHN. Mauritius

39.0

33.7

24.0

0.86

0.71

0.95

120

Total adult shells measured and rib counts

General mean values

0.89

0.71

0.95

118

44.1

Largest specimen observed. MNHN Mauritius (see above)

large fishes; old lots said to be from Sri Lanka or the

Philippines are probably mislocalized.

Description

Shell mediiim-sized, ovoid and alrnost équilatéral,

sometimes vvith a slight fruncation at MPQ mar-

gin and Irequently a slight flattcning of médian

ventral margin. Sliell moderately elongated

(mean L/H = U.8*); range 0.82-0.94) and inflated

(mean W/l - 0.71; range 0.64-0.79).

Lunule narrow, elongated and slightly holltnved

on right valve, very thin to nearly absent on leit.

Lxtcrnal colours beige with variably developed

pinkish patches, sometimes with rvvo umbonal

rays; interior vvbite, except for pink umbonal

rays, Hinge almost symmetrical (mean ratio

D = 0.95; range 0.89- LOO) and moderately

angled (mean <A = 118"; range 110°-125°).

Mean rib number 44.1» range 41 -47,

Rlb morphology: on PQ, anterior part of ribs

wide, posterior part and intermédiare furrow

very narrow and poorly differemiated. Scales

short and sometimes slightly tubcreular; inter¬

stices narrow. On MPQ ribs gcnerally mutidly

triangular ro rounded and asymmetrical, rarely

fully triangular. Posterior flank shorter ai)d more

abrupt, with a clean crestal fold; ribs retro-

tuberculated, rarely entirely smooth, On anterior

half, ribs become rounded, retro-tuberculated or

crenulated, then more or less elearlv top-ridged.

On first ribs of AQ, top l itiges widen and beeo-

me tubercular and irregular, sometimes forming

vaguely concentrical alignments.

Remarks

A. mauritianum is well-characterized by its shape

and colour, and distinctive rib characters on PQ

and MPQ; it differs also from the other species

of the species-group by its less triangular ribs and

the characteristics of AQ, which may be transi-

tional with those of the species-group of A macu-

losum.

Species-group of

Acrosterigma uniornatum n. sp.

Diagnoses. — See Table 5.

Incxuded SPECIES. — 1) Recent: four new species:

A. uniornatum-, A. projitndum; A r amirauté-, A. sulua-

num. 2) Fossil: lack of suffieient information; a new

species described here A, paulayi n. sp.

Acrosterigma uniornatum n. sp.

(Figs 7C-H; 8A; Table 19)

Types. — Ail type lots coinc Iront rhree relatively shal-

low banks on the Melanesian Border Plateau, within

and in the vicinity of Wallis and Futuna Territory,

MUSORSTOM 7 Campaign 1992 in the south-wes-

tern Pacific, in MNHN. Bouchet, Mêtivier, Ricller de

Fors»cs coll. Holocype: a right valve (Fig. 7C, D), stn

DW 29. 12°31'S, 176 WW. Watetwitch Bank,

500 in, Puratype I: a left valve from die saine locality

(Fig. 7t), l’aratype 2: a right valve fmitt the same locali¬

ty, Paratype 3: a right valve front the sanie locality

(Fig. 7F). Para types 4 to 25; odd valves, same locality.

Pararypes ?ù-2~: two valves, stn DW 530. 12 D 33'S,

176°39’W, Waterwitch Bank, 580-600 m. Paratvpes

28 to 36: odd valves, stn DW 538, 12°31’S, 176°40',

ZOOSYSTEMA • 1999 • 21 (2)

293

Fig. 8. — A, Acrosterigma uniornatum, holotype; view of median-anterior part: B, Acrosterigma amirante, holotype: view ot

median-anterior part; C, Acrosterigma profundum, holotype; view of median-anterior part: D. Acrosterigma suluanum, holotype;

E, Acrosterigma suluanum, holotype; F, Acrosterigma suluanum, holotype; view of median-anterior part; G, Acrosterigma paulayi',

type sériés, (PI, P2, P3 = paratypes; H = holotype); H, Acrosterigma paulayi, holotype, view of MAQ and part of AQ; I, Acrosterigma

paulayi, paratype 1 ; detail of PQ and part of MPQ. Scale bars; A-C, F, 4 mm; D, E, G, 10 mm; H, I, 5 mm.

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Waterwirch Bank, 275-295 m. Paratype 37: one

valve, stn DW 542, 12°26’S, 177°28’W, Combe

Bank, 370 m. Paratypes 38-39: rwo odd valves, stn

DW 546, 12°27'S, I77 <, 29’W, Combe Bank,

550-552 m. Paratypes 40 to 44: odd valves, stn DW

588, 12°17’S, 174°45’W, Field Bank, 490-500 ni.

EtymoloGV. — Ornamentation of the ribs unifornt

on ail the parts of the shell.

Material EXAMINED. — The type sériés.

Distribution. — (Fig. 9) Known only from the vici-

nity of Wallis and Futuna Isiands. It is unclear whe-

ther this species lives at 500-600 m whcrc the shells

wcrc dredged, or hâve rafted down from shallower

depths. Anyhow, its depth range is probably greater

than is usual for the genus. This occurs also for the

other species of the spccies-group.

Description

Shell small, ovoid, practically équilatéral, never

truncated posteriorly, moderately elongated (L/H

about 0.83) and rather globose (extrapolated

W/L about 0.82).

Lunule well-delineated, nearly similar on both

valves, small, and slightly hollowed. External

Table 19. — Measurements (in mm) and rib count ot Acrosterigma uniornatum n. sp.

W/L

A°

Ribs

Holotype uniornatum

22.4

18.1

(15.2)

0.81

0.84

130

Paratype No. 1

21.9

18.0

(15.4)

0.82

0,86

=1.0

Idem No. 2

22.1

18.1

(15.4)

0.82

0.80

130

Idem No. 3

21.7

17.9

(14.4)

0.82

0.80

130

Idem No. 4

16.3

14.0

(12.0)

0.86

=1.0

Total adult shells measured and rib counts

General mean values

0.83

0.82

100

128

38.2

Standard déviation

0.02

0.03

4.0

1.4

Largest specimen observed, the holotype

ZOOSYSTEMA • 1999 • 21 (2)

295

Vidal J.

Table 20. — Measurements (in mm) and rib count of Acrosterigma profundum n. sp.

L/H

W/L

A 0

Ribs

Holotype profundum

36.7

28.2

(23.6)

0.77

0.84

120

Paratype No. 1

34.4

288

(22.4)

0.84

0.78

*1.0

Idem No. 2

30.0

24.4

(19.0)

0.81

0.78

*1.0

Idem No. 5

32.3

26.3

(21.8)

0.81

0.83

125

Idem No. 6

21.1

18.0

(13.2)

0.85

0.73

0.95

Idem No. 7

19.5

16.6

(13.0)

0.85

0.78

120

Total adult shells measured and rib counts

General mean values

0.82

0.79

1.03

122

56.9

Standard déviation

Largest specimen observed, the holotype

0.03

0.04

0.06

2.3

3.1

colour uniform: white on juvénile part, pale yel-

low on .idult part. luterior white, verv rarely with

traces of pink; no umbottal rays visible. Hinge

symraetrical (ratio D c. 1.0), and moderately

angled (< A about 130°). On right valve cardi-

nals show a greater degree of fiision at their base

than is typical for fhe genus,

Mean rib number 38.2, range 36-41.

Rib morphology: on PQ (Fig. 7G, H), interstices

thin, ribs liai with a central longitudinal furrow.

Anterior smooth part narrower than posterior

part, which bears elongated, thin, straight,

oblique scales, offen encroaching upon axial fur¬

row and even anterior part. On MPQ (Fig. 7C,

H), saine features occur but posterior zone is

slightly raised and ribs beconte slightlv flattened

triangles. Longitudinal furrow progressively

becontcs less distinct, and anterior smooth part

of rib becontcs its anterior flank. Scales rentain

about the same as on PQ. On anterior naïf

(Fig. 8A), ribs beconte more rounded; scales

become more numérotas and change, firsc into

posterior flank-ridges, tlten into moderately

imbricated transverse top-ridges.

Remarks

The discussions bclow indicate différences bet-

ween Acrosterigma uniornatum and the other four

species of the species-group.

Acrosterigma profundum n. sp.

(Figs 7J-M; SC; Table 20)

TYPES. — Holotype: a right valve (Fig. 71, J),

BATHUS 4 1994, stn DW 894, 20°16’S, 163°52’E,

N of New Caledonia, 245-268 m (MNHN, Bouchet,

Métivier, Richet de Forges). Paratype 1: a left valve

(Fig. 7K. F). New Caledonia LAGON 1 l J87. stn 830,

20°49'S. 165“19’E, NF, zone of I.agçion, 105-1 10 ni

(MNHN idem b Paratypc 2; a left valve (Fig. 7M),

BATHUS 4 1994, sut DW 896, 2()"16'S, 163 U 52’E,

N of New Caledonia, 315-350 ni (MNHN idem).

Paratvpe 3: a left valve, broker and incomplète,

BATHUS 1 1993, sut DW 691, 2H"35 S, I64"59’E,

F, coâst of New Caledonia, 227-250 m (MNHN

idem). Paratvpe 4: -J very small left valve, front same

locality as pars type 1. Paratypc 5: a right valve, stn 14

of 1947 canipaign scaward (N) of west end ol Bikini

Atoll, Marshall Islandï, 177-244 m (USNM 598697,

RW Russel coll.). Paratypc 6: a left valve, same lot as

paratype 5. Paratype 7: a left valve, same lot as para-

types 5 and 6.

Exvmoiogy. — Profundum: deep in Latin; an allu¬

sion to the relative deep environment of tltis shell.

M A 1 1KIA i r x a M INED. — The type sériés.

Distribution. — (Fig. 9) North and east of New

Caledonia, at relahvcly grcat water deptlts for

Cardiidae other than Protocardiinae; présent al.so in

Bikini Atoll, Marshall Islands (paratypes 5 to 7).

Description

Shell medium-sized, ovoid and ncarly équilatéral

with no posterior truncation, moderately elonga¬

ted (L/H range 0.77-0.85) and globose (extrapo-

lated W/L range 0.73-0.84).

Lunule small, rallier intperfectly delineated and

nearly the .same on liotli valves; umbonal margin

raised, fortning a double wall. External colour

pattern is unique: on adult part, PQ cntirely

pink to orange, MPQ bears large, pink to orange,

irregularly concentric splashes, and anterior hall

296

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 21 . — Measurements (in mm) and rib count of Acrosterigma amirante n. sp.

W/L

A°

Ribs

Holotype amirante

23.9

20.0

16.5

0.84

0.82

0.88

130

Paratype

24.2

0.86

of the shell beige, with widely scattered small

splashes. lnterior white with sortie exterior

splashes showing through; no umbonal rays

visible. Flinge synimetrical (ratio D c. 1.0) and

moderately angled (< A about 120°). On right

valve, cardinals show an unusual degree of fusion

at their base, as in the prcceding species.

Mcan rib number 56.f), range 55-63.

Rib morphology, on PQ (Fig. 7M), interstices

thin; ribs Hat with a central longitudinal furrow,

anterior smonth put as widc as or narrower than

posterior. F.longated, thin. straight, oblique scales

de vélo p on posterior part, straddling axial furrow

and encroaching slightly upon anterior zone. On

posterior hall of MPQ (Fig. 7M), the same fea-

tures appcar, but posterior zone slightly taises

and ribs bccomc Hatlv triangular; longitudinal

furrow progressively disappears and anterior

smooth zone becomes anterior flattk of the rib.

On anterior half ofMPQ (Fig. SC), ribs becomc

more rounded, scales becotrte progressively short-

er and more numerous and change into posterior

ridges, rib tops beconiing smooth; at the same

tinte, numerous very thin oblique serrations

appear on anterior flan les of ribs, ehangirig ante-

riorly into small ridges or tuberçles. On MAQ

(Fig. 8C), this sculptural évolution leads to ribs

which are slightly squ.ire-sided, serrated on both

sides, and slightly overhanging interstices. In AQ,

sonie latéral serrations join on top of ribs, forming

thin, free top-ridges, which are not imbricated; the

First ribs of this quarter slightly degenerate, their

ornamentation beconiing less régulât

Remarks

Acrosterigma profiindum has a very elaborate and

spécifie rib morphology that cannot be confused

with any other; it is separated front A, uniorna-

tum by its larger size, higher rib number, and

particular rib ornamentation on the médian part

of the shell, i.e. the small dense serrations on the

anterior flanks of the ribs.

Acrosterigma amirante n. sp.

(Figs 7N, Q; 8B; Table 21)

Types. — I Iolotype: a bivalved shell (Figs 7P, Q; 8B),

stn HI6, Amirauté tsland, Seychelles, 71.3 ni (BMNH

Ref. IM 10-8-31-702, J. S. Gardiner cnlh). Para type 1:

a parrially broken and incomplète lefi valve,

Thontassin Survey, stn D36. 43 U 39’55"W,

23°29'08”S, Tnle.tr, Madagascar, 280 m (MNHN).

EVYMOl.OGY. — Front Amirante Islands, Seychelles.

MateriAL examined. — The type sériés.

Distribution. — (Fig. 9) Indian Océan: Amirante

Islands and Madagascar.

Diagnoses. — Shell small, ovoid, not truncated post-

eriotly, ncariy équilatéral, moderately elongated

(L/l 1 = 0.84) and radier globose (W/l. - 0 83),

Lunule somewhai intpcrlectly delincatcd, small,

almost équivalent on both valves, its niargin r.tised ncur

umbo, t sternal coloui a unilorm pale yellow, exeept

fora pink unlbo. lnterior whire, with a pink concentnc

hand in rhe middle and two pink umhonal rays. i linge

slightly asymmetrical (ratio D = 0.86 and 0.88) and

moderately angled (< A = 130°). On right valve, cardi¬

nals more than usually fused at base once again.

Rib number 55.

Rib ntotphology: ont PQ (Fig. 7P. Q), interstices vety

thin, ribs very ll.it with an axial longitudinal dtin fur¬

row separaring two zones of sintilar widtlt. F.longated,

thin straight, oblique scales occupy entire posterior

zone and encroach upon anterior zone, straddling

axial furrow. Ou MPQ, Inatures are at lirst sintilar,

but rhen (Fig 8B) posterior zone-becomes slightly

raised and ribs hecome llatly niangular; axial furrow

gradtially disappears and anterior sntnoth zone

becontes anterior Fin n k of the rib. Scales shorren,

becoming only more numerous small flank-ridgcs;

rib tops hecome smooth. On anterior part of shell

(Fig. SB), ll.ink- ridges extend again onlp rib tops an cl

cventually beconte transverse rop-ridges, at first free

and thin, then, on AQ, becoming wide, imbricated

and even irregularly tubercular near lunule.

Remarks

Acrosterigma amirante resembles A. uniornatum

in dimensions, shape, and colour, but is distin-

ZOOSYSTEMA • 1999 • 21 (2)

297

Vidal J.

Table 22. — Measurements (in mm) and rib count of Acrosterigma suluanum n. sp.

L/H

W/L

A°

Ribs

Holotype suluanum

25.4

20.5

(15.0)

0.81

0.73

130

Paratype No. 1

43.6

34.0

(30.0)

0.78

0.88

0.95

Idem No. 2

33.3

28.6

(22.0)

0.86

0.77

130

Idem No. 3

27.0

(24.0)

0.89

130

Idem No. 4

30.3

25.7

(19.2)

0.86

0.75

130

Idem No. 5

28.5

23.8

(19.0)

0.84

0.80

130

Total adult shells measured and rib counts

General mean values

0.83

0.80

0.95

130

43.8

Standard déviation

Largest specimen observed, the paratype No. 1

0.03

0.06

1.3

guished from chat species by the lunule shape,

rib number, and details of rib ornamentation. It

is close to A. profundum in lunule shape and rib

number, but difFers in rib morphoiogy having no

high bi-crcnulated ribs on the médian part of the

shell.

Acrosterigma suluanum n. sp.

(Fig. 8 D-F; Table 22)

Type. — Holotvpe a right valve (Fig. 8D-F), stn 5577

of US B F, Tawitawi Island, Sulu Archipelago,

Philippines. 439 tti (USNM 299435). Paratypes l, 2

and 3, a left valve and [wo right valves (rom Sunasald,

Japan, dredged 150-35 in (ZMUC coll. Morrensen).

Paratypes 4 and 5, tvvu right valves, 34°20'N,

130°t0'H, Japan, 113 ni, sanie reposiiory and collec¬

tion.

Etymoloc.y. — From Sulu Archipelago, Philippines.

MaTERIAI F.XAM1NHP AND DISTRIBUTION. — (Fig. 9)

The holorype, from north of Tawitawi Island,

Philippines, ac a great water depth for a Cardiidae and

five paratypes from |apan.

Description

Shell small, ovoid, not truncated posteriori}’ and

almost équilatéral, moderately elongated (mean

L/H = 0.83) and tumid (mean W/L = 0.80).

Lunule small, rather iraperfectly delineated, its

umbonal margin rai.sed. Colour yellow with

more orange splashe.x; inrerior parrially pink.

Hinge slightly asymmetrical (ratio D = 0.95) and

moderately angled (angle A — 130°). On right

valve, cardinals slightly more than usually fused

at base.

Mean rib number 43.8, range 41-45.

Rib morphoiogy: on PQ, interstices very thin,

ribs very fiat with an axial furrow separating rvvo

zones ol similar vvidth. Slightly elongated, tuber-

cular, oblique scales on posterior part encroach-

ing upon anterior zone, srraddling axial luirow.

On MPQ ribs become flatly triangular and

asymmetrical, then rounded. Scales disappear

and are replaced posteriorly with numerous very

thin short ridges or tubercles, situated at base of

rib, touching interstice, On MAQ (Fig. 8F),

large scales reappear on posterior flanks, surirn-

posed ou the lattcr line ornamentation; a similar

Une heading but thinnet and more tubercular

appears also at base ol anterior fiank of ribs. On

AQ (Fig. SF), the fine bcading of both sides

disappears and posterioi large scales proceed to

top and anterior flank of ribs, forming free

top-ridges which arc not imbricated.

Remarks

Acrosterigma suluanum shares with A. profundum

the character of having fine ornaments on both

sides of the ribs on MAQ (différence with

A. uniornatum and A. amirauté thaï hâve orna¬

ments ordy on the posterior sidc). On A. sul.ua-

num, however, the ribs arc rounded and the fine

latéral ornaments are situated at rhe base of the

ribs, touching lhe interstice, whilsr on A. profun-

taon the ribs are squared and the fine ornaments

arc separated from the interstice and slightly

overhang it. In addition, the rib number of pro-

jundum is higher, with an average of 58.4,

against 43.8 on suluanum.

298

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 23. — Measurements (many extrapollated, in mm) and rib count of Acrosterigma paulayi n. sp.

L/H

W/L

A°

Ribs

Holotype paulayi

29.5

25.5

20.0

0.86

0.78

1.27

Paratype No. 1

22.0

19.0

14.6

0.86

0.77

130

Idem No. 2

22.0

19.0

14.6

0.86

0.77

Idem No. 3

22.0

19.0

14.6

0.86

0.77

130

Total adult shells measured and rib counts

General mean values

0.86

0.77

1.27

130

48.7

Standard déviation

Largest specimen observed. the holotype

0.00

0.0

1.8

Acrosterigma paulayi n. sp.

(Fig. 8G-I; Table 23)

TYPES. — .411 matcri.il (Fig. 8G-I) front Niue Island,

Neogene fossil réel, will be stored in USNM, Paulay

colI. Rel. FN1UE 2b. Holotypc: an incomplète lelt

valve. Paratype 1: an incomplète right valve. Paratype

2: an incomplète light valve. Paratype 3: an incom¬

plète riglu valve,

EtyMOLOGY. — In honour ot Dr Cîustav Paulay.

Mateiual kxamined and DISTRIBUTION. — (Fig. 9)

The type material, plus Five valves in poor State of pré¬

servation, ail from Neogene fossil reef of Niue Island.

Description

Shell small to metlium-sized, ovoid and nearly

équilatéral wirh no posterior truncation, ntodera-

tely elongated (l./lf = c. 0.86) and globose

(extrapolated W/l. = c. 0.77).

Lunule small and well-delineaced, about équiva¬

lent on both valves, fiat and slightly bending

towards interior ol shell. Colours not preserved.

Hinge rather symmetrical and moderatelv angled

(< A= 130°).

Mean rib number 48.7. rangé 47-51.

Rib morphology: un PQ (Fig. 81), interstices

narrow, ribs fiat witli central longitudinal furrow

not or hardly ntarked. anterior smooth zone

wider than posterior. Elongated, thin, straiglu,

oblique scalcs develop on posterior part, not

straddling axial furrow and not encruaching

upon antetior zone. On MPQ (Fig. 81), ribs

become flarly rounded, then square-sided with

rounded tops; posterior scales change into more

numerous flanlc ridges or marginal serrations;

possibly appearance of anterior marginal serra¬

tions (observation difficult, due to the imperfect

State of préservation of the shells). On MAQ

(Fig. 8H), latéral serrations procced up to top

and join, forming a herringbone structure. On

AQ herringbone structure changes into straight

concentrical ridging, not imbricaced.

Rl MARKS

Acrosterigma paulayi is close to A, profundum; it

differs in having a more depressed lunule, shorter

oblique scales on PQ not encruaching upon the

anterior smooth zone, no squared ribs elsewhere,

no or less raarked anterior serrations or scales on

the ribs of the médian part, more pronounced

herringbone structure on the anterior part of MPC )

and on MAQ. In addition, the rib number of

A. paulayi is lower (48.7 versus 58.4 in profiirtdum).

Species-group ot Acrosterigma maculosum

(Wood, 1815)

iNCtUDI D SPECIES. — 1) Rccent: A. maculosum

(Wood. 1815) wirh a new subspecies, bowensc;

A. impolitum (Sowerby, 1833); A. transcenderts

(Mclvill & Standen, 1899); A. seurati n, sp.; A. dian-

tbinum (Melvill & Standen, 1899); A. punctolineatum

Healy & Lantprell, 1992; A. bobbsae n. sp.; A. simplex

(Spengler, 1799). 2) Fossil: see above for succint data.

Diagnüsis. — Sec Table 5.

Acrosterigma maculosum (Wood, 1815)

(Fig. 10A-H; Table 24)

Cardinm maculosum Wood. 1815: 218, pl. 52, fîg. 3.

Not maculosum Sowerby in Broderip & Sowerby,

1833: 85 |= C. prislipleura Dali, 1900)].

Not C. maculosum. - Sowerby, 1839: 18, fîg. 123

[= C. vértebratum Jonas, 1844],

ZOOSYSTEMA • 1999 • 21 (2!

299

Fig. 10. — A, B, Acrosterigma maculosum. possible syntype of C. multistriatum Sowerby: C. D, Acrosterigma maculosum, syntype of

Cardium arenicola Reeve: E. Acrosterigma maculosum. specimen (rom Shikoku. Japan. MNHN; detail of PQ and last rib of MPQ;

F, Acrosterigma maculosum, specimen from Japan, MNHN: G, H. Acrosterigma maculosum howense , holotype; I. J. Acrosterigma

impotitum. syntype: K. L, Acrosterigma impolitum. Holotype of Cardium Peauforti Prashad: M. Acrosterigma impolltum, specimen

from Penang (Malaysia), MNHN; N, Acrosterigma Impolitum, specimen from Queensland, MNHN; detail of médian marginal part (an

optical illusion can show the notched interstices as ribs). Scale bars: A-D, F-M, 10 mm; E, 2 mm; N. 4 mm.

ZOOSYSTEMA • 1999

Taxonomie review of the elongated cockles

Fig. 11. — Distribution of A. maculosum, A. maculosum howense and A. sorenseni.

Ca.rd.ium multistriatum Sowerby in Broderip &

Sowerby, 1833: 85. — Sowerby 1841b: Sp. 47, fig. 59.

Cardium arenicolmn Reeve, 1845: Sp, 78.

Types. — Cardium maculosum: the shell figured by

Wood not craced. Cardium multistriatum: the shell in

Cuming collection front Santa Helena, clted and

figured by Sowerby not traccd. Howcver, a loi label-

led C multistriatum front Santa Helena (West

Columbia) is présent in BMNH, Cuming collection

(see also Reeve 1845: Sp. 76). These shells (Reg.

1995079/1-2-3) can be considered possible syntypes

of C. multistriatum , the largest figured here (Fig. 10A,

B). The locality is certainly erroneous, as no similar

Table 24. — Measurements (in mm) and rib count of Acrosterigma maculosum (Wood, 1815).

L/H

W/L

A 0

Ribs

Syntype(?) multistriatum

36.2

29.2

20.7

0.81

0.71

1.14

105

Syntype arerticola

42.0

32.6

23.0

0.78

0.71

=1.0

100

MNHN. Zanzibar

23.2

20.0

15.2

0.86

0.76

= 1.0

120

MNHN, Red Sea

30.0

23.4

17 9

0.78

0.76

= 1.0

120

MNHN, Sri Lanka

44.7

34.7

26.5

0.78

0.76

= 1.0

105

ZMUC. Andaman Is

23.1

20.2

14.2

0.87

0.70

=1.0

115

ZMUC. Philippines

41,6

35,1

25.8

0.84

0.74

=1.0

105

MNHN. Hong Kong

20.2

19.0

12.2

0.94

0.64

= 1.0

120

MNHN. Japan

30.0

250

19.1

0.83

0.76

=1.0

110

USNM 747210, Moluccas

30.0

23.4

(16.4)

0.78

0.70

AMS C077245, Queensland

36.1

29.1

21.8

0.81

0.75

=1.0

120

MNHN. Japan

51.6

42.9

31.6

0.83

0.74

=1.0

105

Tolal adull shells measured and rib counts

102

101

136

General mean values

0.84

0.71

1.02

115

51.7

Standard déviation

0.04

0.05

0.09

5.9

4.5

Largest specimen observed, MNHN, Japan (see above)

ZOOSYSTEMA ■ 1999 • 21 (2)

301

Vidal J.

cardiid lias ever been found on the Pacific coast of the

Americas. Qtrdiunt tv'enicohr. thrcc spécimens ednsi-

dered as sy (Types in BMNH Reg. 197 81 32, front

Ticao, Philippines, thelargcsl figured liere (l'ig. 10C, D).

MATERIAL EXAMINEE), - The Idllowing lois in addi¬

tion to the type ntàterial:

Mauritius. (BMNH Winckworrh).

Madagascar. Stn 237 T ulcar (MNHN Thomassin). —

Nosi Bc (MHNG). — (USNM 718849).

Zanzibar. (MNHN Rousseau 1841).

Somalia. Mogadischio (ANSP 289942).

Djibouti. Gulf ol 1 adfoura (MNHN Gravier

1904). — Tadjoura (MNHN Lavranos 1968). W

ol Tadjoura (MNHN l avranos 1974).

Red Sea. (MHNG). — Elath (MNHN Tel Aviv

University). — Stn 27bis, 27ter, 28 U 14 > N, 33°23’E,

Gulf ol Suc/, 22 m (MNHN Mission Dolluss

1928). — Tor Reacb (MNHN Mission Dolluss

1928). — Mer,sa Thlemel (MNHN Mission Dolluss

1928). — Souakim (MNHN Jonsseaume 1921), — S

of Zeit. Egypt (fossil) (MNHN Plaziat). — Dahlak

Archipelago (BMNH).

Arabian Sea. Aden (BMNH). — Socorra Island

(MNHN lavranos 1968). — 19“22’26"N,

57°53’00"E, oiTMasirah Island, 13.3 m (NMW). —

19°22’36"N, 37°53 00"E off Masirah Island

(NMW). — Gulf Of Oman (BMNH Melvill). -

Makran Coast (BMNH Mae Andrew). — Karachi

(BMNH Mac Andrew).

Persian Gulf. Dubay'ah (MNHN Pras 1981). —

Trucial coast (BMNH). — Calypso 1934, stn 1, 2, k

of Trucial Coast, 70 ni (MNHN Charbonnier). —

Lavan Island (MNHN Eischer-Pîette 1973). — Abu

Dhabi (BMNH).

Maldives. (ANSP 105433).

Sri Lanka. (MNHN Reynaud 1829). — (MNHN

Denis 1945). - - (MNHN Staadt 1969). — (BMNH

Hoidswork). — (USNM 17446). — (USNM

203974).. — OSA^'N, 8l°07'E Nilavel (LACM

84-8). — (AMS C04093).

India. Alidallian Island (ZMUC Roeptorf 1875).

Thailand. Hai N an Beach, Phukei (MNHN

Vidal). — Surin Beach. Phtiket (MNHN Vidal). —

Phuket (ANSP 287283). — Songkhla, Gulf of

Tbadand (ANSP 2S7195).

Malaysia. Pulau Lembu, West Coast

(WAM 33-94). — 04°14'N, I03°27’E, N of

Kampong Kcinarnan, h.ast Coast (AMS C310564 and

ANSP 354005).

Singapore. Pulau Sudong réclamation

(AMS C310550, C310565, Cl 23696), — Senrosa

Island rcçlamatiou (AMS C310549).

Philippines. (ZMUC, 2 lots). — )olo (MNHN

Vidal). - Panglao (MNHN Vidal). — Cebu

(MNHN Vidal), — joie, 38 m (ZMUC Pacific

Expédition Moriensen 1914). — Harbour ol

Zamboanga, Mindanao (Vidal). - Pele 1964, Sibutu,

Sulu Archipelago, 23 m (WAM 878-66). —

Corregidor Island (AMS C310567), — Manda Bay

(ANSP 247060, 247055. 246714). — Siasi Island,

Suiu Archipelago (BPBM 203570). — Marongas

Island (USNM 484414).

Viet Nam. Qui N bon (MNHN Saurin). — Poulo

Cejar de Terre (MN| IN Saurirt).

china. (I ACM 50836). — 2|"07’N, I15°I5 ! E,

I 16-128 m (AMS C143045). — Ping Chan Island,

Hong Kong, 0-8 m (AMS C310599. Cl03243).

Japan. (LACM 13454,13507). (NMW

Melvill). — ANSP 228108). — (NI IMW, 2 lors). —

Shikoku ( M NI IN Staadt 1969). — I osa (MNHN

Staadt 1969). — Wakayaina 10-20 ni (MNHN

Vidal). Tosa (ANSP 209392). — Niï (ANSP

252677,). Ilirado Hi/cti (ANSP 80471). —

Kinu/aru, Awaji fANSP 141828). — Awaji (USNM

273676). — Hirado lliren (USNM 344805). —

Tosa (l ACM 13460). — 32°23'N, 130 D 07’E,

Amakusa, Kuamoto Prei., Kyusliu (LACM 82-25). —

32“3IN,.130°03'E, l amiuka Bay, Amasuka (LACM

82-23) — 34°IUN, I35°09 b,, Banshn /aki.

Wak a va ma pref., Honsfiu (I ACM 13460), —

32"49'N, 128 U 54’E, Nagasaki (BMNH).

indonesia. Makassar, Sulawesi (.ANSP). -- Moluccas

(BMNEI Cuming). — 06°26'S, )33°57 V L, Aru, 50 m

(AVAM 32941. — I.omhok (MNHN Vidal).

MARIEE K1NG Expédition 1970: stn AWl. Ü5°30’S,

134“12'E. Wasi. Island, Aru (USNM 747210,

'47248, WAM 42-94),

Papua New G-uinea. Enrôla Island, SW ol Porr

Moreshv 12-18 m (AMS C210567),

Queensland. Dingo Beach. Cape Gloncester

(AMS C077245). - ' 1 HAAS, I46"37'E, Créât Palm

Island (AMS E! 147234). — Off S end of louer Island

(AMS C310600) — HaVman Island (AMS

Cl47235). — Broven (AMS CQS9470). — Black

Island. t.anglorJ Red, Whitsundav (AMS C310576).

Norfolk Island and Lord Howe Rîse (Australia).

See in A. maculosum howense.

Chesterfield Plateau (New Caledonia). See in

A. maculosum howense.

Description

Shell medium-sizeti to snull. ovoid, ranging in

shape frotn nearly pcrfectly symmetrical to asym¬

metrical with a raised amerior dorsal margin and a

reccding posterior one, sometimes also more or

îess bi- imncaied (on PQ and Ml’Q). Dcspite this

asymmetry, shells always look roughly cquilateral,

and are never expanded backwards, witlt ribs

appearing sttaight in projection, Railler elongated

(mean E/H - 0.84; range 0.73-0.95) and modera-

telv inflated (mean W/l, = 0.71; range 0.59-0.85).

Lunule ver) nariow and liai on left valve, wider

on right valve, and slightly hollowed behind

302

ZOOSVSTEMA ■ 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 25. — Measurements (in mm) and rib count of Acrosterigma maculosum howense n. ssp.

W/L

A°

Ribs

Holotype howense

40.2

33.3

(22.0)

0.83

0.66

115

Paratype No. 1

36.8

30.5

(20.0)

0.83

0.66

0.93

Idem No. 2

36.6

29.0

(22.0)

0.79

0.81

Idem No. 3

38.0

30.2

21 1

0.79

0.70

=1.0

115

Idem No. 4

34.5

27.3

18.8

0.79

0.69

=1.0

120

Idem No. 5

51.1

40.4

(28.8)

0.79

0.71

0.90

Total adult shells measured and rib counts

General mean values

0.83

0.64

0.85

113

54.1

Standard déviation

0.03

0.04

0.08

4.6

Largest specimen observed. paratype No. 5 (see above)

exteriorly raised shell margin. Exterior almost

always strongly staincd with two sorts of shades

of brown, purplc or orange; large diffuse irregu-

lar ones and short darker more regular concentric

ones (mainly in young shclls). Interior white to

almost entirely purplc. Hinge symmetrical (mean

D close to 1-0), and ratber lirtle angled (< A

range 105°-I25°).

Rib number rather high, but variable according

to the particular form (see below), gencrally ran-

ging 43-61. Strong internai ribbing along mar¬

gin, very weak or not discernible on rest of shell.

Rib morphology: on PQ (Fig. 10E), smooth

anterior part of ribs about as widc as axial furrow

and posterior part, which are ofteu confused in a

depressed /.oue that usually bears irrcgular, main¬

ly tubercular, seules, ohen slightly encroaching

upon anterior part. Interstices narrow and very

thin. On MPQ, ribs asymmetrically slightly

rounded, with posterior flank steeper; tins llank

bears oblique, elongated svide tubertles which

encroach upon top zone, and produce festooning

ot posterior margin. 1 hese ridges stronger on last

ribs, in continuity with scales of PQ but more

numerous (Fig. 10E); interstices U-shaped, gene-

rally smooth but occasionally and locally not-

ched, independencly front flank sculpture. On

anterior hall of shell ribs appear as on MPQ or

become progressively top-ridged; fîrst ribs, until

the fourth or the fifth, progressively degenerate.

Distribution and remarks

(Fig. 11 ) Very schematically, ail rhe characters

described above are common to two morpholo¬

gie groups, with a rough correspondence to two

geographical distributions;

1) "Multinriiuunî form: (Fig. I0A, B) shells

rather small (30 mm in height), mainly symme¬

trical, with numerous ribs (50-61), umbo mode-

rately aaue (A range I 15-125°), shell smoother:

Red Sea. Per si a n Gu If, tropical east coast of

Africa, islands of the Indian Océan, Philippines,

soine localities in Indonesia. Shells front rhe Red

Sea are generallv smoother: scales on PQ are fre-

quently absent; retro-ridges in MPQ, and

top-ridges on the anterior balf are rare.

2) “Amiicola" form: (Fig. 10C, D, F) shells larger

(up to 51 mm), more often asymmetrical and

truncatcd, ribs less numerous (43-50), umbo more

acute (< A range 105°-I15°): N Indian Océan

(Srilanka, Andaman Sea), NW Pacific (China,

Japan), castern Indonesia (Moluccas, Papua New

Guinea), Australia (Central Queensland).

These two groups cannot constituie subspecics

beeause mahy exceptions and intermédiare lorms

exist, and a given individual is not necessarily

identifiable as bclanglng to one of the two groups.

The différences could be of cco-phcnotypic origin:

for cxample, large asymmetrical forms wîth a few

ribs in littoral waters, smaller more symmetrical

forms with more ribs in deeper water. However, a

subspecics with peculiar characters, can be des¬

cribed in the south-east corner of the range of

the species (see below).

Acrosterigma maculosum howense n. ssp.

(Fig. 10G, H; Table 25)

Types. — Holotype: a right valve (Fig. 10G, H), New

ZOOSYSTEMA • 1999 • 21 (2)

303

Vidal J.

Fig. 12. — Distribution of Acrosterigma impolitum.

Caledonia CORAIL 2 Campa ign 1988, stn DW 10,

20°52’S, 161 D 4l'E, Lansdowne- Fairwuy Shclf, 60 ni

(MNHN Richet Je Forges-ORSTÜM j. Paratypc I: a

left valve, saine data, l’ara type 2: a left valve, samc

data. Para type 3: a bivalved shell, 29 I, 02'S, 167°57’E,

Norfolk Island (AMS C05944G Bell de Iredale).

Paratype 4: a bivalved shell, same data. I’aratype 5: a

left valve. 31 n 33'S, !39°05'E, Lord Howe Island

(AMS Cl47228 Baxter).

EtymoI.OCtY. — From Lord Howe Ridge, between

Coral Sea and Tasman Sea. east of Australia.

M A II Kl Al KXAMINED. — The type specimens and the

lollowing lots:

Table 26. — Measurements (in mm) and rib count of Acrosterigma impolitum (Sowerby, 1833).

L/H

W/L

A°

Ribs

Lectotype impolitum

48.3

38.1

30.2

0.79

-1.0

120

Holotype beauforti

27.3

20.3

17.8

0.74

0.88

= 1.0

125

Paratype beauforti

23.3

17.8

14.1

0.76

0.79

Holotype dilmunense

43.1

35.7

26.8

0.83

0.75

1.05

125

Holotype couvriti

46.6

36.6

27.5

0.79

0.75

1.10

110

Holotype vtamtngi

27.4

24.2

(16.6)

0.88

0.69

0.96

Holotype rosemariensis

20.2

17.0

(11.8)

0.84

0.69

Holotype dampierense

39.3

32.4

(19.5)

0.82

0.60

= 1.0

105

Paratype dampierense

45.1

37.4

25.8

0.83

0.70

1.13

125

MNHN, Penang

43.8

35.0

25.7

0.80

0.73

0.95

105

MNHN, Singapore

32 5

27.4

19.0

0.84

0.69

1.10

120

MNHN, China

42.2

32.9

23.5

0.78

0.71

1.06

110

MNHN, NW Australia

21.5

18.4

13.0

0.86

0.71

1.10

125

MNHN, Queensland

16.0

15.3

9.6

0.96

0.63

0.96

125

QM 97-5707, Queensland

56.3

44.4

33.8

0.79

0.76

Total adult shells measured and rib counts

102

101

136

General mean values

0.85

0.70

1.04

118

39.5

Standard devialion

0.05

0.06

7.0

6.1

Largest specimen observed, QM 97-5707 (see above)

304

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cocldes

Australia. Norfolk Island (AMS C059446 Bell and

Iredale, AMS C14~222 Mort & Woolacort).

Tut 1962, Norfolk island: stn 62028, 28°54’S.

I67“59’F, 23 rn (MNZ M.224552). Stn 62029,

28°56'S, 167"58'E, 38 m_(MNZ M.224925). — Stn

62030, 28 0 59’S. 167°58'S, 38 rn IMNZ M.225104);

Lord Howc Island (AMS 0)29ISS, (.147228).

Kimbla 1976, Lord Howc- Rise: srn I Hl, 31"35'S,

159°00’E, 73 m (AMS C123975). — Stn LH5,

30“25’S, I S9"06 E. 50 m (AMS G123719). — Stn

I.H2, 31“38'S, I 59"04 L, 44 m (AMS CI24463),

New Calcdonia (MNHN). Coriatis MUSORSTOM 5

1986 Cotai Sea: stn 264, 25°20’S. 159°44’£, Banc

Capel, 56 m.

Chalcal 1984: stn 07, 20"51’S, 161°37’E, Banc

I.andsdowne-Fairway, 62 rn.

Coriolis CORAIL 2 1988, Bancs Landsdowne-

Fairway: stn DW 01, 20"56’S, 16T41E, 59 m. —

Stn DW 02, 20°50'S, I61°37'E, 62 m. — Stn DW

04, 20°52’S, 16l°37'f, 64 m. — Stn DW 10,

20°52'S, |6|°4rE, 60 m.

Distribution. — (Fig. 11) In New Caledonia, only

in Lansdowne-Fairway Bank and near Capel Bank,

both siruated on the Chesterlield Plateau. In

Australia, Lord Howe Island and Norfolk Island.

Description

As for Acrosterigma maculosum s.s. except for

three distinctive characrers:

1. Colour: shell exteriorly white. sontetimes wirh

vague lighr pink stains; interior gcncrally either

entirely pink or pink-striped, but always white in

umbonal cavity and on margins. Nymphal plate

always characteristically pink ( The nominal sub-

species is lyever pink, except for the interior of

some specimens from Queensland close to the

area of distribution of this subspecies),

2. In New Caledonia rib number higher than in

nominal subspecies, ranging from 53 to 61.

3. Rib morphology: rib.s lower and smoother

than in nominal subspecies, practically without

ornamentation (except on PQ and on First part

of AQ); on sortie shells First ribs ol AQ become

indistinct, forming a smooth zone beside lunule.

Acrosterigma impolitum (Sowerbv, 1833)

(Fig. 101-N; Table 26)

Cardium impolitum Sowerbv, 1833: fig, 6: 1841a:

107.

Cardium beauforti Prashad, 1932: 270, pl. 6,

figs 27; 28.

Fig. 13. — Distribution of Acrosterigma dampierense. A. rose-

mariensis and A. vlamingi.

loematrdium ( 7 rrtchycardium) couvrili Fischer-Piette,

1977: 57, pl. 5, fig. 2.

Acrosterigma vlamingi Wilson & Stevenson, 1977: 92,

pl, 6, figs 21; 22.

Aerosterigma memariensis Wilson & Stevenson, 1977:

95. pi. 6, figs 23; 24.

Acrosterigma dampierense Wilson & Stevenson, 1977;

98, pi. 6, figs 1-5.

Acrosterigma n. sp.(a) Oliver, 1995: 246, fig. 1089.

Tnt chy cardium impolitum dilmunensis Oliver &

Chesnev, 1997: 65. figs 25 ; 26; 29-36,

TYPES. — Cardium impolitum rhree shells acessioned

1991.042 in BMNH, Cuming collection, China, are

apparenrly syntypes. The largest (Fig. 101, J), certainly

the one figured by Reeve (1845: Sp. 80), is possibly

th.it mçasured ami figured by Sowerby (1840: fig. 66)

and was selected as lectotvpe by Voskuil &

Onvcrwagt (1991b: 119). Cardium beauforti'. two

shells in ZMA No. 314-4, Iront Siboga Expédition,

stn 133, Salibabu Island (between Philippines and

Moluccas), the holorype (Fig. 10K, L) sclccted and

figured by Prashad. A paratype from the santé area in

NMW, Melvill-Fomlin collection, 55.158.1245.

Laevicardium (Tracbycardium) couvrili : holotype in

ZOOSYSTEMA • 1999 • 21 (2)

305

Vidal J.

MNHN, ihc only reportcd specimen, svtkl to be front

off Gabon (West Africa). Acnisterigma ularningî ; holo-

type in WAM 1029-66, a single left valve Iront Shark

Bay, Western Australia. Eig.hr para types, ail Iront the

central west toast of Western Australia, front

Abrolhos Islande to Shark Bay. Acrosterigmct rosetnd-

riensis: holotype in WAM 650-66, a specinten from

Rosentary Island, Western Australia. T en paratypes,

ail from the northern enast of Western Australia, (Vont

Point Clones- to Dampier Atchipelaeo. Acrosterigma

dampierentf. holot)’pe in WAM 3334-68, a specimen

front Port Hcdland. Western Australia. hifteen pitra-

types, ail from the northcrnniost parr of Western

Australia, north front Dampier Archipelago (with one

exception). Northern lerritory, Queensland, and Arti

Island (Indonesia). Trachycanlium impolitum dilmu-

nensts : holotype B.VIN H 1994.100 front Trucial

Coast, Pcrsian Gulf. Tvvo paratypes, Biggs collection,

from Persian Gulf, one in BMNH, one in NMW.

Materiai, EXAMINE!». — The types of impolitum ,

beauforti, couvrtli, impolitum dilmunensr, one pararype

of rosemarietise . liamingi-, tsvci paratypes of dampieren-

se. The following additional lots:

Persian Gulf. Calypso 1954, E of Trucial Coast

(MNHN Charbonnier): stn 1. 54 Q OO N, 26°35’E,

35 m; sin 2, 70 m. — Bahram (BMNH). — Abu

Dhabi (BMNH 1944-100, 1944-101). —

26°N,51'E, Grand Mosque Beach, Jaffair, Bahrain

(NMW). 26-28’N. 50°05'E, Tarut Bay, Saudt

Arabia (AMS C310598).

Sri Lanka. (MNHN Staadt 1969).

India. Mandapam, grill of Manaar (ANSP

302324). — KarikaI (1RSNB).

Burma. S of Akyab, Bay ni Bengal (ANSP

239957). - Tavoy Island'(ANSP 292951). —

13°06’N, 96" 16'F., Tavoy Island (ANSP 292951).

Thailand. Phukc-t, fiai Nan Beach (MNHN

Vidal). — Phuker (ANSP 286323)- — Songka Beach,

E coast S Thaîland (AMS).

Malaysia, l’cnang (MNHN Vidal, IRSNB).

Singapore. (MNHN). — (BMNH Archer). —

(BMNH Wiuckworth). — (NMW Melvill). —

(USNM 128478). - (ANSP 5421 1). — Puiau Pasva

(ANSP 319360). — Puiau Sudong réclamation

(AMS 123696).

Cambodia. Ream (MNHN Fischer 1972).

Viet Nam. Qui Non (MNHN Saurin).

Philippines. (USNM 292297), Manila Bay

(ANSP 247226, 246659). Corregidor Island

(AMS). — Mindoro (USNM 617485). — Bentavan

(USNM 2931.35). — Balabac (USNM 237977). —

Jolo (USNM 235633, 235565, 235516. 23656.3). -

Jawî Jawi (USNM 283380, 236190). — Bacataan

(USNM 235971. Panay (USNM 248405). -

Iataan (USNM 236300). — Peltr 1964, N ot Subi

Archipelago, .37-40 tu (WAM 655-66). — Pt/e 1964,

S Eagoniy Sibutu, Sulu Archipelago, 16-27 m

(WAM 662-66).

China. (BMNH). — (MNHN Denis 1945). —

(MNHN Jousscaume 1921). —(USNM 120168).—

Hong Kong (AMS). — Hong Kong (ANSP

262933). — Hong Kong (7.MA von Hcukelom). —

Hong Kong (IRSNB Daur/cnberg). — Dongshan,

Fujian Province (MNHN Vidal). 22'‘33’N,

1 Ci 24'E, near Hong Kong. 0-8 m (AMS C103

243). — Tanin,n 1943, 2T17 N. ! I4 U 52'E, off Hong

Kong, 87-97 m (AMS C310540),

Indonesia. Siboga 1899-1900, stn M, Macassar

(ZMA), — Siboga 1899-1900, stn .311, Sumbawa

(ZMA). — liai m aéra, Moluccas (USNM

761777). — Saparoea Bay, Moluccas, 19 m

(ZMUC). — 5 D 30‘S, I43°12 E, Wastr Island, Aru,

30-40 m (USNM 747248). — 5°32'S, !32°4l’E > N

of Nuhu Rowa, Kai Island, 37 m (USNM

746926). —Japon Island, Irian Jaya (ANSP 206749).

Papua New Guinea. Manubada Island, Port Moresby

(AMS C310597).

Western Australia. Off Rottnci Island

(WAM 907-66). Port Hedland (QM). — La

Grange Bay (ANSP 325325).

Northern Territory. 12"09'S. 130°18’E. Charles

Point. W ol Darwin (WAV! 4-95). — 32 km off

Point Charles, Darwin (AMS C3I0512,

C060742). — Gove Pcninsula, NE of Arnhem

(AMS C.310568).

Queensland. AMS collections: Dingo Beach

(C084369). — Mapoon, Gulf of Caipemarîa

(COI418I). — Masthead Island, Capricorn Group

(C018857). — Albany Passage, Cape York

(CO36208). — Off Murray Island, Torres Srrait

(C036336). — Albany Passage (CO55693). —

Lindéman Island (058782). — Olf Burnet Heads

(G66185), — Big Sartdy Play, Swain Reef

(C066188) — lady Musgrave Island (C06618 7 ). —

2r42‘S, 1 52"26’E, .3 km NE ofW side of'Gillet Cay,

Swain Reef, 64-73 m (C12.3512). - Magnecic Island,

Townsville (C3 10541). — North West Island,

Capricorn Group (C.310543). — Eow lsles

(C.310569). — N ol Direction Island (C.310570). —

Bargara (C310571). — Eownsville (C310572). —

Witsunday Passage. 24 m (C310573). — llook

Island. Whitsunday Passage (C310574). — Hayman

Island ((2310575). — Black Island. Langford Reef,

Whitsunday (C310576). — Seaford. N of Mackav

(C310577). — Mackay (C3IÜ 578). - E of Mackav

(C.310 579). — E of Satina <C310580) — Hroad

Sound (C3I0581, C.3 10582). — Humpy Island,

Keppel Bay (C.310583), — Créât Keppel Island,

Keppel Bay (010584). — Noith West Island,

Capricorn fjroup (6.310585). — Yeppon (C.310586,

C.310587). — Hervey Bay (C310588, C.310590). —

OtT Monron Bay (C.310589). — Pialha. Hervey Bay

(( 510591). — Keppel Bay (C3I0592). — 6)11 Shaw

Island. N of Mackay (C310593). — Quoin Island,

Port Curtis (C'310594). — Tamnun Sands

(010595).

Lamprell collection (MNHN 1996): Gull of Carpen-

306

ZOOSYSTEMA • 1999 - 21 (2)

Taxonomie review of the clongated cockles

caria. — Nhelbourne. — Kurrimine Beach. — Dingo

Beach.— Rnrrnm Heads.— ülioal Point. — Turkey

Bay. — Woody Lsland;

Vidal Beach Survcy 1994, (MN1 IN): Airly Beach. —

Shoai Point, 15 km N of Mackay. — T'ammin Satids,

Canoë Point, SF, of Gladstone. Hervey Bay. —

Noosa Heads. Caloundra. Moreron Bay. —

Southporr.

Other muséums: Pialba, Hervey Bay (QM

52-002). — Peel lsland, Moreton Bay (QM 52-001,

97-5707). — Moreton Bay (BMNH Corning). —

Port Curtis (BMNH 1K81 -11-10-160-1 ). — Flinder’s

Entrance (BMNH 1881-11-10-162-3).

Distribution. — (Figs 12; 13).

Description

Shell medium-sized to small, subovoid, with

umbonal area more or less poinred. Young spéci¬

mens almosr perfectly symmctrical, then bccom-

ing slightly asymmetrical, with a recéding

posrerior dorsal margin and a raiscd and inflated

anterior one. Postcrior Iralf of PQ olten llatte-

ned, in relation to a straightened margin, form-

ing a roundcd angle with anterior part of PQ,

which can also be slightly truncated. Shells

always elongated (mean L/H = 0.85; range

0.76-0.95), but extern varies with populations

and individuals; a same variation is observed in

amount of compression (mean W/L = 0.70;

range 0.60-0.88).

Lunule narrow and fiat on left valve, slightly

wider on right valve. (Jnder a typical opaque

light yellow-brown periostraeum, regular over

entirc shell, ex te rival colour gencrally light, whitc

to yellow, sometimes with pttrplc mottling.

Inrerior whitc, sometimes slightly stained with

purple. Lfinge sometimes slightly asymmetrical

(average ratio D - 1.04; range 0.92-1.24) and

modérately augled (average < A = 118“; range

110°-130°), but thèse values seem to dépend

both oti population and shell size, large and

adttlt shells having a gcnerally smaller umbonal

angle and looking more poinred.

Mean rib number 39.5, range 29-52, possibly

reflecting an environ mental variation (see

below).

Rib morphology: on PQ, interstices always deve-

loped and often confused with hollowed post-

erior part and axial furrow; smooth anterior part

wider, with numerous small tubercular scales,

sometimes fused and reduced to a thin scar, or

occastonally absent. On médian part of shell

(MPQ and MAQ), ribs slightly rounded, trian-

gular and asymmetrical with posterior flank

steeper and shorter; posterior flank of ribs more

or less retro-ridged on different individuals;

interstices becomc notched with crescent-shaped

hollows, proceeding up posterior flanks, and

sometimes slightly up anterior flank as wcll,

a condition apparently relaced to the retro-

grooving of the ribs (Fig. ION). Sometimes a

longitudinal split euts hollows in rhe bottom of

interstice; this split can be double producing a

riblet. Sometimes ribs become rrapczoidal, with a

fiat top. In ail populations of Australia, and rare-

ly elsewhere (e.g. specimens of " beasijbttS' and

others from the Gulf of Matinal ), ribs of certain

individuals bear small coneentrically aligned gra¬

nules, gencrally on flanks but somelimcs on tops

or in interstices. On anrerior part ol shell, latéral

ridging progressively procceds up to top of ribs,

forming simple straight eross-bars. These

cross-bars are never imbricated, and may be rare

or absent. The tliree or four first ribs following

lunule always degenerate anteriorly, becoming

smooth, low and sometimes poorly marked.

Remarks

Acrosterigma impolitum is unusually variable in

rib number (29 to 52); it is also rather variable in

size. Very schematically, two extreme forms can

be observed: ( 1 ) large shells (30-45 mm hlgh or

more) with a few ribs (29-40), and (2) smaller

shells (20-30 mm high) with more ribs (4 1-52).

However, many transitional forms exisr and I

hâve been unablc to find signilicant breaks. Lite

hypothesis niade conccniing A. maeulantm is

probably appropriate here: large shells with fewer

ribs occurring in shallower zones are probably

ecotypes.

Acrosterigma impolitum can easily be separated

from the rclated species of the group by several

characters, the most importani being:

1. The “pointed’ look of the shell and the uni-

form thick light brown periostraeum; these cha¬

racters may not always lie definitive.

2. The regular, crescent-shaped notching of the

interstices (Fig. ION).

ZOOSYSTEMA • 1999 • 21 (2)

307

Vidal J.

3. The trapezoizal profile of the ribs Ln section.

4. The possible presence of local longitudinal

splits in the interstices, somedmes creating small

“riblets".

5. The rib-top cross-bars of tbe anterior part,

ofien disappearing anteriorly, but never degene-

rating into tubercles,

6. In Australia and occasionalJv elsewhere, the

presence ofaiigned small pustules on the ribs on

the médian part. Such pastules can be présent on

other species of the genus, but are smaller, gene-

rally less nurnerous and limited ro the juvénile

parts of the shells.

In spite of thèse nurnerous possible characters of

identification, several different nominal species

hâve been creatcd, winch I consider to be syno-

nyms of impolilum:

A. beauforti (Fig. IÜK, L), whîch differs only by a

slightly more elongated shape and the presence

of unusual reddish colours.

A. dampierense , rosetrwriensis and viamingi, from

Western Australia, hâve manv characters of irnpo -

litum, and, in my opinion, no character allowing

a spécifie séparation from it, nor among tltem-

selves. According to Wilson & Stevcnsons’s data,

these species are distributed in three different

areas (sec above the distribution of types and

Fig. 13), and are pracrically parapatric, with two

exceptions only. This suggesrs ecoryptc

influences, but the différences among these three

taxa are not sufficieutly constant to justify even a

subspeeifîc séparation. In Queensland, a compa¬

rable diversity of forms is présent, but without

apparent géographie or écologie ségrégation.

Such ecophenotypic variations are admirted in

Western Australia by Wilson & Stevenson (1977:

99), but for A. dampierense only: they separate:

(1) “fresh paired valves collected in shallow svater

or on beach"; (2) “smaller spécimens [ ta ken by

dredging in deeper vvater], more tumid with

slightly higher rib count rhan orhers (probablv

ecophenotypic variations...)”. On tbe other

Itand, Olivei & C.'hesncy (1997: 68), altliougb

they agrée with tbe synonyme ol dampierense

with impolitunn think that “wc could not accept

the amalgamated [Australien] cotnplex as a single

population", These autbors (197"’. 65) separate

also a subspccics impoli tum dilmunense in tbe

Persian Gulf, based on small différences on

shape, rib count and morphology, but above ail

on geographical isolation,

A. eotivrilr. tbe only specimen is a typîcal forra of

A . hnpolitum. It seerns that the only tesson for

this nevv species is an erroneous label iront West

Africa.

Acrosterignia transcendens

(Melvill & Standen, 1899)

(Fig. 14A-D; Table 27)

Cardium ( Iraehycardium) transeendens Melvill &c

Standen, 1899: 191, pb 11, fig. 21.

Card/tmi persiriatum Kuroda, 1928: 11 [ l'idc Habe,

1981: III].

Latvieardium pulcberrimum Sakurai &l Habe, 1966:

293.

I YlU-s. — Cardium transcende nr. bolorype, a left valve

Irom lorres Strait, BMNII 1899.23.6. l’ararype: a

specimen front sanie locality, Melvill coll. NMW

55.158.696, Cardium perstriatum : no types data.

Latvknrdium pulchtrrimuor. holotype and two para-

types, front Kakeroma-Iinia, Anianti lsland, in

National Science Muséum, |apan, ref. NSMT-MO

18405 and MO 70S79.

Ma i l 8IM. P SAMINI ti. — The following lots in addi¬

tion to the' type material of C. transcendens :

South Africa. N /ululant! from off Kosi Bay to off

Sodwana Bav (in Natal Muséum). Meinug Naudé

1987:. sm ZA9. 26”54.6’$. ,32°55.3’E, 50 m

(S48”4). Stn /.bl. 27 Ü 01.4"S. 32“54.2T.. 50 m

(D6836). Sm ZB6. 27 U 0I.I'S. 32"55.2F.. "8 m

(D7509). — Stn ZC3, 27 < ’06.5 S. 32°52.9'F., 70 m

(D6458). - Stn ZI 13, 27°32.8’S, 32°42.6'E, 68 m

(D6731). — Stn ZH4, 27 tt .33.2"S, 32*42.8’E. 85 m

D6772). Stn ZH16, 27 U 3S O S, 32 C '4I.8F., *0 m

(DS493).

NM DP 1990: stn ZA37- 26 M 54 OS. 32“55.5E, 50 m

(S3957). Stn ZA4S, 2f"53.5'S. 32°55.6'F.. 51 m

(S4039). — Stn ZA50, 2Û"55.0'S, 32"55.2’E. 41 m

(S7373). — Sm ZB19, 27°00."'S. 32'55.2'E. 70 m

(S4894). — Stn ZB22, 27°02.4 S, 32°54.9i:. 75 m

(S5369). - Sm ZCI0. 27”06.0'S, 32°53.3'S. 74 m

($6483). Sm ZOI2. 27”07.6'S. 32 u 52.4'f, "6 m

(88965). Stn ZD 10, 27' , |I.5'S, 32 # 50.4'F.. 78 ni

(S4635). - Stn ZH19, 27“'32.8'S. 32“42.8E, 77 ni

($4''771. - Stn ZH24, >7"32.2*S, 32‘’42.2'E,

49-53 m (s-T3 1).

Mozambique, (ialatheo 1951, stn 209, 20°08’S,

35"33T, olY Beira, 75 m (ZMUC). — ]omise. Boira

(USNM 718568).

La Réunion. Marion Dufresne 1982 (MNHN):

stn DC.4I., CHMZ and CP43, 21°21’S, 55°27’E,

73-77 m. — Stn DR47, 21°23'S, 55"37’E,

205-215 m. — Stn CP55, 21°05’S, 55°13’E,

308

ZOOSVSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

97-110 m. - Srn DC56, 2r()5’S, 55' J 12’E,

170-225 m. - Stn CP57, 2P05’S, 55°1I’E,

210-227 m. — Stn DC124, 20 ,, 52’S, 55"37’E, 40 m.

Mauritius. (MNHN Carfié 1911). — (MNHN

Arnould 1927).

Madagascar. Thomassin Survey 1962-1973. I ulear

area, MNHN': stn D3Ü, 23 n 23’S, 43°36’E,

70-75 m. — Stn 240, Grand Récif, 36 m. — Stn

621, Tulear Lagoon, S Pass.

Seychelles. Stn 1221. Amirante Island, 55 m, (BMNH

1910-8-31-7041.

Sri Lanka. N ol Trintomalee, 29 ni (AMS C310547).

Singapore. Sistcr Island réclamation (AMS

C310563). — Pu tau Island réclamation (AMS

310550). — Sentosa Island réclamation (AMS

C310548).

Philippines. Punta F.ngano, Mactan Island, Cebu,

80 ni (Hobbs), — Bentayan Island (USNM

293135), — Bentayan Island, 20-40 m (Hobbs)- —

Pelc 1964, off Mactan Island, Cebit (WAM 8-95). —

Bohol (Lamprell). — Pele 1964, SL of Balaba Island,

Palawan. 55 m (WAM 35-94). — Sibnga 1899-1900,

stn 98, 06“09’N, 120“21'F, Sulti Archipelago, 350 m

(/.MA).

Japan. Anianii O.shirna, Kyushu, 9-18 m (Hobbs). —

Onna Village, Okinawa (USNM 8385S4); 26°29'N.

127°50’F,{ (LACM 79-75); 26°29,6’N. 1 27°50.5’E,

(I.ACM 78-101). — Off Honian, Okinawa (BPRM

4337d), - 26"30,OO’N, I27“50,54’E (USNM

341664).

Indonesia. Bangka Island, Sumatra (Hobbs), —

8-48'S, 1 15" 14"F., Nusa Dua, Bali (LACM

86-166). — Aru Island (USNM 755486). — Aoeri

Island, Irian |ava (ANSP 208924, 205663). — E

Padaido Island. Irian Java. 45-90 m (ANSP 206104).

Siboga 1899-1900, (/.MA); stn 43, Polit Sarassa,

Postillon Island. — Stn 310, 873075. I I9°07.5’L,

Sumbawa, 73 ni. — Stn 315, Paiera osier Island. —

Stn 299, S of Rom Island. Timor. — Sut 240, Banda

Island.

Papua New Guinea. I Itinsa Bay, S of Madang, 45 m

(IRSNB 26132, 25955).

Queensland. Murray Island, Terres Strait

(AMS C029784, C030282, C310545). —Tories

Strait (BMNH 1937-7-9-29-30). - Off St Crispin

Reef. 40 ni (AMS C044684). — Kurrimine Beach

(7.MA Pïni 1977). — Swain Reef (AMS Cl23512).

Kimbla 1981 stn CIO, NE of Cairns, 55 ni

(AMS C310542).

Kimbla 19 7 7, stn 19. E of North Reef, Capncorn

Group (AMS t ’116523).

BRU ISM EXPEDITION 1928: stn 17, N of

Cooktown, 34,5 m (AMS C310561). — Low Isles

(AMS C.310557).

Western Australia. (29 km NNE of Port Hedland

(AMS C310555).

New Caledonia. Vaubau LAGON 1984, SW zone of

Lagoon (MNHN): stn 119, 22“28.0'S, I66°46 T1:.,

20 m. — Stn 130, 22°29.1’S, 166°48.3’E, 32 m. —

Stn 267. 22‘*21,5’S, 166°15'E 70 m; Vaubdn

LAGON 1984-85, S zone ol Lagoon (MNHN): stn

316, 22735.3'S, I 66°54.0’E, 68 ni. — Stn 392,

22**48.2'S, 167' I 02.3’E, 80 m. — Stn 398, 22737.0’S,

167**11.8 E, 71 m.

Vat.iban LAGON 1986, SE zone of Lagoon

(MNHN): stn 600, 22**|7.9’S, 167°04. i"E,

62-65 m. — Stn 624, 21“59.7'S, 166°52.0'E,

44-46 m. — Stn 626, 2l°57.94’S, 166°52.5'F,

47-48 in. — Stn 632, 2I°57.3'S, 166"49.6'E,

44-45 m. — Stn 6.33, 2I"55.6'S, ]66‘'48.2'F,

50 m. — Stn 658, 21 ”46.578, 166'’35.2 E,

49-51 m, — Stn 667, 21 o 42.07S, 166“27.7"E,

33- 3 7 m — Stn 688. 21 ”31,4’S, 166°15.2 E,

36-40 ni. — Stn 713, 2P22.67S, 166**00-7’E,

34- 35 m. — Stn 716, 21**22. US, I65"58.9’E,

30 m. — Stn 726, ’1**20.4'S, I65‘*55.0'E, 50-51 tn.

Van-ban I.AGON 198'-88, NE zone ol L.agoon

(MNHN); stn 748, 2I“I6.9’S, 1(^5' 5 49.9 E, 35 m. -—

Stn 761, 2P13.I5’S, 165“44.35’E, 41-44 m. —

Stn 782, 2r06.TS, 165736.7'E, 30 m. — Stn 789,

21°03.25’S, 165“33.55'E, 29 m. — Stn S0 7 ,

20 o 59.US, 165°28.75'E. 55 ni. — Stn 814,

2 I °55.5'S, 1 65°26.0’L, 38-30 m. — Stn 813,

21/54.1 'S, 165°26.95'E, 32 m. — Stn 816,

2T*52,6’S. Ki5“25.4'F, 31 m. — Stn 821, 20°51.9'S,

165 S 23.2*E, 32 m. — Stn 836. 2() a 46.4'S,

163<*15.'5’i:. 57 m. — Stn 83', 20**45.5*S,

165 3 l 3.9’F, 28-36 m. — Stn 900, 20°I4.6*S,

164723. TE, 40 m.

Cor/olis CHALCAL 1984, Lansdowne-Fairway Banks

(MNHN): stn DI 0, 20°36.09 , S, 161**05.82'E,

87 ni, — Cbcsierfield-Bcll'Otia Plateau (MNHN):

srn D46, 20°'32.26'S, ! 5 8 73.3,74'E, 65 m. —

Stn D4T 20*’50.85'S, 158**36.03’F, "0 m. —

Stn D50, 21°04.40'S, 158"40.70"E, 70 ni. —

Stn D51, 21 °13.21 'S. 1 58"42.50'E, 55 ni. —

Stn D52, 21°13.40'S, 158* , 49.20 E, 69 ni. —

S. n D5 5, 2 1 C 23.90'S, 158 C 59.60 E. 55 m. -

Stn D56, 2I°24.40’S, 159H)8.80'E. 60 ni. —

Stn D59, 2P40.367S, 1 59°2 1,29’E. 56 ni. —

Stn D61. 21 ”42.40 S. 159'’29.()()'E, 50 ni.

Coriutii CORAIL 2 1988, Landsdowne-Faitway

Banks (MNHN): stn DW 04, 20°52S, 16r37'E,

64 m, — Stn DW 08, 20**52 S, I6TT38T, 63 m. —

Stn I)W 21, 20"36’S, 16I°02’E, 86 m. — Stn DW 40,

19°29‘S, 158**3 5’E. 58 m; Conn/is CORAI1 2 1988,

Chesterfîcld Plateau (MNHN); stn DW 83, 19**12’S,

158°54'F. 59 m. — Stn DW 137, l'9734'S, 158°15’S,

32 m.

MONTROUZ1ER EXPEDITION 1993. Touho

area, (MNHN): srn 1261, 20°46’-20°4' r 'S,

165° l 54-l65“U)‘H, Chenal de Touho, 45-56 ni.

Al fi BATI IUS I 1993, east coast, (MNHN):

sm DW 678, 20°497S, 165”19’E, 94-100 m.

Vanuatu. (MNHN).

Wallis and Futuna. A lis MUSORSTOM 7 1992,

(MNHN): stn DW 538, 1273TS, 176"40’W,

Waterwitch Banc, 275-295 m.

ZOOSYSTEMA • 1999 • 21 (2)

309

Fig. 14. — A B, Acrosterigma transcendens, specïmen from Mauritius, MNHN; C. Acrosterigma transcendens, a left valve from

Chesterfield-Bellona Plateau, New-Caledonia. MNHN, D, Acrosterigma transcendens, a right valve Irom New-Caledonia Lagoon.

MNHN; detail o( anterior partshowlng concentrlcal leatures; E, F, Acrosterigma seurati, holotype; G, Acrosterigma dianthinum, syn-

type; H, I, Acrosterigma dianthinum. specimen from Passe de Koumac, New-Caledonia, MNHN; J, K, Acrosterigma punctolineatum,

specimen from Touho, New-Caledonia, MNHN; L, Vasticardium sewelli. specimen from New Caledonia, right valve, MNHN; detail of

rib morphology on the médian part of Shell. Scale bars: A-F, 10 mm; G-l, L, 5 mm; J, K, 20 mm.

ZOOSVSTEMA • 1999 • 21 (2)

VÊmS

vp*

Fig. 15. — Distribution of Acroslerigma transcendens and A. seurati.

Society Islands. Moorea (ANSP 250482). —

Opunohav Bay, Moorea (USNM 630645). — Bora

Bora (USNM 630006). — 16°42.8'S, 151°02’W

(LACM 74-37). — Raiatea (MNHN). — Tahiti

(ANSP 250348), — Tahiti (USNM 879714).

Tahiti. Taone Reef (USNM 671626). — Afaahiti,

Tahiti, 40-60 ni (MNHN Boutet).

Tuamotu Archipelago. (ANSP 53998, 53999). —

S Marutea (MNHN). — Rangiroa (USNM

789666). — Mataira (USNM 711626).

Tuvalu. 8°31’S, 179°13’E, Funaluti (AMS

C006184).

Marshall Islands. Bikini, 46-55 m (USNM 583037).

Distribution. — (Fig. 15) In addition to the loca¬

tions listed above, Acrosterigma transcendens is rccot-

Table 27. — Measurements (in mm) and rib count of Acrosterigma transcendens (Melvill & Standen, 1833).

L/H

W/L

A°

Ribs

Holotype transcendens

25.0

21.0

15.0

0.84

0.71

0.92

Paratype transcendens

26.8

23.1

16.3

0.86

0.71

MNHN. Mauritius

302

27.1

18.9

0.90

0.70

MNHN. Madagascar

24.7

22.0

16.4

0.89

0.75

0.95

125

AMS C310547, Sri Lanka

24.6

20.0

(15.2)

0.81

0.76

=1.0

AMS C310547, Singapore

22.7

19.2

(12.8)

0.85

0.67

120

Hobbs, Mactan Is

29.4

25.3

18.3

0.86

0.80

=1.0

MNHN, Japan

24.6

21.2

14.7

0.86

0.69

MNHN. New Caledoma

24.0

20.7

15.0

0.86

0.72

0.95

120

AMS C006184. Tuvalu Is

27.5

19.2

(12.8)

0.85

0.67

120

MNHN. Tahiti

26.3

22.3

17.5

0.85

0.78

=1.0

125

Total adult shells measured and rib counts

General mean values

0.87

0.73

0.98

123

64.4

Standard déviation

0.03

0.05

0.07

3.1

3.9

Largest specimen observed, MNHN, Mauritius (see above)

ZOOSYSTEMA • 1999 • 21 (2)

311

Vidal J.

Table 28. — Measurements (in mm) and rib count of Acrosterigma seurati n. sp.

L/H

W/L

A°

Ribs

Holotype seurati

43.6

37.9

(30.0)

0.87

0.79

125

Paratype No. 1

37.5

32.2

(25.4)

0.86

0.79

=1.0

Idem No. 2

40.9

35.5

(27.0)

0.87

0.76

130

Idem No. 3

37.6

33.0

(23.2)

0.88

0.70

=1.0

Idem No. 4

34.2

30.2

(22.6)

0.88

0.75

=1.0

Idem No. 5

32.2

28.5

(20.0)

0.89

0.70

130

Total adult shells measured and rib counts

General mean values

0.89

0.72

1.03

127

62.8

Standard déviation

Largest specimen observed, the holotype.

0.02

0.04

0.03

4.9

3.6

ded from Southern Honshu, Japan (Kuroda & Habe

1981: 1 11); it does noc live in littoral waters and is

only recoverable by dredging or deep diving, which

explains its relative rarirv. It is probably présent in the

main part of the tropical Indian Océan, and in many

Central Pacific archipelagoes.

DESCRIPTION

Shell medium to small, almost perfectly ovoid

and équilatéral (anterior dorsal margin somc-

times ver) slightly raised), and ne ver truncated;

also weakiy elongated (mean L/H = 0.87; range

0.82-0.93) and rat lier globose (mean

W/L = 0.73; range 0.66-0.86).

Lunule narrow, almost equal on both valves, and

poorly dilièrentiared, with iis umbonal margins

slightly raised. Extet'nal colour variable, olten

vivid and bright, liglit beige-yellovv variously

spotted and splaxhed with darker pink, orange,

or brown. Interior variable in colour, white to

pink or orange, with two umbonal rays which

are often widc and well-developed. Hinge nearlv

symmetrical (ratio D r. 1.0) and moderately

angled (< A range 120°-125°).

Mean rib nttmber 64.4, range 56-74.

Rib morphology: on PQ, interstices narrow and

both parts of ribs about équivalent in width, with

a variable axial furrow; scales irregular, mainly

tubercuiar, sometimes becoming very small, or

absent altogether. On posterior part ol MPQ, ribs

become mort or le.ss flatly triangular and asym¬

metrical (the anterior flank sborter and steeper),

sometimes still bearing an axial furrow; wider

posterior flank smooth or bearing elongated

ridges which are analogous to the scales ol PQ,

but more numerous. On anterior part of MPQ,

axial furrow and posterior long ridges eventually

disappear and ribs become progressivelv more

symmetrical and slightly rounded, ornamented

only with posterior crenulations; interstices beco¬

me wider and often very finely notched. On ante-

rior half of shell, ribs at ftrst remain rounded and

retro-crenulated, and then become somewhat

top-ridged; on ftrst ribs of AQ, top ridges become

tubercuiar and irregular, tendtng to be arranged

in concentric alignments which can replace the

longitudinal ribbing (Hg. 14D).

RhMARKS

This species is very close co A. maculosum, parti-

cularly as far as rib morphology is concerned.

A. transcendent can be separated Iront maculosum

bv a more vividly coloured, tbinner and often

smaller .shell, less elongated (mean L/II = 0.87

comparée! to 0.84), more globular (mean W/L =

0.73 compared to 0.71 ) and with more ribs (range

56-74 as opposed to 43-61 in maculosum).

A. trameendens is general ly rather consistent as

fàr as si/e, colours and tbickness of the shell are

concerned.

Acrosterigma seurati n. sp.

(Fig. 14E, F; Table 28)

Cardium maculosum Wood, 1815 - Lamy 1906; 214.

Cârdium unicaiot Sowvrhy. 18.34-Ranson 1967; 125.

Tirts. — Ail the existing material, nincreen single

valves, coilected by !.. G- Son rat, probably on h caches,

in ihe Marutea du Sud Atoll, Tuamotu Archipclago

(1902-1905), is selected as type sériés, stored in

MNHN. Holotype: a right valve (Fig. 14E, F).

312

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Table 29. — Measurements (in mm) and rib count of Acrosterigma dianthinum (Melvill & Standen, 1899).

L/H

W/L

A°

Ribs

Holotype dianthinum

14.0

12.0

8.0

0.86

0.67

M .0

130

MNHN, New Caledonia

12.3

10.6

8.0

0.86

0.75

1.05

120

Idem

10.0

9.0

6.2

0.90

0.67

=1.0

120

AMS C053697. Queensland

15.2

12.7

(8.6)

0.84

0.68

130

Idem

14.2

13.2

(8.4)

0.93

0.64

0.95

Idem

13.5

12.6

(9.0)

0.93

0.71

130

Total adult shells measured and rib counts

General mean values

0.89

0.69

1.00

126

46.1

Standard déviation

0.05

0.04

4.5

2.5

Largest specimen observed, AMS C053697 (see above)

Para types 1 to 18: icn riglit valves and eight left valves

(H = 40.9 to 25.2, L = 35.5 to 23.3, extrapolated

W = 27.0 to 16.2 mm).

Etymolocy. — In honour to Dr L. G. Seurat.

MaTERIAI EXAMINER AND DISTRIBUTION. — The type

material from Tuamotu archipelago, South Central

Pacific (Fig. 15),

Description

Shell medium-sized almost perfectly ovoid and

équilatéral (anterior dorsal margin sometimes

very slightly raised), rarely slightly rruncared pos-

rerîorly, weakly elongated (mean L/H = 0.89;

range 0.85-0.93) and rather globose (mean

W/L = 0.72; range 0.64-0.79),

Lunule narrow, almost equal on both valves, and

poorly differenriated, with its urrrbonal margins

slightly raised. External colour almost entirely

vvhite, exccpt foi a lew pink spots or small

splashes and part of PQ sometimes ycllow; inter-

ior white, sometimes with very pale yellow

umbonal rays. Hinge ncarly symmetrical (ratio D

c. 1.0) and variably angled (< A range 11 5°-

130°).

Mean rib number 62.8, range 56-68.

Rib morphology: on PQ, interstices very narrow

and both parts of ribs about équivalent in width;

scales irregular, tubercular. On posterior part of

MPQ, ribs become more or less Harlv roundcd,

sometimes still bearing an axial hirrow; entire rib

or wider posterior part can bear elongated ridges,

analogous to the scales of PQ, but more nume-

rous. On anterior part of MPQ ribs ornamenred

only with posterior crenulations; interstices beco¬

me wider and often very finely notched. On

anterior halfof shell, ribs at first remain rounded

and retro-crenulated, and then become finely

top-ridged; on first ribs of AQ, top ridges

become tubercular and irregular, tending to be

arranged in concentric alignaient* which can

replace the longitudinal ribbing.

Remarks

A. seuruti lias scveral characters in common w'ith

A. transcende)!s [1 treated it as a Tuamotu sub-

species of A. transcendent until 1 discovcred in

rliis archipelago several perfectly cypical lots of

the latter], but has a much larger, thickcr, heavier

shell and slightly less elongated shell (mean

L/LI - 0.89, rather than 0.87 in A. transcendent).

lt is also noticeably less colorful and the hinge is

a little less angled (mean < A = 127°, as compa-

red to 123° in A. transcendent).

Acrosterigma dianthinum

(Melvill & Standen, 1899)

(Fig. 14G-1; Table 29)

Cardiurn ( Tracbycardinm ) dianthinum Melvill &

Standen, 1899: 190, pl. I I, fig. 25A.

! Y!*FN. — Ail type séries from channel berween

Himimon Island and Wcduc-.day Splii Tornes Strait

(Austialia). Figurcd symype: a bivalvcd specimen in

BMNH, Rcg. 1899.2.23.11 (Fig. 14G). I hrce svn-

types: two bivalvcd shclls and onc righi valve in

NMW Reg. 1955.158.693. Five syntypes: three bivnl-

ved shells and two single valves in Manchester

Muséum (according to Trew 1987: 36).

ZOOSYSTEMA • 1999 • ZI (Z)

313

Vidal J.

MATERIAU EXAMJNKU AND DISTRI BU MON. — Five lots

in addition 10 the syntypes in BMNH and NMW

(Fig. 17).

Queensland. Eight small single valves, oft Murray

Island, lorres Strait (9°56’S, 144 < ’4'E), 9-15 ni

(AMS Ç310544, Hedley et ni coll., 1907). —

Nineteen single valves, Michaelmas Cay off Cairns,

16°36’S. 145"59'E (AMS C05.3697, Iredalc et ni

coll., 1926).

New Caledonia. MONTROUZ1ER EXPEDITION

1993, Koumac area, (MNHN): one bivalved spéci¬

men, stn 1.319. 20°44 S, 164°15.5’E, Passe Deverd

10-20 nv, one bivalved specimen (Fig. 14H, 1), stn

1310, 20 r '39.7’S, 164°14.9'E, Passe de Koumac,

15 m, Touho area, (MNHN), four bivalved spéci¬

mens, stn 1271,20°52,7’S, 165°19.5’E, Tié, 5-25 m.

Descri ki ion

Shell small, transversally ovoid in shape and

asymmetrical, but not posteriorly truncated,

élongation variable, but moderate (L/H range

0.84-0.98), and moderately inflared ÎW/L range

0.61-0.76).

Lunule small, slightly wider and hollowed on

right valve, and well-delineated. External co lotir

uniformly beige, only MPQ being colonred

brown-purple; interior slightly yellow in umbon-

al cavity and purple on MPQ (and sometimes

part of PQ) by transparency; nympli yellow; no

umbona! tavs observed. Hinge slightly asymme¬

trical (ratio D a little higher than 1.0) and mode¬

rately angled (< A c. 125°).

Mean rib number 46.1, range 42-50.

Rib morphology: on PQ, ribs liât, interstices

thin but well-marked; anterior smooth part of

ribs slightly narrower, and posterior part bearing

elongated, strong, regular and regularly disposer!

scales. On MPQ, ribs low, slightly rounded, rare-

ly somewhat flatly triangular, and smooth excepr

for presence on posterior third ol top of regularly

disposed small rounded tubercular pusiules. On

médian part of shell (MPQ and MAQ), ribs

beconre flatter and very low, are sometimes

slightly retro-crènulated, still bearing pustules.

Relatively wide interstices, ofren bearing regular

fine grooves in adult part, sometimes with a very'

thin, fairly clear, ribler on bottom. On AQ, pus¬

tules lengthen, and become progressively wider

and srronger top-ridges.

Remarks

Acrosterigma dianthinum is easily separated from

tire other small speeies of the genus both by its

characteristic shape and by the consistent rib-cop

pustules. The regular elongated scales of PQ and

the well-delineated lunule are châracters of the

species-group of A. vdriegatiun, but it is placed in

the A. maculosum species-group because of its

shape and low and poorly ornamented ribs.

Acrosterigma punctolineatum

Healv & Lamprell, 1992

(Figs 14J, K; 16A, B; Table 30)

Acrosterigma pnnctolineata Healy & Lamprell, 1992:

87, pl. 3, lîgs c-h.

Cardium foveolatum Sowerby — Reeve 1845: Sp 87

(Here Fig. 16,4, B),

l’robably not Ctirdium jvveohttum Sowerby, 1841a:

111 fwhich is unidentifiable],

Types, —Acrosterigma jmm tolineiU/t: hcilotypc: a nair-

ecl specimen Irom Finie Trunk Reef. N Queensland

(I8°20’.S, 146"46’E), in QM ref. MO32905.

Paratypes; four paired specimens from N Queensland,

in AMS.

Matériau examined. — The followittg lors:

Philippines. I3”46’N, 120“44'E, Batangas Province,

Luzon CW AM 661-66). — Tabango, Luzon

(AMS Cl04739), — 13°45'N, I20°46’E, Luzon

(WAM 663-66).

Indonesia. 1"28.4’N, 124“49.5 F, Menado, N of

Sulawesi (I.ACM 88-58). - Moluccas (ZMA

Huclit). — N coast of Ambon Island (WAM

41-94). 03"35'S, I28°02'L. Kg Said. N coast of

Ambon Island (WAM 12-95).

Papna New Guinea. Matlung (WAM 40-94). —

Fians.t Bay, S of Madung (IRSNB 26132). — Hansa

Bay, 10 m (IRSNB 26253). — New Britain

(AMS C45574).

Australia, Northern Territory. O rentes Reef, off

Port Fssingtort (WAM 42-95).

Queensland, lisard Island, Mrs Warson's Beach

(MNHN Vidal). — Ribhon Reef ( AMS Cl38337).

Australia, Lord Howe Island. (AMS C13~85).

New Caledonia. Poindimie, Monitel Hôtel Bcach

(MNHN Vidal 1992). — Poindimie (AMS

C87767). — Nchouè Bay, SI, of Poum (MNHN

Vidal 1992). — Alis LAGON 1989, N /une of

Ltgoon: sut DW 1195, I9°3Ô'S, 163°I9’E, 35-38 m

(MNHN).

MONTROUZIER EXPEDITION 1993, Fouho

area, (MNHN): stn 1271, 20°52.7'S, 165’19.5’E,

Haut Fond de fié, 5-25 m. — Stn 1272. 20 Ü 49.5’S,

165° 19, 6 F, Passe de Touho, 10 m. — Stn 1255,

20“43.0'S, 165°08.0’L, îlot Ouao, Il m. -

Stn 1246, 20°42.8'S, 165°08.7’E, îlot Ouao, 0 m.

314

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review ol the elongated cockles

Vanuatu. F.rakor Lagoon, Port Vila (MNHN Vidal

1989). — Efate Island (ANSP 787382). - Santos

Island (ANSP 787442).

Solomon Island. 9°3l’S, 160°48’E, Nuda Island

(LACM 78-69-42).

Distribution. — (Fig 17) Kccordcd only in the eas-

tern part of the Western Pacific, in littoral, reef envi¬

ronment. The species has beeo cited and fîgured trom

Taiwan by Kuroda (1941: 161. pl. 5. lig. 74) and from

Okinawa by Kubo fit Kitrozumi (1995: 181, lig. 3).

Description

Shell niedium-sizcd, ovoid, cquilaterally (only

anterior dorsal margin slightly raised) to trans¬

versale (MPQ niargin expanded) but not poste-

tiorly truncated, modcratcly elongated (niean

L/H = 0.86; range 0.80-0.93), and variably

compresscd (mean W/L = 0.73; range

0.66-0.85).

Lunule of medium size on right valve, with its

umbonal margin raised and encroaching on left

valve, whcre lunule i.s almosr non-existenr

External colotir beige with small, slightly elongat¬

ed brown spots, limited to smooth anterior part

of ribs on PQ, and to rib tops on other parts of

shell; on PQ thèse spots are longer and darker

than elsewhere. Inrerior shows the same features

by transparency. I linge symmetrieal (ratio D

c. 1,0) and moderately angled (< A I20°- 125°).

Mean rib nuinber 48.9, range 42-56.

Rib morphotogy: on PQ, ribs fiat with thin inter¬

stices; anterior smooth part of ribs narrower than

posterior part and terminating abrupdy against

interstice. Axial furrow relatively wide and readily

visible; oblique seules slightly elongated, régulât

in shape and rcgularly disposed in axial furrow,

encroaching upon posterior part of rib, but usual-

ly not upon anterior part. On médian part of

shell (MPQ and MÀQ), ribs roundly uiangular

co rounded, slightly asymmetrical, and gcnerally

smooth, cxcept for chose closest to PQ, which

sometimes bcar numerous long ridges on poste¬

rior face; on sortie young shells, small rounded

tubercular pustules randomly scattered on rib

tops; interstices narrower than ribs, bearing a

well-delineated small riblet in bottom, and fineiy

notched in places on adult parts. On AQ, ribs

become retro-crenulated, then top-ridged (wide

ridges, somewhat imbricated, becoming even

wider and irregular approaching lunule).

Remarks

Acrosterigma putictolineatum has typicaî charac-

ters (colourcd spots, riblets In interstices etc...)

and cannor be confused; ît is close to the ncxt

species A, hobbsae , but can easily be separated

from it (see below).

Acrosterigma hobbsae n. sp.

(Fig. 16C-H; Tables 31; 32)

Types. Hololype: a bivalved shell (Fig. 16C. D)

from tangle nets at 110 m, P un ta Hngano. Mactan

Island (Philippines), in MNHN, lee of Mrs Sue

Hobbs. Paratypc 1: a bivalved shell, same data.

Pararypc 2: a bivalsxd shell (iig. 16F, F), same locali-

ty, in Mis Sue Hobbs' private colI. Paratypc 3: a

bivalved shell Iront Molucca.s, in BMNH, Cuming

coll., Reg. 1996423. Paratypc 4: ,t bivalved shell,

slightly broken in both umbones, wirh soit parts, Aura

Harbor, Western Shoals, Cin.im. 2 m (’UGML, Paulay

coll.). Pararype 5: a left valve (Fig. I6G, H), santé

localicy and repository, Paratypc 6: a bivalved shell

slightly broken in ventral margin of right valve,

14 U 39 S, I45“29.5T, Macgillivrav Cav, Greac Rarrier

Réel, Queensland, 9-15 m (AMS C I 38397. Ian l.och

cl tl/.). Pa ta type 7: a loir valve, Montrouzier

Expédition 1993, New Caledonia: stn 1256,

2(J°45.0'8. l65 o 09.8’E, Vieux I ouho Lagoun,

15-20 ni. Patatype 8: a small bivalved specimen, from

CORAIL 2 Campaign 1988 New Caledonia: stn DW

117. 19°12’S, 158°36T, Chesterlleld Plateau, 60 m.

Etv.MOLOGY. — In honour to Mrs Sue Hobbs.

MaTERIAE EXAMINEE) AND DISTRIBUTION. — (Fig. 17)

The type sériés only; a rare species so fer fourni exclu-

sively in the eastern part of the Western Pacific, in

relatively "deep" warer (Philippines) as well as in lit¬

toral areas.

Description

Shell medium, generally ovoid and équilatéral,

rarely with a very slight oblique élongation and a

small expansion uf MPQ, but never posterutrly

truncated; weaklv elongated (niean L/H = 0.88;

range 0.84-0.95), and moderately inllated (mean

W/L - 0.72; range 0.68-0.82).

Lunule relatively wide and slightly hollowed on

right valve, with umbonal margin raised very

slightly or not at ail, narrower on left valve; to

sonie estent on borh valves, but tnainly on right,

posterior limir of lunule well-delineated, although

ribbing is degenerate near this limir (Fig. 16H).

External colour beige with fairly numerous irregu-

ZOOSYSTEMA • 1999 • 21 (2)

315

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«whii i w ibu.] i ii kî rsri T fl«fi[sî» •îilkMA i *!!k'C

Jmi T» 1» ; (3n»itïfcw$hïr»i[î):m$ *tjî*îii ir:îmu*iu■ ü ravM*f:ir;i»r*iil hW A1 Jl■ W VJuV* ?hi

ViIII»B*llK#T*[îî i Tr*»rt , TîT , [«[tk«l*BII*B ,, r«]l«]iI*][»T , l T M#lk'--i^h

Taxonomie review of the elongated cockles

Fig. 17. — Distribution of Acrosterigma dianthinum, A. punctolineatum and A. hobbsae.

lar, brown to purple splashes, more frequent on

posterior part and rarelv darker on PQ; both ribs

and interstices coloured. Interior shows saine

colours by transparency. Hingc asymmetrical

(rnean ratio D = 0.86; range 0.72-1.00), and

moderately angled (< A range 125°-130°).

Mean rib numbet 61.9, range 58-65.

Rib morphology: on PQ (Fig. I6G), interstices

very thin, but usualty distinct. Ribs fiat, not

abrupt anteriorly, both parts rather irregular and

variable in width, and axial furrow sometimes

indistinct. Scale.s small, not elongated, but irregu¬

lar in shape and si/e, vaguely spiniform or .sparuli-

form, and often connected by a small thin ridge.

On médian part of shell (MPQ and MAQ), ribs

low and Hat, with a small, abrupt posterior flank,

those on MPQ slightly more rounded; ribs

smooth, except onc to three ribs at posterior edge

of MPQ, vvhich can bear posteriorly a few fine

ridges; anrerior ribs of this zone, beside AQ, can

Table 30. — Measurements (in mm) and rib count of Acrosterigma punctolineatum Healy & Lamprell, 1992.

H L

Holotype punctolineatum

35.7

29.9

Type Reeve's foveolatum

32.1

26.5

ZMA. Moluccas

35.0

30.8

IRSNB Hansa Bay, PNG

29.5

25.7

MNHN, Queensland

31.2

25.9

MNHN, New Caledoma

34.9

28.0

MNHN, New Caledonia

23.1

18.4

MNHN, Vanuatu

34.0

31.7

Total adult shells measured and rib counts

General mean values

Standard déviation

Largest specimen cited, the holotype (see above)

ZOOSYSTEMA • 1999 • 21 (2)

L/H

W L

A”

Ribs

24.0

0.84

0.80

120

19.6

0.83

0.74

1.10

120

21.4

0.88

0.69

18.8

0.87

0.73

(19.0)

0.83

0.73

125

20.9

0.80

0.75

1.10

(15.6)

0.80

0.85

=1.0

(21.0)

0.93

0.66

=1.0

0.86

0.73

1.04

122

48.9

0.03

0.05

2.4

3.3

317

Vidal J.

Table 31. — Measurements (in mm) and rib count of Acrosterigma hobbsae n. sp.

L/H

W/L

A 0

Ribs

Holotype hobbsae

24.9

22.0

15.0

0.88

0.68

0.84

130

Paratype No. 1

25.0

23.7

15.5

0.95

0.65

0.86

130

Idem No. 2

40.2

36.0

24.4

0.90

0.68

0.72

130

Idem No. 3

32.3

29.0

20.1

0.90

0.69

0.81

130

Idem No. 4

45.4

38.0

31.0

0.84

0.82

0.88

125

Idem No. 5

39.4

33.0

(24 4)

0.84

0.74

=1.0

Idem No. 6

35.5

31.0

22.0

0.87

0.71

0.85

125

Idem No. 7

32.2

28.5

(19.0)

0.89

0.67

0.83

Idem No. 8

16.6

11.5

6.9

non mature

Total adult shells measured and rib counts

General mean values

0.88

0.72

0.86

128

62.1

Standard déviation

0.03

0.05

0.07

2.4

2.6

Largest specimen observed, the paratype No. 4 (see above)

become increasingly finely retro-crenulated. On

AQ, these crenulations change into straight or

curved, relatively thin, free top-ridges, First regu-

larly disposed, then lengthening and tending to

align concentrically (Fig. 16H).

Rf.marks

Acrosterigma hobbsae resembles A. punctolineatum

as far as general appearance, shape and colours

are concerned, but dUTers by eharaccers tabulated

in Table 32. The two species are sympatric at

Touho. New Caledonia (MNHN).

Acrosterigma simplex (Spengler, 1799)

(Fig. Kd-P; Table 33)

Cardium simplex Spengler, 1799: 17.

Cardium unicolor Sowcrhy. 1834; fig. 29; 1841a: 107.

Cardium nebulosum Reevc, 1845: Sp. 99.

Laevicardium soyeri Ftscher-Pietre, 1977: 19, pl. 1,

figs 4-7.

Types. — Cardium simplex: holotype, a paîrcd speci¬

men in ZMUC without locality, Spengler coll.

(Fig. 16K. L). Cardium unicolor: six paired syntypes

from Ticao (Philippines), in BMNH, Cuming coll.,

the largest of which Ih’ig. ]61, J) fus the .specimen

cited and flgurcd hy Sowerby, and was selected as

lectotvpc by Vosknil & Onverwagr (1991hr 116).

Cardium nebulosum: holorype. a paired specirnen from

the Maldives m BMNH 1900-2-13-22, ftgured by

Reevc. Laevicardium soyeri: holotype and ilirce para-

types, ail paittd spécimens, labellcd (incorrectly)

“Corsica", in MNHN.

MaTERIAI. 1.XAM1NED. —- The following lots in addi¬

tion to the type sériés:

Mozambique, Conducia Bay, S of Choca (Natal

Muséum 1 14189 and G3042 legs Grosch).

Mauritius. (1RSNB Dauizenherg).

Madagascar. (MNHN Powis 1840). — (MNHN

Tcvnr de Ravin 1853). - (MNHN Petit 1921). —

Amhatnloakj (MNJ1N Petit 1921). — Nosi Be

(MNHN Boivm 1853). — 5m 230. 255. Tulear

(MNHN Thomas*in 1969). - Ambatoloaka, SW

Table 32. — Character différences between Acrosterigma punctolineatum Healy & Lamprell, 1992 and A. hobbsae n. sp.

A. punctolineatum A. hobbsae

Lunule small. Hat, limit vague

Colour spots only on top or anterior part of ribs (PQ)

Rib number range 41-57

Anterior margin of ribs on PQ abrupt; scales large and

regular

Ribs on médian part high; rounded interstices notched,

w/a intermediary riblet

Top-ridges on AQ thin and long

Lunule larger, liollowed, clear limit

Colour pattern rtol so confined

Rib number range 58-65

On PQ, anterior margin not abrupt: scales small, irregu

lar

Ribs on médian part low, Hat; interstices smooth, no

intermediary riblet

Top-ridges on AQ wide and tubercular

318

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

Fig. 18. — Distribution of Acrosterigma simplex.

Nosî Bc (MNHN von Cusel 1986). — Ambariaonka,

SW Nosi Bc (MNHN von Cosd 1986).

Comores. (MNHN Jousseaumc 1921). — Mayotte

(MNHN Clarct 1874). — Mayotte (Natal Muséum

K2273 Roseoe).

Zanzibar. (MNHN Rousseau 1841). — (MNHN

Boivin 1853). — (ANSP 213976, 54225, 597142,

643452).

Kenya. Likoni, near Mombassa (MNHN Lavranos

1970). - Shimoni (MNHN Bemley-Buckle 1972).

Seychelles. (MNHN Boivin 1853). — (BMNH

Taylor). — Aldabra Atoll (ANSP 837717). —

Olhuveli blet, near Mahè (LACM 84-1).

Red Sea. (BMNH 1844-6-3-11).

Sri Lanka. (MNHN Denis 1945).

India. Ofi Madras (Natal Muséum K2029

Honkcr). — Port Blair, Antiaman Island (BMNH

Winckworth).

Table 33. — Measurements (in mm) and rib count of Acrosterigma simplex (Spengler, 1799).

L/H

W/L

A°

Ribs

Holotype simplex

32.7

29.3

23.6

0.90

0.81

=1.0

125

Lectotype unicolor

45.5

37.7

28.5

0.82

0.76

=1.0

115

Holotype nebulosum

27.2

25.2

17.6

0.93

0.70

=1.0

125

Holotype soyeri

423

35.7

27.9

0.84

0.78

= 1.0

MNHN. Madagascar

40.5

35.3

27.0

0.87

0.76

1.10

120

HNHN, Sri Lanka

32.8

27.9

(22.8)

0.85

0.82

120

Hobbs coll., Sumatra

20.8

18.8

14.2

0 90

0.76

Hobbs coll., Philippines

220

196

15.7

0.89

0.80

MNHN, Queensland

34.1

29.2

22.9

0.86

0.78

1.10

125

MNHN, New Caledonia

42.5

35.4

31.7

0.78

0.89

1.05

115

Hobbs coll.. Carolines

28.1

25.0

19.5

0.89

0.78

USNM 614342, Marshall

39.5

32.4

(27.6)

0.82

0.85

120

MNHN, New Caledonia

52.8

39.4

36.0

0.75

0.91

1.10

115

Total adult shells measured and rib counts

General mean values

0.85

0.80

1.07

119

47.6

Standard déviation

0.05

0.06

5.0

3.8

Largest specimen observed, MNHN, New Caledonia (see above)

ZOOSYSTEMA ■ 1999 • 21 (2)

319

Vidal J.

Thailand. Koh T.to (ANSP 419740).

Philippines. Caholoa, Polillo Group, Quezon

Province (LAC’M 89960). — Qff Buyong Beach,

Mactan Islam! (AMS C143113). — Olongo Reef,

Mactan Island (Hobbs). — Ticao (MNHN I.etellier

1949). — Ticao (IRSNB). — Davao (I.ACM

13362). — Tabaco Bay, Albany Province ( I ACM

89222). — Bubuan Island (ANSP 248415). — Sa

Cruz Island, Zaïnboanga (ANSP 248298). — Siasi

Island, Sulu Archipelago (BPBM 203571).

Palau. (.ANSP 2020311. - Malakal I larbour (ANSP

202781). — Babcltiiunp Island, Palau (ANSP

203340). — Palau (USNM Paulay Bl'AI. 14, BPAL

16, BPAL 17, BPAI , 52).

Japan. Awaji (BMNH Mac Andrew).

Indoncsia. Sumarra, Bengkalis Island (Hobbs). —

Waiara Maumere, Flores (Hobbs). — Moluccas

(ZMA, 2 lots). Moluccas (BMNH). — Biak

Island, Irian Java (ANSP 600635).

Papua New Guinea. Hansa Bay, S of Madang

(IRSNB 25681, 26132). — Louisiadcs Archipelago

(BMNH 1856-126-8 22). — Samarai Island (Galatea

stn L396, ZMUC).

Solomon. I vous Point, Florida Island (BPBM

198603).

Queensland. Uatnlcy Island, Torres Sirait (WAM);

(BMNH 1846-8-31-203). —Stephen Island, Torres

Strait (WAM). — 09'’09'S, 143*5 .VF., Bramble C.n,

Torres Suait (AMS 0)51330). — 16’15’S, 145*5017,

Opal Reef, N of Catrns (AMS C310558). — 16 U 46'S.

145*58’E, Green Island, olï C.iirns (AMS

C310559). — Two l.slc.s (MNHN Vidal 1994). —

Four Miles Beach (MNHN I.amprcll). — Lizard

Island (MNHN Vidal 1994). Mrs Watson’s

Beach, Lizard Island (Vidal 1994). - Lizard Island

(ANSP 704706, 352683); (LACM 79-53, 79-55,

79-57). — 2I°42’S, I52°26'F:. Gillet Gay, Swain

Reef. 64-73 m (AMS 0 23512). 23*32’S,

151°45’E, Masthead Island, Oapricorn Group,

31-37 m (AMS CQ21200).

New C.aledonia. MNHN various cnil.: (Marie

1872). (Lambert 1876). — (Crosse & Fischer

1974). - (ORS IOM). — (Laboure 1982). — Ile

aux Canards (Catula 1953). — Baie des Citrons,

Nouméa (Btrih.utlr 1994). — Anse Vata, Nouméa

(WAM 3567-65). — Baie de Kuio, Ile des Pins

(MNHN Vidal 1989). — Ncltoué Bay. near Poum

(MNHN Vidal 1992). — Kountac Harbour (MNHN

Vidal 1992). — Platier du Ouen Tort) (MNHN

Vidal 1989). Anse Vata, Noumca (MNHN Vidal

1992). —Grand Recil Sud (MNHN Laboutc 1982).

Vuubah LAGON I9S4, SW zone of Lagoon,

(MNHN): sen 4, 22"22.5 S, 166"2(L~T, 9 m. Stn

5, 22*24..VS, 166*22.01., 10 m. — Stn 7, 22“24.0’S,

166° 19-7’E, 14 m. - Stn 10, 22*19.9'$, 166 n 20.4’L,

15 ni. — Stn 21, 22*22.8'S, 166°23.4’K, 10 m. —

Stn 49, 22 n l 8.5’S, 166*13.S’L, 10 m. — Stn 50,

22*12.6 ! S, 166“12.2‘E, 12 m. — Stn 51, 22 t T4.7’S,

166°11.1’E, 10 m. — Stn 63, 22°26.0’S, 166°26.3’E,

20 m. — Stn 66, 22"27.5'S, J66°27.4’E, 15 m. —

Stn 95, 22*31.3’S, I66°32.8'E, 14 m. — Stn 99,

22"32.6'S, 166°34.6’E. 14 m. — Stn 127. 22*30.6’S,

1 66*45.9’K,

— Stn

128.

22*30.2’S

166*44.017,

- Stn

150.

22*30. rs

166*50.4 E,

— Stn

161,

22*34.4’S

166*38.4'E,

— Stn

163,

22*12.0’S

166*07.5'E,

ni.

— Stn

185,

22*04.8’S

I66°02.2 , E, 15 m. — Stn 251, 22°J 9.3’S, 166“25’F„

20 m. — Stn 253, 22°22.l'S, 166°23’E, 16 m. — Stn

281, 22*23.~'S, 166°24 E, 10 m. — Stn 284,

22°25.8'S, 166°25’E. 6 m.

Vaubâti I AGON 1984-85. S zone of Lagoon,

(MNHN): stn 293. 22"4l.5'S. 166°40.9’K, 20 m. -

Stn 294, 22"43.7'S. 166*41.8L, 21 m. — Stn 296,

22°40.6'8, 166*44.4'E, 26 ni. — Stn 312, 22°4l.9’S,

166*48.8’E. 26 m. — Stn 340, 22*47.7’S,

166*46.6’E. 27 m. — Stn 357. 22*29.8’S,

167*06.7’E, 77 m. — Stn 546, 22*53.3’S,

166°51.6’F, 33 m. — Stn 547 22“54.5‘S, I66“53.0’E,

29 m. Stn 554, 22"50.2’8, 3 66*53.5'E, 27 m. —

Stn 564, 22'’46.8’S, 166*56.017, 35 m. — Stn 592,

22°34.2’S, 167°22.0’E, 22 ni. —Stn 593, 22°33.4’S,

167*20.017, 25 m,

Vauban LAGON 1985, (MNHN): Huon Atoll,

stn 443, 18*00.05, 162*55. |’E, 40 m. — Surprise

Atoll, sm 448, I8°2I.5'S, 163°07.0’E, 30 m. —

Stn 452, 18"27.4'S, 163'12,3’E, 27 ni. Stn 455,

18*29.5'S, I63"07.9E, 40 m. — Stn 465, 18*22.l’S,

163’05.0'F, 45 m. — Stn 473, 18*24.2’S,

I63°03.3'E. 50 m.

Vauban LAGON 1987-88, NF. zone of Lagoon,

(MNHN): sut 855, 20°38.35 , S, 1 65*09.1 I F„

22 m. Stn 885. 20*26. ES, I64“42,15’E, 32 m.

Vititban I.AGQN 1988, NW zone ul Lagoon,

IMNLIN): stn 916, 20°55.5S, 164*28.317, 13 m. -

Stn 923, 20°48.~’S, I64°24.2 Î E, 9 m. -- Stn 936,

20*4()7’5, 164’16.4 L. 15 ni. - Stn 940. 20*38. l’S,

164*15.5’F., 10 m. — Stn 941, 20*38.9’S,

164° 1.3.3’E, 16 m. — Stn 984, 20*21.2’S,

163*56.4’E, 23 m, — Stn 1026, 20*04.6’S,

163*47.6’E, 29 m. — Stn 1060, 20" 14.3’S,

164*15.4’E, 13 ni.

A lis LAGON 1989, N zone ol Lagoon. — Stn 1063,

20*03'8, 163*47’F., 31 m. — Stn 1065. 19’58’S,

163*51 ’E, 28 m. — Stn 1084, I9“5I'S, I6,3"50’E,

35 m. Stn 1088, I9"46'S, 163*58,E, 23 m. —

Stn 1094, 19*54’S, 163°41'E, 26 ni. — Stn 1104,

19*42’$, 163*59,E, 22 tn. Stn 1105, 19*40’S,

163*5717. 25 m. — Stn 1118, I9°35'S, 163°52'E,

30 m. — Stn l 126, 19°33’S, 163°46’H, 41 nv —

Stn 1140, ! 9*24’S, 163*4417, U m — Stn 1145,

19*21 ’S, 163"45’E, 38 m. Stn 1146, 19"08’S,

163*31 ’E, 185 m. — Stn 1154, 19”09’S, I63°19’E,

40 m. Stn 1157, 19 M 10'S, I63°10’E, 48 m. —

Stn 1158, I9"10’S, I63"07’E, 48 m. — Stn 1168,

19*16’S, 163*09’E, 50 ni. — Stn 1169, 19*19'S,

163"1 LE, 47 m. — Stn 1174, I9 n 2l’S, 163"l4’E,

53 m. — Stn 1182, 19°27’S, 163°16’E, 48 m. —

320

ZOOSYSTEMA • 1999 • ZI (Z)

Taxonomie review ol the elongated cockles

Stn 1195, !9°30'S, 163°19'E, 38 m. — Stn 1205,

19°42’S, 163°26’E, 38 m. — Stn 1213, 19°50’S,

163 0 33’E, 32 m. — Stn 1217, !9“52 > S, 163“36’E,

30 m.

(Inriolis Corail 2 1988, Chesterfield Arol! (MNHN):

stn DW 51, 19"18.50’S, 1 58"36.55’E, 69 m. Stn

DW 148. 19"5-108'S, ! 58 A 27.12'E, 34 ni.

A lis MUSORSTOM V! 1989, Loyauté Island Ri.se,

(MNHN): stn DW 445, 20°20.56'S. 16(> n 07.83’E.

— Stn DW 436, 20"20.27'S, I66 Ü 07.49'E, 33 m.

A lu PLOUVE.A1 1992. Ouvea Lagoon (MNHN): stn

1219, 20°30'S. 166 U 28F., 15 ni.

Vidais Survcy. b caches of main island 1989-92,

(MNHN): Néhouê Bav. — Koumac Harbour. —

Ouen Toro reef fiat. — Kuto Bay, lie des Pins. —

Pointe aux Longs Cous. Nouméa. — Anse Kuenda,

Presqu’île Nou. — Baie des Citrons, Nouméa.

MONTROUZIER EXPEDITION 1993, Koumac

area (MNHN): stn 1277, 20 Q 34 , S. I64 P I6’E. Baie de

Ouanap, 0-2 m. - Stn 1282. 20°33’S, 164’13'B, îlot

Tangadiou, 0 m. — Stn 1283. 20°33.5'S, 164°I2’F„

îlot Magone, 0 m. — Stn 1286, 20 B 38’S, 164°17’E,

Plateau Karembé, 0 m. Stn 1292, 20"22.4'S,

I64“06.8'E, Pointe de Barbouillât, 0 m. — Stn I 301,

20“37-3’S, 164“15'F, Récit de l'Irifernei, 1-5 iti- —

Stn 1303, 2l)°37.7’S, 164" 16 F., Plateau Karcmbe,

0-8 m. — Stn 1304, 20“38.6'S, 164"13.2'H, Chenal

de l’Infernet, 12-15 m. — Stn 1306, 20"39.TS,

164“ 1 2.4’E, Chenal de l'Infernet, 11-13 m.

Stn 1307, 20"33.7'S, 164°I0.3'E, Passe du Baron,

12m. — Stn 1308, 20”40S, 164°15.2 E, Ilot Kendec,

15-20 m. — Stn 1309, 20°40.5S,164'M3.4'E. îlot

Kendec, 18 m. — Stn 1314, 20°39.8’5, I64°15.3’E,

Passe de Koumac, 30-63 m.

MONTROU71FR EXPEDITION 1993, Touhc.

area, (MNHN): stn 1242, 20°46.2’S, 165“I4.5’E, reef

fiat off Touho wharf, 0 m. — Stn 1246, 20°42.8'S,

165"08.7’E, 0 ni.

A lis BATHUS I 1993, easr coast (MNHN):

stn DW678, 20°49'S. 165°19'E, 94-100 m.

Kiribati. King's Mill Island (USNM 76105). —

Tarawa Atoll (MNHN Paulav BTAR1 ).

Caroline Islands. Ponapc Hôtel (Holihs). — Truck

Lagoon (Hobbs). — I lelen Réel (ANSP 208013).

Marshall Islands. 0.5 miles a If Bikini Island 27 m

(USNM 582996), — 15 miles S ol'Vena, Bikini

(ANSP 585230) — Uierilc (ANSP 615609) —

Caaranbira (ANSP 584859), — Fniwetok (ANSP

285297, l.ACM 65-31). Wotho (USNM

614342). — Over Eniwerok (USNM 542916). —

Taka (USNM 615569). — Ailuk (USNM

615256). — N end of Lijeron Island. Jaluit (USNM

660006). — W of Rongelap Island, 37 ni (USNM

585487).

Dis tribution. — (Fig. 18) Acrosterignm simplex lias a

large but patchy distribution in the Indo-Pacific; it

seems inexplicable absent from the north-western

Indian Océan and from the south-eastern west Pacific

(eastern Melancsia and Polynesia).

Description

Medium-sized; adult shells generally asymmetric¬

al on dorsal margin with a reccding posterior sidc

and a raised, inflated anrerior one, and with an

obtuse angle on dorsal margin at the level of

anterior laterals; ventral part also asymmetrical,

with an expansion of posterior side and ribs

generally curved backwards in projection.

Generally slightly truncared at PQ-MPQ limic; a

weak furrow sometimes séparâtes these two

zones, producing an obruse notch on ntargin.

Moderately but variably elongared (mean

L/H = 0.85; range 0.73-0.96), and rather inflat¬

ed (mean W/L - 0.80; range 0.70-0.91).

Lunule rather largo and fiat on lcft valve, very

large and apprectably depressed on right valve. An

additional thick dark brown laver of lamellat-ed

periostracum présent on PQ and part of MPQ of

adult shells, Exterîor of the young shells generally

marked with irregular concentric Jîght putple

stains; on adules, which generally remain uniform

pale white to yellow, these aligned stains can per-

sist on MPQ. Inreriorlv white to some-what

purple b) transparency; PQ always white. Hinge

slightly asymmetrical, with ratio D always higher

than 1.0 (mean D = 1.07; range 1.0- 1.18); angle

A rather small, in a range of I I0°-I25 u .

Mean rib number 47.6, range 42-59.

Rib morphology: on PQ (Fig. 16P), smooth

anterior pari of ribs fiat, with an abrupt anterior

flank, and much wider than posterior pai r, repre-

sented by a thin, low ridge between axial turrow

and interstice; diis ridge, in turn, bears a very

thin top lurrow in which very small irregular

scales oecur, often connected by a thin ridge.

These features présent only on juvénile part of

shell; on adult part (excepi for some of last ribs),

posterior ridge and scales disappear and posterior

part of ribs becomes a “pseudo-interstice”, with

or without an axial cicatrice; simultaneously

anterior part ot ribs lowers, shell surface beco-

ming almost smooth on most adult margin of

large shells. On médian part of shell (MPQ and

MAQ), therc are low ribs of slightly rounded

profile, at fîrst smooth beside PQ, then beco-

ming finely and regularly retro-ridged. Interstices

ZOOSYSTEMA • 1999 • 21 (2)

321

Vidal J.

rounded, smoorh, and half width of ribs. On

AQ, posterior rhin ridges of ribs disappear or

change inco scarce, irregular, very thin

top-ridges; ribs beside lunule bccome very low,

someiimes wirh somewhat wide, tubercular

top-ridges, sometimes entirely smooth and

almost indistinct on fully adult part of shell.

RliMARKS

Acrosterigma simplex can be separated from the

other species' of the species-group mainly bv its

large lunule, the lack of bright colours (though

exceptions cxisr). unique character ofPQ tib

morphology [particularly wirh ontogénie disap-

pearance ol scales and smoothing ol surface

(Fig. 16P) and the weak ornamentation of the

ribs elsewhere]. As discussed below, this species

shows several transitional characters between the

species-group of A. maculusum and the following

species-group of A. biradiatum.

Species-group of Acrosterigma biradiatum

(Bruguière, 1789)

Diagnosis. — See Table 5.

INCI.UDED SPECIES. — 1) Recent: A. biradiatum

(Bruguière, 1789); A. attenuatum (Sowerby, 1841).

2) Fossil: no data.

Remarks

These two species bave been refctred by ail other

authors to the genus Laevicardium , subfamily

Laevicardiinae Kcen, 1951 (Keen 1936 is inva-

lid). This subfamily has often been vievved as

represetuing a wide group cncompassing a num-

ber of gcncra and subgenera (Keen 1951; Thiele

1935; Adam 8e Lcloup 1939; Fischer-Piettc

1977; Schneider 1995). Kafanov (1975; 145;

1980: 299) reduced the number ol généra to

only Laevicardium and Fulvia. Fuma has been

removed from the group by Wilson & Stevenson

(1977). I hâve shown (Vidal 1994) that Fulvia

rnust be placed in the subfamily Cardiinae (tribe

Vepricardiini), so thaï Laevicardiinae now would

contain only the genus Laevicardium.

Even restricted in ch is way, this subfamily still

lacks homogeneity, as remarked by Wilson &

Stevenson (1977: 57): “Shell characters of species

assigned to Laevicardium s.l. are variable [...]

This variability of shell characters within

Laevicardium suggests that subdivision of

Laevicardium [....] may be necessarv".

The séparation from this genus of the

species-group of A. biradiatum consdcures one of

the subdivisions considcred necessarv by Wilson

& Stevenson. The two species of this species-

group bave mûrie tous characters typical of

Acrosterigma and are here included in thaï genus,

in agreement wirh Wilson & Stevenson who sta-

ted (197”: 57) that “in sonie (species assigned to

Laevicardium ] shell characters approach the shell

characters ol the Ttachycardiinae very closely”.

The characters shared wirh Acrosterigma are:

simîJar shape; similar hirige structure (short,

massive, and strongly augled); presence ol two

coloured, internai umbonal rays and a scerigma;

on PQ ribs bipartite, wuth scales in juvénile

shells; crenulare posterior margin and retro-

crenulated ribs on part of the anterior half; and,

in the siphonal area ol rhe soir parts, a similar

disposition of tentades bearing no ocular organs.

In addition to these factors, several peculiar cha¬

racters of this species-group alreadv appear, inore

or less developed, in sonie species of

Acrosterigma. In addition to general loweringand

smoothing of ribs, and increasing rib number,

these are:.

1. Progressive ontogénie smoothing of PQ,

concurrent with the création of “pseudo-

interstices” (particularly in A. simplex and some

orher species ot rhe species-group ol A. rnaculo-

sum).

2. Latéral lowering and disappearance ol ribs in

AQ, in two stages separated by “diachronous”

(not occurring on the saine rib during growth,

but on successive ribs) démarcation lines (parti-

cularly visible in A. simplex ; manv other species

show only graduai degenerarion. even disappea¬

rance of the First ribs).

3. Appearance of concentric arrangements in the

anterior part ot the shell (more or less marked in

rhe species-group of A. nutculosum).

In conclusion it appears that the two species of

the group in question here are very close ro

Acrosterigma, and must bc placed in this genus.

Nor hâve I any doubts that this species-group

phylogenetically dérivés from Acrosterigma and

322

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cookies

particularly (rom the species-group of A. maculo-

suma , A. simplex representing an intermédiare

stage of this évolution.

The genus Laevkardium remains represented in

the Indo-Pacific by only two species, Laevi-

cardium multipunctatum (Sowcrby, 1833) and

L. lobulatum (Deshayes, 1855).

Acrosterigma biradiaturn ( Bruguière, 1789)

(Figs 16Q, Rt I9A-F; Table .34)

Cardium biradiaturn Bruguière, 1789: 231.

Laevkardium ntbropicium II abc- & Kosuge, 1966b:

153, pl. 59, (Ig, 2.

Not Cardium serratmn Linné, 1758 |= Laevkardium

laevigatum (Linné) sensu CL ne h & Smith, 1944: 22].

Tyi'HS. — Cardium biradiaturn: not traced. Sotne

authors considcr dut a possible type specimen is a

shell front Sri Lanka, Ml ING 1085/60 Laraarck coll.

(Fig. I6Q, R); I herc select this shell as neotype of

C. biradiaturn. l.aevicardmm rubropictum: holotype

and rwo paraty'pes. Iront Zamboanga (Philippines), in

National Science Muséum (Tokyo).

MATERIAL EXAMINE!). — The following lots:

South Africa, N Zuluknd, front ofl Rosi Bay to off

Sodwana Bay (lu Natal Muséum);

Mrinng Naudr 1987: sut ZA2, 26"56.0S. 32°54.7T.,

50 m (D7304). — Stn ZA5, 26“54.7’S, 32°55.LL,

45-37 m (D6265). — Stn /.Ail. 26“55.3\S,

32°55.4’L, 50 m (D8757). — Sut ZA30, 26*^3.H'S,

32°55.5'E, 50 ni (D7917). Stn ZB2, 27*00.8*5,

32 D 54.3'E 50 m (D9173). - Stn ZB5, 27*00.0’S,

32°55.2'F., 70 m (D6419). — Stn ZH7, 27°32.5’S,

32°42.0'F., 48-58 m (D6557). — .Stn ZJ6, 27*42.7’S,

32°39.9'E, 50 m (F,#23),

N MDP 1990: ZB14, 27°Û0.8'S, 32“54.3’E, 51 m

(S6542). - Stn ZB26. 27*03.8’S, 32*53.411, 44 m

(S8951). — Stn ZI 122, 27*31.8’, 32*43-0’E, 70 m

(S3884).

Mauritius. (MNHN Vidal, rwo lors). — (MNHN

Cloué 1850). - (MNHN Charret 1874). -

(MNHN Arnould 1927). - (MNHN Carné 1941).

Madagascar. (MNFIN Pawis 1840). — Nosi Be

(MNHN Vidal). — Nosi Bc (MNHN von Cosel

1986). — Nosi Bc- (MNHN Thornassin).

Thomassin Survey 1962-1973. l ulear area (MNHN):

stn DI6, 23"29'36”S. 43*41 ’48"W, Tulear Lagoon,

13-17 m. — Stn 726,. 733, 737. 738. 822, S Pass,

Tulear Lagoon, 12 m.

Mozambique. (LACM 50865). Conducia Bay (Natal

Muséum H42I I, H42I2, H4213, H42I4, H4216

Groscb 1975).

Comores. (MNHN Jousseaume 1921). — Mayotte

(MNHN Joly 1923).

Seychelles. Rêves 2 1980 (MNHN): stn 4. 05°08'S.

56°35’E, 32 m. — Stn 5. 05"05'S, 56“24'F.. 33 m. —

Stn 7, 04*53'S, 56*0l’E. 57 m. — Sut 32, 04’23’S.

54°F6’F, 51 m. — Stn 37, 04°35 , S, 55°12’L,

65 m. — Stn 47. 04"03’S, 55°59’E, 50 m. — Stn 50.

03*55’S. 55“40*E, 45 m.

Zanzibar. (MNHN Rousseau 1841). — (ANSP

213623).

Somalia. Near Mogadisciu (MNHN lavranos

1969). — 14 km N ofMogadisdo (ANSP 2958.37).

Yenien. Socotra Islaud (MNHN Lavtanos 1967).

Red Sea, (LACM 13471).— El Eliath, Aquaba Gulf

(ZMUC).

Sri Lanka. (MNHN Staadt 1969). — (LACM

50879). — (ANSP 54174).

Maldivc Islands. (ANSP 305458).

Philippines. Luzon (MNHN). — Cape Calavite,

Mindoro (LACM 89958). — Burms Island (MNHN

Staadt 1969); (ANSP 54.310, 225865). — Benrayan

Isknd, Cebu (MNHN Vidal). Mindanao (ANSP

223705). Zamboanga (LACM 50862). — Sulu

Archipelagq (MNHN Vidal). — Sulu Archipclago

(ANSP 223681). — |olo Island. Sulu Archipclago

(MNHN, ZMUC). — Siasi Island, Sulu Archipclago

(LACM 89-913. BPBM 203567).

Japan. Oshima (LACM 61161). — Lookoos,

Kumejima Island, Okiniwa (ANSP 321631). —

Yagaslti Island, N of Nagao, Okinawa (LACM

282' t 5). — OU Hnmân, Okinawa (BPBM

10232a). — Ryukyti Isknd (ANSP 252733).

Palau. Koror Island (ANSP 202421).

1 ndonesia. Java, 08”30’-08 I, 35’S, U4°28’E.

70-150 m (ZMUC Mortensen 1929). — Aocrî

Island, Irian Java (ANSP 208880). — Lontbok

(MNHN Vidai). '

Papua New Guinca. Hansa Bay. S ol Madang, 45 m

(IRSNB 26132).

Australia. Off Darwin, Northern Territory (ANSP

219289).

New Calcdonia. Coriolis C11ALCAL 1984,

Ghesterfield Atoll. (MNHN): stn D)4, 19*26.9Q’S,

158*31.90E, 246 m. — Stn Dlû, 19*1 1.90'S,

158°57.00'L, 63-67 m. — Stn D17, 19*1 1.90’S,

158*55.80’É, 44 m. — Stn D26. 19*10.72’S,

1 58*34.95’H, 48 m. — Stn D28. 19*24.18’S.

158*31.40‘K, 51 m. — Stn D33. 19*44.80’S.

1 58*25.80'L. 205 m. — Stn D39. 20*28.90’S.

148*48.?0T„ 40 rn.

Coriolis CHALCAl. 1984, Bellona Reefs (MNHN):

stn Di 4, 2 ()Vi 6,O.VS, 158*33.73% 79 m. — Sm

D46, 20*52,26’S, 158*33.74’E. 63 m , — Stn D48,

20*46,25’.S, 1 58*4 1,64’E, 70 m. — Stn D49,

20*58.20’S, 1 58*39.OO’F., 48 m. - Stn D51,

21*13,2 I’S, 158*42.50’E, 55 m. — Stn D53,

21*19.50’S, 158*59.30’L, 60 m. — Stn D95,

21*2.3.90’S. ! 98"99.60’F-, 55 m. — Stn D99,

21*40.,56’S. !59"21.29’E, 56 m. — Stn D60,

21*48.65’S, 159°27.95’E, 45 m.

Vauban LAGON, (MNHN): SW zone of Lagoon

ZOOSYSTEMA • 1999 • 21 (2)

323

Fig. 19. — A, B, Acrosterigma biradiatum, a specimen from Mauritius, MNHN; C, D, Acrosterigma biradiatum, a specîmen trom Sri

Lanka, MNHN; E, Acrosterigma biradiatum, detail ot juvénile PO in a specimen trom Chestertield Plateau, New-Caledonîa, MNHN,

showing the scaled ribbing of Acrosterigma type; F, Acrosterigma biradiatum, detail oi AQ in a left valve from Mauritius. MNHN,

showing the variation and disappearance of ribbing; G, H, Acrosterigma attenuatum, a specimen from Chesterfield-Bellona Plateau,

New Caledonia, MNHN; I, Acrosterigma attenuatum, a specimen from west coast of Gulf of Thailand (Thaiiand), MNHN; detail of PQ,

showing particularly the thin remaïning two last ribs white coloured. Scale bars: A-D, G, H, 20 mm: E, F, 5 mm: 1,10 mm.

294 ZQOSYSTEMA • 1999 • 21 ( 2 )

Taxonomie review of the elongated cockles

1984, stn 290. 22°36.6’S, 166 o 45.0'F., 35 m. —

S zone of Lagoon 1985, stn 410, 22°45.9’S,

167°22.2’E, 35 m. — SE zone of Lagoon 1986,

stn 716, 21 “22.1'S, 165°58.9’E, 30 m.

Vaul/an LAGON 1985, Surprise Atoll (MNHN):

stn 465, 18°22.1 ’S. I63°05.0’E, 45 m. — Stn 473,

I8°24.2’S, 163 0 03.3 , E, 50 m, — Stn 474, 18 D 02.4’S.

I63°0I . 8 ’E, 52 m.

CoriolU CORAIL 2 1988, Eairway Bank (MNHN):

stn DW 04, 20"52.30'S, 161°36.56’E, 64 m.

Coriolis CORAIL 2 1988, Chesterfield Atoll

(MNHN): stn DW 31, 19°24.86’S, 158°45.03’E,

57 m. — Stn DW 32, 19“24.90’S, 1 58' J 48.75 , E,

55 m. — Stn DW 34, 19°21.62’S, 158"55-77’E,

47 m . _ Stn DW 41, 19 t '2l.52’S, 158“31.87\E,

52 m. - Stn DW 42, 19°21.53’S, 158°28.83'E,

45 m. — Stn DW 44, 19°21.82 , S, 158°22.95’E. —

Stn DW 48. 19°18.,30'S, 158 a 27.00 : E, 44 ni.

Stn DW 50, 19°!8.30'S, l58 n 33.57’E, 50 m. —

Stn DW 51, 19 U 18.50'S. 158°36.55’E. 69 m. —

Stn 54, 19 , ’18.57’S, 158°43.50'E, 71 m. —

Stn 59, !9“18.50’S, 15R“56.5Î’E, 50 m. — Stn 60,

19°14.98’S, 158"56.98'E, 45 m. — Stn 63,

19°15.15’S, 158°47.73’E, 71 m. — Stn 65,

Table 34. — Measurements (in mm) and rib count of Acrosterigma biradialum (Bruguière, 1789).

L/H

W/L

A°

Ribs

Neotype biradiatum

40.0

33.1

21.0

0.83

0.63

0.95

110

MNHN, Madagascar

45.4

34.2

24.9

0.75

0.73

«1.0

105

MNHN, Zanzibar

51.3

39.7

28.2

0.77

0.71

«1.0

110

MNHN, Sri Lanka

45.6

33.5

23.4

0.73

0.70

=1.0

100

MNHN, Philippines

42.7

34,3

22.4

0.80

0.65

«1,0

115

MNHN, Surprise Atoll

48.0

36.2

26.3

0.75

0.73

0.95

100

MNHN. Chesterfield Atoll

42.4

34.2

20.8

0.81

0.61

0.90

105

MNHN, Socotra Is

56.8

45.2

(30.0)

0.80

0.66

«1.0

Total adult shells measured and rib counts

General mean values

0.80

0.67

0.98

110

50.9

Standard déviation

0.05

0.04

0.06

4.5

3.9

Largest specimen observed, MNHN, Socotra Island (see above)

ZOOSYSTEMA • 1999 • 21 (2)

325

Vidal J.

19°15.Q0'S, 158°40,64’E, 62 m. — Stn DW 67,

19°14.92'.S, 158°36.94’E, 66 m. — Stn DW 71.

19° 15.37',S, 158°24.37’E, 55 m. — Stn DW 72.

19°15.30'S, 158°20.89’E, 32 m. — Stn DW 76,

19°12.25'S, 158°32.90’E, 53 m. — Stn DW 77,

19°12,0rS, I58°35.9&E, 60 m. — Stn DW 79,

19°11.55"S, 158"43.40’E, 58 m. — Sm DW 80,

19°11.98'S, 158“47,81’E, 66 m. — Stn DW 87,

19 D 06,14'.S, 158 <I 59.94'E, 31 m. — Stn DW 88 ,

19°05.98'S. 158 t, 55.85'L, 32 m. — Stn DW 93,

19°05.92'S. 158°53.00'E. 59 m. — Stn DW 94.

19°06.0Q'S. 158“50.00’E, 40 m. — Sm DW 96.

19°06.00'S. 158”41.92’E, 41 m. — Stn DW 102,

19°09.03’S, 158 C 29.99'E, 58 m. — Stn DW 103,

19 o 01.01'S, 158°31.94'E, 58 m. — Stn DW 104,

19°08.95 8, 158 D 35.o7'E, 49 m. — Stn DW 106,

19°09.00’S, 158°42.62 , E, 62 m. — Stn DW 110,

19°08.95’S, 158 C '55.S2’E, 40 m. — Stn DW 1 18,

19°25.06’S, 158°28.35'E, 52 m. — Stn DW 119,

19°25,00’S, 158°24.60’E, 56 m. — Stn DW 122,

19°28.17’S, 158°17.Ü6’E, 32 m. — Stn DW 126,

19°28.07'S, I58°2~.00'F., 38 m. — Stn DW 128,

19°27.89’S. I 58"30.44’E, 46 m. — Stn DW 133,

19°31.U)’S f 1 58“25.35'E, 45 m. — Stn DW 137.

19°34.00’S. 32 m. — Sm DW 143,

19°37.40\S, 1 58 D 25.16'E, 45 m. — Stn DW 144.

19”27.73’S, 15{rZ3.28’E, 50 m..— Stn DW 145,

19°37.00’S. 158"! 9.12’E, 54 ni. — Stn DW 146,

19°37.00’S, l58 a 16.28'E, 44 m. — Stn DW 147,

19°36.87’S, l58 a 1.3.52’E, 25 m. — Stn DW 154,

19°52.04’S, I58°26.50’E, 35 ni. Sm DW 158,

19°46.00’S, 1 58“16.50'F., 28 m. — Stn DW 160.

19°46.00'S, 158°23.00’E, 41 m. - Stn DW 164,

19°4l.48'S, 158*'I8.79'E, 58 m.

A lis LAGON, N zone 1989 (MN11N): stn 1157,

19°10’S, 163“10’E, 48 m. — Stn 1 168, 19 D l6’S,

163°09'£, 50 m.

Wallis and Futuna. Alis MUSORSTOM 7. 1992

MNHN: stn DW 538, 12"31'S, I76 Ü 40’W,

Waterwitch Bank, 276-295 m.

Distribution. — d'ig. 20) In addition to the loca¬

tions abovc, this spcdes bas bccn rccordecl from the

Seychelles, Ni cobar Island, Central Indoncsia (Siboga

campaignj, Queensland and north Western Australia

(Wilson & Stevenson, 1977: 60).

Description

Shell of medium size, roughly pear-shaped

(maximum length measuted rnuch doser to ven¬

tral titan dorsal margin), witli médian ventral

margin straightened, and aJmost équilatéral with

posterior dorsal margin often more receding;

MPQ margin sornetimes sligluly expanded and

PQ often slighdy truncated with an obtuse notch

in margin, Variably elongated (mean

L/H = 0.80; range 0.72-0.88), and somewhat

compressed (mean W/L = 0.67; range

0.61-0.75),

Lunule s,s. not delineated, because there are no

rtbs on the anterior-mosr part (sublunule);

umbonal margin rising slighdy on right valve.

External colour Variable, bejge-ydlow wirh fairly

numcrous irrcgulâr red-brown splashes (sorae-

titncs lacking); on l’Q there are sevcral radially

disposcd darker stains, often triangiil-ar and well-

deljneated. Hinge rather symmetrical (mean

ratio D = 0.98; range 0.85-1.09) and appreciably

angled (mean < A = 110°; range 100°-120°). A

thin sterigma in the umbonal cavity of some

shdls in alloue 10% of lots.

Mean rib number 50.9, range 46-61. On the

most anterior part, this number is based on

internai marginal marks ol ribbing, which is

cleatcr thnti the degenerated ribbing itself.

Rib morphology: on PQ, the chârâcteristic rib¬

bing features ut the genus are only discernible on

the juvénile part (Fig. 19E)i ver)' thin interstices;

fiat ribs divided into two parts roughly équiva¬

lent in width and separâted by a thin axial fur-

row; irrcgulâr, tubctcular, more or less prjsmatic

scales présent on the last ribs, becoming progres-

sively smaller. Présence of scales restrkeed to a

small part of juvénile shell, nvainly on the last

ribs (Fig. 19E); then the axial furrow and post¬

erior part of rib unité to form a wide

''pseudo-interstice" (sec équivalent proccss in

A. simplex). On MPQ, ribs very low, slighdy

rounded and smooth (excepi (or some ribs near

PQ which hâve a little fine posterior ridging);

interstices smooth. On MAQ, ribs beçonie pro¬

gressive!)' retro-ridged with numerous thin

oblique ridges regularly disposcd, causing crenu-

lations of posterior sidc of top zone. On AQ,

posterior half idemical to MPQ with retrô-

ridgetl ribs (Fig. 19F). Then a zone occurs of

three to ftve unornamented ribs that are al most

indistinct, but still well-marked on margin

(Fig. 19F); this is lollowed by a smoorh zone of

equal width, with neither ribs nor marginal-

inrernal marks, posteriody contused with lunule,

i.e., a sublu-nule (Fig. 19F).

In addition to features described above, anterior

and posterior parts of shell also with a rather irre-

gular concentric ornamentation.

326

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the eiongated cockies

Table 35. — Measurements (in mm) and rib count of Acrosterigma attenuatum (Sowerby. 1841 ).

L/H

W/L

A 0

Ribs

Lectotype attenuatum

77.1

50.2

35.0

0.65

0.70

MNHN, Mozambique

60.0

41.1

27.8

0.69

0.68

=1.0

MNHN, Madagascar

52.3

35 7

23.8

0.68

0.67

1.05

MNHN, Kenya

42.3

30.6

19.7

0.72

0.64

1.10

MNHN. India

74.4

48.8

34.9

0.66

0.72

1.10

MNHN, Thailand

52.3

36,5

22.5

0.70

0.62

=1.0

MNHN, New caledonia

82.5

50.0

33.6

0.61

0.67

0.91

MNHN, New caledonia

768

45,5

33.3

0.59

0.73

=1.0

MNHN, Philippines

87.6

55.7

37.6

0.64

0.68

1.07

Total adult shells measured and rib counts

General mean values

0.66

0.68

1.03

56.9

Standard déviation

0.05

0.04

0.06

4.5

3.9

Largest specimen observed, MNHN, Philippines (see above)

Re MARKS

Some authors thought that C. biradiutum is a

junior synonym of Cardium serrât uni Linné,

1758 [for example Clench & Smith (1944: 23)].

I hâve examiner! the synrypes of C. sermtwn in

Linné’s personai collection in London, three

single valves that belong undoubtedly to the

American Atlantic Laevicardium luevigatum

(Linné) sensu Clench & Smirh, 1944. The lar-

gest, a left valve [37.7 X 31.5 X (24.0)J, with

Linné’s hand marks 73 and 89 (numbers in rhe

1758 and 1767 listings) still discernible, is here

selected as lettotype of Cardium svnamm Linné.

As for Cardium lacvigatum , Linné left three syn-

types, rwo bivalved shells that are Fttîsra papyra-

eea (Bruguière, 1789) [see Vidal 1994: 99] and a

single valve ot Fulvia austmlis (Sowerby, 1834).

The largest of the two bivalved shells [46.2 X

46.5 X 31.1], hand niarked 88 by Linné (1767

listing) is here selected as lcctotype of Cardium

laevigatum Linné, senior synonym of Cardium

papyraceum Bruguière, 1789. I lie two above

Linné’s species hâve been misidentified by ail the

subséquent authors [except Hanley (1855: 51),

cited by Weinkauff (1867: 148) and Bucquoy et

al. (1892: 301)].

Acrosterigma attenuatum (Sowerby, 1841)

(Fig. 19G-I; Table 35)

Cardium attenuatum Sowerby, 1841a: 109.

Types. — Several bivalved shells, bearing data

“Ceylon, Zanzibar, Philippines, M. C.” (although

only “Ceylon” [Sri Lanka| is mentioned in Sowerby’s

description), are considérée! as syniypes (BMNH, Reg.

1971-26). A lectotype was selected by Wilson ht

Stevenson (1977: 57).

MATKKIAL LXAMINIT). — The following lots in addi¬

tion to rhe type specimens:

South Africa. N Zululand, from off Kosî Bay to off

Sodwana Bay (in Natal Muséum):

N MDP 1990: stn ZA56, 26°56.4 , S, 32°54.2’E, 35 m

(S7665). — Stn ZA57, 26°56.8S, 32°53.6’E, 29 m

(S8803). — Sm ZII24, 2^°32.2'S. 32°42.2’E,

49-53 m (S4722).

Madagascar. Nosi Be (MNHN Plaute 1968). —

Nosi Bc (ANSP 261742).

Vhotnassin Survey 1962-1973, Tttlear area (MNHN):

stn D36. 23"29’36’S, 43°41’48"W, Tulear l.agoon,

13-17 ni. — Stn DI7, 23"29'00”S, 43°4l 39"W,

Tulear Lagoon, 7-8 m. — Stn 630 735, 740. 741, S

Pass pf Tulear Lagoon, 12-13 m.

Mozambique. (MNHN Staadt 1969). — (ANSP

2475-20). (1 ACM 13483, 50850, 50851). — SW

Conduira Bay, N of Choca (Natal Muséum H4204

Grosch 1975),

Zanzibar. (ANSP 226416, 213419),

Kenya. Shirrtoni, 80 kni S of Mombassa, 18 m

(MNHN Bendey-Buckle 1972).

Red Sea. Aquaba Gulf (MNHN Dolfuss 1929, sm 39).

India. (MNHN Vidal).

Sri Lanka. (ANSP 54313).

Thailand. h coast ol pcninsula, Gulf of Thailand

(MNHN Vidal).

Philippines. (MNHN). — Cape Calavite, Mîndoro

(LACM 89958). — Zamboanga (LACM 50881 ).

Japan. ÜlTHoman, Okinawa (BPBM Thaanum).

Indonesia. Padaido Island, Irian Jaya (ANSP

ZOOSYSTEMA • 1999 • 21 (2)

327

Vidal J.

206098). — Aoeri Islsnd, Irian Jaya (ANSP 205664).

Papua New Guinea. Ilansa Bay, S of Madans’

(IRSNB 26)32).

Australia. Darwin, Northern Territorv (ANSP

219279).

New Caledonia. Vauban LAGON 1984, SW zone of

Lagoon (MN1IN): stn 80, 22°30.5’S, 166°27.7’E,

33 m. — Stn 113, 22*22.9'S, I66°48.3'E, 32 m. —

Stn 115. 22°25.2’S, 166"46.2’E, 26 ni.

Vauban LAGON 1984-85, S zone of Lagoon

(MNHN): stn 336, 22°4I.5’S, l66 a 5l.4’E, 26 m. —

Stn 339, 22°46.2'S, 166 t ’47.9 , F„ 26 m — Stn 346,

22°44.8’S, 166°5I.6'L, 40 m. — Sin 544, 22°50.8'S,

166°48.5’E, 25 m. - Stn 596, 22“31.0'S,

166°21,0’E, 35 ni.

Vauban LAGON 1986, SE zone of Lagoon

(MNHN): stn 626, 21 “57-9’S, 166°52.5’E. 48 m. —

Stn 633, 21°55.6’S. I66 0 48.2’E, 50 m. — Stn 724,

2I°19.7’S, 165 g 57.8'E, 37 m.

Vauban LAGON 1987, NE zone of Lagoon

(MNHN): stn 885, 20”26.1’S, 164°42.Î5’E,

32 m. — Stn 900, 20°14,6'S, l64' : ’23.rE, 40 m.

Vauban LAGON 1988, NW zone of Lagoon

(MNHN): stn 937, 20°39.5 S I64°15.4’E, 53 ni. —

Stn 989, 20°18.rs, 163 <, 57.I’E, 21 m. — Stn 990,

20°19.0’S, 163°55,3’E, 23 ni. — Stn 1008,

20°11,0'S, 163°53,4'E, 27m. — Stn 1009, 20°09.9'5,

163°55.1 ’E. 19 m. — Stn 1024, 20"05.5’S,

163°50.3’E, 26 m.

A Lis LAGON 1989, N zone of Lagoon (MNHN): stn

1068, 19“5?’S, 1Ê3°53’F., 26 ni. — Stn 1069,

19°59'S, I63°53'E, 30 m. — Stn 1085, 19°50 S.

163°53’E, 33 ni. — Stn 1105. 19"40’S, 163 U 57'F„

25 m. — Stn 1118. 19 r, 35AS. 163 l1 52'E, 30 m. -

Stn 1128, 19°31’S, 163°52'E. 26 m. — Stn 1138,

19°27’S, 163°47’E, 42 m, — Stn I 155, 19°09’S,

163°16 E, 48 m. — Stn 1168, 19"16 S. 163°09E.

50 m. — Stn 1181, 19°24’S, 163 U 15'E, 45 m. —

Stn 1190, 19“34’S, 163°31’E, 40 m. Stn 1205.

19°42’S, I63”26’F., 38 m. — Stn 1214, 19“50'S,

163°37’E, 29 m. Stn 1217, 19 n 52 S. 163“36'E,

30 m.

Coriolis CHALCAL 1984, Lansdowne-l airway Banks:

stn D02. 21 Q l4.4l'S, 162°1 6.27'E, 80-120 m. —

Stn D03, 21°l4.00'S. I62°!6.4Ü'E, 120-150 ni. —

Stn D07. 20°50.86‘S, 16l”36.99'E, 62 ni. —

Stn D10. 20°36.Ü9‘S, I 61 “05.82'F., 87 m. —

Stn D12, 20°31.33'S, 16r06.5l'E, 80 m.

Coriolis CHALCAL 1984, Llellona Réels, (MNHN):

stn D52, 21°13.40’S, 158°49.20'E, 69 m, — Stn

D55, 2l“23,90’S, 158°59,60'E, 55 m.

Coriolis CORAIL 2 1988, l.andsdowne Bank,

(MNHN): stn DW 19, 20 °h1.72’S, 161°00.17’E,

77 ni. — Stn DW 20, 20“38.97’S, lOrOl.Ol’E,

88 m. — Stn DW 21, 22"07.88'S, 16T01.75’E,

86 m, - Stn DW 22 20°32.89’S, 161°01.09’L,

88 m. Stn l)W 23, 20 t *30.60’S, 16H03.55'L,

80 m. — Stn DW 28, 22 n 07.88’S, 160*56.34’E,

78 m.

Coriolis CORAIL 2 1988, Chesterfield Atoll

(MNHN): stn DW 48, 19°18.30’S. 158°27.00’E,

44 m. — Stn DW 75. 19°12.00’S, 158°29.50’E,

65 m. — Stn DW 100, 19°05.99’S, I58“26.89’E,

40 m.

DlMRIKUTtDN. — (Fig. 10) No additional informa¬

tion in the literature.

Description

Shell medium-sized to (rarely) large; ovoid and

almnst équilatéral when juvénile, hecoming more

and more incquilateral with expansion of MPQ

and scrong straightening of entire PQ margin;

adules very elnngated (mean L/H - 0.66; range

0.58-0.77), and moderatdv inllated (mean

W/L - 0.68;. range 0.62-0.74) '

Lunule somecimcs marked on right valve by a nar-

row hollovved zone; umhonal margin raised and

slightly sparuliform. External colour generally

white and yellow, rarely purple, PQand AQheing

darker. Inrerior white, with yellow margin and

umhonal cavity. Hinge almost syrnmerrical (ratio

D slightly variable but generally greater than 1.0),

and strongly angled (mean < A = 88°; range

80 ü -95 u ). A v/c.ik sterigtna is présent in tire untbo-

nal cavity ofsnme shc-lls in about 40% of lots.

Mean rih number 56.9, range 52-65 (again, rib

counr based on internai marginal Yibbing”).

Rib morphology: on PQ, characieristic ribbing

features of genus only discernible on juvénile

parr: interstices very tlhn; fiat ribs divided into

two rougbly equal parts, separated by a thin axial

fijrrow; irregular, tubercular, more or less prisma-

tic scales présent on last ribs, hecoming progres¬

sive!)' smaller; presence of scales restricted to a

small part ot |uvenile shell, mainly on last ribs.

Three wide pseudo-interstices (see équivalent

process on A. simplex and A. biracliatum) are

already tormed on very juvénile part of shell,

separating the lasr ribs that tbereby become well-

marked; generally rwo (rarely three) ol these last

ribs remain prominent and always readily discern¬

ible in adult shclls (Fig. 191); otber more anterior

ribs of PQ less discernible but strongly marked

on margin which is always crenulate (Fig. 19G, I).

On the médian part of shell (MPQ and MAQ),

ribs, not marked on rhe external surface of shell,

but oberi discernible by colours and confirmcd

by internai marginal “ribbing”, which is visible at

328

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

some distance from margin. On post-erior part

of AQ, ribs become again apparent ver) occasio-

nally with posterior crenulations as on other spe-

des of genus, then disappear and are rcplaced bv

rather irregular longitudinal striation; tins disap-

pearance happens, on each rib, more and more

backwards as shell grows; as il is rather abrupt, it

forms an oblique, "diaohronous" démarcation

line. On anterior striated zone, vague ribbing ean

still be visible by cojouring, and snll marked on

internai margin; rhis internai marginal ribbing,

and associated eoloured ribbing, disappear even-

tually, always at the santé distance from “lunule”,

most anterior part of shell being totally sntooth

(Fig. 190).

In addition to features described above, anterior

and posterior parts of shell bear a rather irregular

concentric ornamentation.

Acknowledgenients

I am especially grateful to Philippe Bouchet,

Bernard Métivicr and Rudo von Cosel, in the

MNHN of Paris, (or cheir help in many ways. I

am also indebted to Bertrand Richer de Forges

(ORSTOM, Nouméa).

I thank the Idllowing pcople for allovving me to

visit muséums, for loans, informations, and

donations to the MNF1N; Bruce Hayward

(AIM, Auckland); lan Loch (AMS, Sydney);

Gary Rosenberg (ANSP, Philadelphia); Robert

Cowie (BPBM, llottolulu); Kathie Way

(BMNH, I.ondon); J. van Gocthem and Claude

Massin (TRSNB, Bruxelles); James McLean and

Lindsev Groves (LACM, Los Angeles); Yves

Finet (MNHG, Genève); Bruce Marshall (MNZ,

Wellington); Richard Kilburn (Natal Muséum,

Pietermaritzburg); Graham Oliver (NMW,

Cardiff); Kevin Lamprell and John Stanis’ic

(QJvf, Brisbane); Andeis Warén (Swedish

Muséum of Natural History, Stockholm); Harold

and F.mily Vokes (Tularte University, New

Orléans); Gustav Paulay (University of Guarn

Marine l.ahoratory, Mattgilao); Alan Kabat and

M. G. Harasevvych (USNM, Washington); Lars

Wallin (Uppsala University Zoological Muséum,

Uppsala); Frcd Wells and Shirley Slack-Smith

(WAM, Perth); Robert Moolcnbeek (ZMA,

Amsterdam); Godtlred Petersen and Tom

Schiotte (ZMUC, Copenhagcn).

Very spécial thanks for the friendship and help of

Mrs Sue Hobbs, of Cape May, New Jersey, USA,

who provided me müch information and mate-

rial from ber large collection, and made several

donations to MNHN. I am also indebted to

Mrs Elizabeth Ruggeri, for editing and correcting

the English of the manuscript.

INDEX OF SPECIFIC AND SUBSPECIFIC TAXA

(A.= Acrosterignia, F.- Fulviti , L.= Laevicardium, T.= Trachycardium ,

V.= Vastuardium , Vp. = Vepricardium)

abrolhense (A.) n. sp.284

alternatum ( 14) see angulatum .269

aminmte (A) n. sp...297

angulatum ( D .269

arenicoloides (A?) foss..272

arenicolum (A.) see mdculosum .299

asiatîcurn (Vp.) .264

assimile (V.) .269

attenudtum (Z.) . 327

australis (F.) .327

heauforti (A.) see impolitum .305

belcheri (T.) .264

biradiatum (A.) .323

burchardi (A.) .276

caparti (Vp.) . 264

cipangense ssp. of elongatum (V) .268

compuncnm ( If) see mbicundum .269

consors (X) . 264

condense .ssp. of élongation (V) .268

couvrili (Z.) sce impolitum ...305

cygnoruin (A.) . 279

dalli (.4.) foss... 272

dampierense (A.) see impolitum .305

déclive (A.) foss.272

delcçtabile (A.) foss. .. 277

delicatulum ( Vpi) . 267

denticostulatum ( 17) foss... 270

dianthinum (A.) .313

ZOOSYSTEMA • 1999 • 21 (2)

329

Vidal J.

dilmuneme ssp. of impolitum (A.) .305

disais (,40 n. sp.290

dominicense (77) foss.264

dupucbense ssp. oïflavum (17) .269

egmontianum (77) . 264

elongatum (VT) . 268

mode ssp. of elongatum (VT).268

fidele ( V0 .268

fimbriatum (ty.) .267

flavum (VO ......269

foveolatum (A ) see tygnorum .279

fraternum ( Aî) foss. ......272

fucatum ( VO sce flavum .........269

fultoni ( 17) sec ornatttm ..269

gaillard! (/I. ) sec tygnorum .279

gortanii ( 17) foss. ...268

gratiosum (VO s ce flavum ...269

hadratatum ssp. o ï dominicense ( 77) foss.264

hawaiensis ssp. of orbita ( VT) ... 268

hobbsae (A.) n. sp. 315

boerleorum (A.) foss...272

bornelli (A) sce pristipleura ..272

boivense n. ssp. o ï maçulosum (VT) ..303

impolitum (A.) . 305

inconspkuutti [A.) foss... 272

indioceanum ssp. of elongatum (VT) ..268

insulare ssp. of luteomarginatum ( Vf) .268

isocardia (77). 264

kantoense (Ai) foss. .. 272

kengaluorum (VO see rubicundum ..269

kerslakae (A.) ...281

lacunosum ssp. of assimile ( VT) ..269

biddi ( 17) foss. sce sewelli .....269

laevigatum (L.) . 327

leucostoma (A.) sec magnum .....274

linguatigris (Æ) foss. 272

lobulation ( L.) . 327

luteomarginatum ( Vf) . 268

maculation {A.) see pristipleura . 272

maçulosum ( A. ) Sow. see pristipleura .272

maçulosum (A.) Wood ..299

magnum (A) . 275

manueli ( 77) . 264

marerubnon ssp. o (luteomarginatum (17) .... 268

marielae (Æ) . 283

marmoreum (À.) see magnum . 275

mauritiamtm (A.) . 291

mendanaense ( Vf) ssp. of orbita (Vf) .268

mindanense ( Vf) see rubicundum .269

multicostation ( Vp. ) foss .264

multipunctatum ( L .) . 327

mulustriatum (A) see maçulosum .299

muricatum ( 77) . 264

nclntlosum (A. ) see simplex .318

nigrapunctatum ( VT) see pectiniforme .269

okinawaense (Vf) see élongation .268

olivifer ( 17) see vertebratum .269

orbita (17) .268

ormuum (VT) ...-.269

otukaî (Ai) foss.272

axygontm (/!.). 288

pana merise (77) . 264

papuanum (V.) . 268

papyraeea (F.) . 327

paulayi (A.) n. sp. foss...299

pectiniforme ( 17) . 269

peregrinum ( V!*) sec pectiniforme .269

perstriatum (/I.) see transcendens . 308

plnlippinense ssp. of orbita ( Vf) ...268

praeeygnorum (A.) foss. 277

pristipleura (.4.). 272

procerum ( 7.) . 264

prveet'dens (Ai) foss. 272

profundum (A.) n. sp.......296

protosu b rtigostcm ( Vi) foss...270

pseudoanguhnum (VT) see orbita .. 268

pulchenimum (A.) see transcendens .308

punctolinentum (/t) ..314

quadragenariim (T) .. 264

reeveanum ( Vf) see vertebratum ..269

régulant ( 17) see pectiniforme ...269

rhegminurn ( Vf). 269

rosemariense (A.) sec impolitum .. 305

rubicundum (V.) . 269

rubropicntm (A.) see biradiatum .. 323

rugosum ( VO see pectiniforme . 269

sanguincotinctum (A.) see oxygonum . 288

sedanense ( V?) foss.270

selene (A.) n. sp. 289

senticosum ( 77) . 264

serrât uni (77.). 327

serricastdtum ( VT) see elongatum .268

serrulatum (Vp.) .264

seurati (Æ) n. sp. 312

sewelli (V.) .269

simplex (A ) .318

sorenseni (A.) .280

sowerbyorum (17) see ornatum .269

330

ZOOSYSTEMA • 1999 • 21 (2)

Taxonomie review of the elongated cockles

soyeri (A ) see simplex .318

spolongettse ( V?) foss.270

subalternatum (V?) foss.270

subelongatmn (A.) s cl- magnum .274

subrugosum ssp. ot flavum (Vf) .269

suluanum (A.) n. sp.298

talahabense ( V?) foss.270

thomassini (Vf) .269

tokyoense (AF) foss.272

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Submitted on 14 May 1998;

accepted on 6 November 1998.

ZOOSYSTEMA • 1999 • 21 (2)

335

Confirmation de la validité du genre Hormiops

Fage, 1933 avec redescription d ’Hormiops

davidovi Fage, 1933 (Scorpiones, Ischnuridae)

Wilson R. LOURENÇO

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

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

arachne@mnhn.fr

Lionel MONOD

Muséum d Histoire naturelle, Route de Malagnou 1,

case postale 6434,

CH-1211 Genève 6 (Suisse)

Lourenço W. R. & Monod L. 1999. — Confirmation de la validité du genre Hormiops Fage,

1933 avec redescription d Hormiops davidovi Fage, 1933 (Scorpiones. Ischnuridae).

Zoosystema 21 (2) : 337-344.

MOTS CLÉS

Scorpion,

Hormiops ,

Ischnuridae,

redescription,

Poulo-Condore,

Vietnam.

RÉSUMÉ

L’étude des types ainsi que des quelques exemplaires de Scorpions apparte¬

nant au genre Hormiops Fage, 1933, déposés au Muséum national d’Histoire

naturelle à Paris, autorise la revalidation de ce genre, avec la proposition

d’une redescription pour Hormiops davidovi Fage, 1933. Cette espèce est

connue uniquement de l’île de Poulo-Condore, dans la région sud du

Vietnam. Les caractères différentiels sont la présence de deux yeux latéraux,

la trichobothrie db de la pince sur la face externe, un hémispermatophore

très simple avec des crochets réduits.

Kl. Y WORDS

Scorpion,

Hormiops,

Ischnuridae,

redescription,

Poulo-Condore,

Vietnam.

ABSTRACT

Confirmation of tbe validity of the genus Hormiops Fage, 1933 with redescrip¬

tion «/Hormiops davidovi Fage, 1933 (Scorpiones, Ischnuridae).

Study of the types and specimens of scorpions bclonging to the genus

Hormiops Fage, 1933. deposited in the Muséum national d'FIistoire natu¬

relle, Paris, leads to the revalidation ot this genus, with a redescription of

Hormiops davidovi Fage, 1933. fhîs species is known only from the Island of

Poulo-Condore in the south région of Vietnam. Differenrial characters are

presence of two latéral eyes, a trichobothriurn db of cltcla on the external sur¬

face, and an hemispermatophore very simple, with reduced hooks.

ZOOSYSTEMA • 1999 • 21 (2)

337

Lourenço W. R. & Monod L.

INTRODUCTION

Dans une note consacrée aux scorpions collectés

par M. C. Dawydoff en Indochine française,

Fage (1933) propose la création d’un nouveau

genre, Hormiops appartenant à la famille des

Ischnuridae et voisin du genre Hormurus Thorell,

1876. Dans sa diagnose du nouveau genre, Fage

propose les caractères distinctifs suivants entre les

genres Ifurmiops et Hormurus : (1) Présence de

deux yeux latéraux au lieu de trois comme chez

Hormurus ; (2) absence de carènes ventrales aux

segments 111 et IV' du metasoma ; ces carènes

étant présentes chez Hormurus. Fin outre, Fage

analyse la régression oculaire observée chez

Hormiops et l'associe à celle observée chez les

Heteroscorpion Birttla, 1903 de Madagascar, t e

genre Heteroscorpion , au départ placé clans la

famille des Ischnuridae est désormais classé

comme une famille à part, celle des Hetcro-

scorpionidae Kraepelin, 1905 (Lourenço 1996).

Après la description du genre Hormiops par Fage

(1933), ce genre n’a été le sujet d'aucune révision

ou réanalyse. Ceci tien t à deux, voire à trois fac¬

teurs : (1) les spécimens disponibles en collection

sont extrêmement rares, (2) le genre Hormiops est

de toute évidence endémique de la petite île de

Poulo-Condore située au sud de l’ancienne

Indochine française, aujourd’hui Vietnam, (3)

l’évolution meme de l’histoire de la région indo¬

chinoise depuis un demi-siècle, n'a pas autorisé

des recherches ou des inventaires précis sur la

faune existante.

Dans une étude relativement récente (Lourenço

1985, 1989), visant à situer de manière précise la

position taxonomique de la famille des

Ischnuridae par rapport à celle des Scorpionitlae,

la totalité des genres appartenant aux deux

familles ont été révisés. Au sein des Ischnuridae

deux genres ont posé des problèmes et sont restés

dans une position d'impasse. Le statut de l'un

d’entre eux, Chiromacbetes Pocock, 1899 alors

monotypique et connu d’après un seul spécimen

femelle, a pu être finalement clarifié avec la

découverte d’une deuxième espèce décrite à par¬

tir d’un spécimen mâle (Lourenço 1997), autori¬

sant ainsi l’étude des hemispermatophores. Le

second genre à poser des problèmes au cours de

la révision de Lourenço (1985, 1989) était bien

Hormiops , en raison de la fragilité des caractères

annoncés par Fage (1933) pour le distinguer du

genre Hormurus à présent synonyme de Uorbeles

Sundevall, 1833-

Danx le cadre de la révision du genre Liocheles

entreprise par un de nous (L. M.), une définition

précise du genre Hormiops s’avérait nécessaire.

L’étude des types A'Hormiops, ainsi que d’une

femelle et sa portée, collectées également à

Poulo-Condore, nous a permis la mise en éviden¬

ce d’autres caractères diagnostiques. Ces carac¬

tères ont été tirés de la trichobothriotaxie, mais

en particulier de l’examen des hémi.spermato-

phores du seul mâle existant dans la série

typique. L’hémispermatophore à'Hormiops pré¬

sente une structure plus primitive avec les cro¬

chets bien plus réduits, étant ainsi nettement

différents de ceux du genre Liocheles , mais égale¬

ment de l’ensemble de ceux observés chez les

autres genres de la famille des Ischnuridae.

SYSTÉMATIQUE

Genre Hormiops Fage, 1933

ESPÈCE-TYPE. — Hormiops davidovi Fage, 1933, par

tnonotypie.

REPARTITION GÉOGRAPHIQUE. — île de Poulo-

Condore, Vietnam.

NOUVFT.1 H DIAGNOSE DU GENRE

Le genre Hormiops , tel que son nom le suggère a

été associé au génie Hormurus (à présent syno¬

nyme de Liocheles) par Fage (1933), ce dernier

étant également distribué dans la partie méridio¬

nale de I Indochine française. Scorpion de taille

moyenne à petite, avec une longueur totale

d’environ 4 à 4,5 cm. Les différentes régions du

corps et des appendices très peu sculptées,

presque lisses. Tranchants des doigts mobiles des

pédipalpes avec une granulation digitale unifor¬

me. disposée en deux séries longitudinales, avec

absence des granules plus gros. A remarquer, la

sculpture des granulations des doigts mobiles des

pédipalpes est un caractère diagnostique pour les

genres appartenant à la famille des Ischnuridae

(Lourenço 1985). Le genre Hormiops peut être

distingué du genre Liocheles par les caractères sui-

338

ZOOSYSTEMA • 1999 • 21 (2)

Confirmation de la validité du genre Hormiopi

Fig. 1. — Hormiops davidovi (d’après Fage, 1933), lectotype 6, habitus. Échelle : 2 mm.

vants : (1) présence de deux yeux latéraux au lieu

de trois ; (2) trichobothriotaxie du type C ortho-

bothriotaxique, avec cependant la trichobothrie

db de la pince sur la face dorsale alors que chez

ZOOSYSTEMA • 1999 • 21 (2)

d’autres Ischnuridae elle est le plus souvent

proche de la limite entre les faces interne et dor¬

sale, voire sur la face interne. Par ailleurs, la tri¬

chobothrie Dt se situe à la limite des faces

339

Confirmation de la validité du genre Hortniops

dorsale et externe ; (3) hémispermatophore très

simple avec des crochets réduits; lamelle distale

de taille équivalente à la portion basale.

Honniops davidovi Fage, 1 933

(Figs 1-4)

Hormiops davidovi Fage, 1933 : 32, 3 figs. - Lourenço

1985 : 34, 1 fig. ; 1989 : 170.

Matériel examiné. — Types : MNHN-RS-0562.

1 6 désigné lectotype, 2 $ 9 désignées paralecto-

tvpes. Potilo Condore, forêt de Sudonnai (sous

pierres), M. C. Dawydoff coll.. 1I.1930-1V.1931. leg.

1932.

Remarque. — Fage (1933) indique pour la série

typique un mâle avec une interrogation pour sa condi¬

tion d’adulte, une femelle et un juvénile. 1. examen de

ce matériel nous permet de confirmer un mâle et deux

femelles tous adultes. Le spécimen femelle supplémen¬

taire collecté avec sa portée de I I petits (trois mâles et

huit femelles), se trouvait dans les collections du

MNHN, R.S-0499 avec les indications suivantes : tle

Poulo-Condor, leg. Germain (sans date). Det =

Hormurus awtrdlaûae Kraep. det 1900. Hormurus

complanatus E. Simon det.

Redescription fondée sur le male lectotype

Coloration. La couleur de base est brun rou¬

geâtre (la coloration actuelle est estompée).

Prosoma : plaque prosomienne de couleur brun

rougeâtre. Tubercule oculaire et régions des yeux

médians et latéraux noirâtres. Mesosoma de

même couleur que If prosoma. Metasoma avec

les cinq anneaux de couleur brun rougeâtre, plus

foncés que le prosoma et le mesosoma. Vésicule

jaunâtre avec l’aiguillon rougeâtre. Sternites jau¬

nâtres. Peignes, opercule génital, sternum,

hanches et processus maxillaire jaunâtre clair.

Pattes jaunâtres. Pédipalpes brun rougeâtre avec

la face intérieure du fémur et du tibia rouge

noirâtre. Chéliccres jaunâtres sans taches bru¬

nâtres ou trame.

Morphologie. Lront de la plaque prosomienne

moyennement échancré. Tubercule oculaire plus

ou moins au centre de la plaque prosomienne,

mais légèrement antérieur ; yeux médians séparés

par moins d’un diamètre oculaire. Deux paires

d’yeux latéraux. Sillon interoculaire très peu pro¬

fond, s’allongeant légèrement jusqu’à l'arrière des

yeux médians. Toute la plaque prosomienne est

Tableau 1. — Mensurations (en mm) du 4 lectotype et d'une 9

paralectotype d' Hormiops davidovi.

Prosoma

Longueur

4,5

5,1

Largeur antérieure

2.3

2,4

Largeur postérieure

5,1

5,4

Anneau caudal 1

Longueur

1,5

1,7

Largeur

1,1

1,2

Anneau caudal V

Longueur

2,3

2,1

Largeur

0,8

Hauteur

1.0

1,0

Vésicule

Largeur

0,7

0,8

Hauteur

0,6

0,8

Pédlpalpe

Fémur longueur

6,0

5,2

Fémur largeur

1,6

1,8

Tibia longueur

5.6

5,2

Tibia largeur

2.0

2,1

Pince longueur

9.8

9,8

Pince largeur

2,2

3,2

Pince hauteur

1,6

1,8

Doigt mobile

Longueur

3,8

4,2

lisse sans granulations. Mesosoma : tergites lisses.

Carène axiale très peu perceptible. Metasoma à

anneaux arrondis, sans carènes marquées ;

quelques granules spiniformes sont présents sur

la face ventrale du cinquième. Telson très

allongé ; aiguillon proportionnellement très

court. Vésicule dépourvue de carènes. Peignes

petits avec 7 -~ dents. Pédipalpes ; fémurs à cinq

carènes ; tégument intercarénai avec une granula¬

tion très fine. Tibia et pince avec très peu de gra¬

nules et avec la présence de quatre carènes

complètes. Présence sur la face interne du tibia

d’un grand éperon dans la région basale, avec

deux granules à l’extrémité. Pince aplatie ; face

dorsale dépourvue de granulations ; faces laté¬

rales avec des granulations moyennes ; face ven¬

trale brillante et lisse. Doigts lisses ; tranchant

des doigts mobiles avec la granulation disposée

en deux séries uniformes longitudinales, non

ZOOSYSTEMA • 1999 ■ 21 (2)

.343

Lourenço W. R. & Monod L.

fusionnées à la base et non interrompues par des

granules plus gros. Tarses des patres avec deux

rangées de soies parallèles. Chélicères avec la den¬

tition caractéristique de la famille des Ischnu-

ridae. Trichobothriotaxie du type C, ortho-

bothriotaxiquc ; la trichobothrie db de la pince

située sur la face externe ; trichobothrie Dt se

située à la limite des faces dorsale et externe.

Valeurs morphométriques exprimées dans le

Tableau 1.

Paralectotypes femelles

Mêmes données que pour l’holotype. Coloration

légèrement plus foncée. Morphologie similaire à

celle du mâle, mais avec des valeurs morphomé¬

triques différentes (voir Tableau 1). Taille légère¬

ment plus grande ; peignes plus petits avec

6-6 dents. La distinction entre mâles et femelles

est possible, comme pour tous les Ischnuridae,

par la morphologie de la plaque qui renferme

l’opercule génital : divisée en deux chez les mâles

et fusionnée chez la femelle.

Nombre de dents des peignes observé : 6-6 pour

la femelle adulte et pour les huit juvéniles

femelles ; 7-7 pour les trois juvéniles mâles.

Remerciements

Nous remercions vivement MM. M. Gaillard et

J. Rebière, Laboratoire de Zoologie (Arthro¬

podes) MNHN, pour la réalisation de plusieurs

illustrations et le Dr S. Jourdan (Paris) pour la

révision du manuscrit. Le second auteur (L. M.)

tient à remercier tout particulièrement le

Département des affaires culturelles de la Ville de

Genève pour son soutien financier.

RÉFÉRENCES

Fage L, 1933. — Les Scorpions de l’Indochine fran¬

çaise. Leurs affinités, leur distribution géogra¬

phique. Annales de la Société entomologique de

France 102:25-34.

Lourenço W. R. 1985. — Essai d'interprétation de la

distribution du genre Opisthacanthus (Arachnida,

Scorpiones, Ischnuridae) dans les régions Néotropicale

et Afrotropicale. Etude taxinomique, biogéographique,

évolutive et écologique. Thèse de Doctorat d’Etat,

Université Pierre et Marie Curie, Paris, France,

287 p.

Lourenço W. R. 1989. — Rétablissement de la

famille des Ischnuridae, distincte des Scorpionidae

Pocock, 1893, à partir de la sous-famille des

Ischnurinac Pocock, 1893. Revue Araclmologique 8

(10) : 159-177.

Lourenço W. R. 1996. — Scorpions. Faune de

Madagascar 87 ,• 1-102.

Lourenço W. R. 1997. — Considérations taxono¬

miques sur le genre Chiromachetes Pocock, 1899

(Chelicerata, Scorpiones, Ischnuridae). Zoosystema

19 (1) : 81-89.

Soumis le 8 juin 1998 ;

accepté le 13 octobre 1998.

344

ZOOSYSTEMA • 1999 • 21 (2)

Un Heterocarpus nouveau (Crustacea,

Decapoda, Pandalidae) du Pacifique Sud-Ouest

Alain CROSNIER

ORSTOM, Laboratoire de Biologie des Invertébrés marins et Malacologie,

Muséum national d’Histoire naturelle, 55, rue de Buffon, F-75231 Paris cedex 5 (France)

crosnier@mnhn.fr

Crosnier A. 1999. — Un Hetocarpus nouveau (Crustacea, Decapoda, Pandalidae) du

Pacifique Sud-Ouest. Zoosysiema 21 (2) : 345-351.

MOTS CLÉS

Crustacea,

Decapoda,

Caridea.

Pandalidae,

eau profonde,

océan Pacifique,

Nouvelle-Calédonie,

espèce nouvelle.

RÉSUMÉ

Une espèce nouvelle, Heterocarpus intermedius, confondue lors de premières

identifications avec H. woodmasoni Alcock, 1901, est décrire d'après des spé¬

cimens récoltés sur la côte est de l’Australie, en Nouvelle-Calédonie, au voisi¬

nage des îles Loyauté et Chesterfield et sur les bancs Combe et Tuscarora.

Elle se caractérise, en particulier, par l'absence d'une crête postrostrale et la

présence de deux paires d’épines dorsolatérales seulement sur le telson. Un

complément à la dé d identification des Heterocarpus publiée par Crosnier

(1988) est proposée.

KEYWORDS

Crustacea,

Decapoda,

Caridea,

Pandalidae,

deep water,

Pacific Océan,

New Caledonia,

new species.

ABSTRACT

A new Heterocarpus (Crustacea, Decapoda, Pandalidae) from South-West

Pacific.

A new species, Heterocarpus intermedius, confused until now with H. wood¬

masoni Alcock, 1901, is described after specimens caught off die east coast of

Australîa, New Caledonia, the Loyalty and Chesterfield islands, and ihe

Combe and Tuscarora banks. It can be separated mainlv by the fact that it

has no postrostral cresr and only two pairs of dorsolateral spines on the tel¬

son. An addition co the identification key of the Heterocarpus species publî-

shed by Crosnier (1988) is proposed.

ZOOSYSTEMA • 1999 • 21 (2)

345

Crosnier A.

Parmi les Heterocarpus A. Milne Edwards, 1881

récoltés en Nouvelle-Calédonie depuis 1985,

nous en avions identifiés de très nombreux à

H. woodrmtsoni Alcock, 1901, espèce bien carac¬

téristique et supposée largement répandue dans

le Sud-Ouest Pacifique.

Un réexamen récent de ces spécimens nous a

maintenant convaincu qu'ils appartiennent à une

espèce nouvelle que nous décrivons dans les

pages qui suivent.

Sauf indication contraire, les spécimens examinés

sont déposés dans les collections du Muséum

national d Histoire naturelle (MNHN), Paris.

Abréviations utilisées dans les i-iglires

LT longueur totale

P3 - P5 troisième à cinquième péréiopodes

Famille PANDAl.lDAE Hayworth, 1825

Genre Heterocarpus A. Milne-Edwards, 1881

Heterocarpus mtermedius n. sp.

(Kg. 1)

Types. — Une femelle ovigère dont la carapace mesure

29,6 mm, récoltée à la station CP .163 de la campagne

MUSORSTOM 5 aux îles Chesterfield, est l'holotype

(MNHN-Na 13142). Les autres spécimens, à l’excep-

tion de ceux récoltés en Australie dont tous les rostres

sont cassés, sont des paratypes.

Matériel examiné. — De nombreux spécimens dont

la longueur de la carapace est comprise entre 10 et

36 mm, ont été examinés.

Australie. Côte est. Southern In trader, 23°52’S,

153°02’E. 650 m, 29XLI983. 4 ex. (QM-

W113I7). — 23“45 S, I53“07’E, 550 m,

29.XI.I983, 3 ex. (QM-Wt 1309). — Valkyrie

Voyager, 15"58'S, 149°56'E, 590 m, 30.VL 1989,

1 ex. (QM-WI62II).

Nouvelle-Calédonie. 13IOCAL : stn CP 40, 650 m,

1 ex. — Stn CP 109.495-515 m, t ex.

MUSORSTOM 4 : stn DW 156, 525 m, 2 ex. — Stn

CP 178. 520 m, I ex. — Stn CP 198, 585 m, 31 ex.

— Stn CP 2T6, 490-515 m, 1 ex. — Srn CP 236, 495-

550 m, 2 ex. — Stn CP 241, 470-480 m, 1 ex.

(MNHN-Na 9929) — Stn 242. 500-550 m, 2 ex.

BATHUS 3 stn CP 845. 592-622 m, 11 ex.

BATHUS 4 : stn CP 910, 560-608 m, 1 ex. — Srn

CP 921, 610-613 m, 3 ex. - Stn CP 922, 600 m,

2 ex. — Stn CP 948, 533-610 m. 1 ex.

HALIPRO 1 : stn CH 872, 620-700 m, 3 ex. — Stn

CH 873, 640-680 m, 11 ex.

îles Loyauté. MUSORSTOM 6 : stn CC 470,

560 m. 4 ex.

îles Chesterfield. MUSORSTOM 5 : stn DW 341,

675-680 m. 1 ex. — Stn DC 358, 680-700 m, 1 ex.

— Sut CP 359, 700-720 m, 3 ex. — Stn CP 363,

685-700 m, 47 ex. (dont 3 au NHM, Londres et 3 au

NMNH. Washington).

CO RAI I 2 : stn DE 13, 700-705 m, I ex.

Vanuatu. MUSORSTOM S : stn CP 992, 775 .

748 ni, 7 ex. — Stn CP CP 1007, "80-8.10 m, 1 ex.

— Sm CP 1028-, 624-668 m, 5 ex. — Stn CP 1089,

494-516 ni) 3 ex. —Stn CP 1090, 470-502 m, 2 ex.

Banc Combe ( 12°27*S, 177°28’W). MUSORS¬

TOM 7 : stn DW 540, 600 m, 2 ex.

Banc Tuscarora (ÎPAJ'S, 178’20’W). MUSORS¬

TOM 7 : stn DW 560, 697-702 m, 1 ex.

Répartition. — Cette espèce n'est encore connue

que de la cote esc d’Austiulie, de la Nouvelle-

Calédonie, des îles Loyauté et Chesterfield, du

Vanuatu et des bancs Tuscarora (ll f '47'S, 178°20’W)

et Combe ( I2 0 27'S, l'TS'Tl, entre 500 et "50 m

au moins. Heterocarpus woodmasotti n’ayant jamais été

trouvé en mélange avec elle, il semble qu elle prenne le

relais de cette espèce, dans le Pacifique Sud-Ouest, à

l’est de l'Australie et de l’Indonésie.

Description

Le rostre modérément recourbé vers le haut, n’a

pas de carène latérale nette ; sa longueur est com¬

prise entre 1,4 (grands adultes) er 2,1 fois (petits

spécimens) celle de la carapace ; son bord dorsal

est armé de neuf à onze (le plus souvent dix)

dents dont la première est en arrière de l'orbite et

la seconde au niveau du fond de l’orbite ; son

bord ventral porte de sept à dix (le plus souvent

huit) dents. Il n’y a pas tic crête rostrale marquée

(un semblant de crête, toutefois, chez les juvé¬

niles — Fig. JD - qui disparait totalement par la

suite), mais par contre une carène postrosttale,

bien en relief, à section arrondie, qui atteint

presque le bord postérieur de la carapace. Les

faces latérales de cette dernière portent chacune

deux carènes très nettes qui prolongent directe¬

ment les épines atitennaire et branchiostège et

atteignent presque le bord postérieur de la cara¬

pace. L’épine branchiostège, plus courte que

l'antennaire, a son extrémité qui se situe en deçà

de celle de cette dernière.

Les yeux, bien développés, sont sans ocelle ; leur

cornée est colorée.

Le stylocérite s’étend jusqu’au tiers, environ, du

346

ZOOSYSTEMA • 1999 • 21 (2)

Un Heterocarpus nouveau

Fig. 1. — Heterocarpus intermedius n. sp. A-C, S ovigère 29,6 mm, holotype, îles Chesterfield, MUSORSTOM 5, stn CP 363, 700-

685 m (MNHN-Na 13442) : A, partie antérieure du corps, vue latérale ; B. abdomen, vue latérale ; C, telson vu de dessus D, iuvé-

nile 14,2 mm, Nouvelle-Calédonie, MUSORSTOM 4, stn 241, 470-480 m (MNHN-Na 9929). partie supérieure de la carapace.

Échelles : 1 cm.

second segment du pédoncule antennulaire. Le

scaphocérite est, chez l’adulte, environ 3,3 lois

plus long que large ; sa lame dépasse très nette¬

ment l’épine distolatéraîe externe.

Les troisièmes maxillipèdes n’atteignent pas tout

à fait la pointe vie l'épine distolatérale externe du

scaphocérire ; leur exopodite est bien développé,

atteignant presque F extrémité de l'antépénul¬

tième segment ; leur dernier article est de 1,75 à

1,9 fois plus long que l'avant-dcrnier.

Les seconds péréiopodes sont inégaux ; le plus

court atteint l’extrémité du pédoncule antennaire,

le plus long dépasse des doigts de sa pince le sca¬

phocérite ; le plus souvent le plus court et le plus

fort est le droit, mais pat lois c’est le contraire qui

est observé ; le carpe du péréiopode court et fort

compte de onze à quatorze articles (le plus sou¬

vent douze), celui du péréiopode long et plus

grêle de ving-quatre à vingt-six articles. Les troi¬

sièmes péréiopodes ont un dactyle dont la lon¬

gueur est très légèrement supérieure à la moitié

de ceile du propode ; il en est de même du carpe

par rapport au propode. La répartition des épines

sur l ischion, le mérus et le carpe des trois der¬

nières paires de péréiopodes est donnée dans le

Tableau 1. Tous les péréiopodes, à l excepdon des

cinquièmes, portent un épipodite.

Les premier et second segments de l’abdomen

sont sans carène dorsale, le premier présente, en

arrière de son bord antérieur, deux gros granules.

Les trois suivants sont carénés dorsalement,

aucune des carènes ne se terminant par une

épine ; le troisième porte une forte épine dorsale

qui s’élève, à parrir de la carène, vers son milieu ;

le quatrième montre l’esquisse d'un tubercule

dorsal aux trois quarts de sa longueur : le sixième

porte une paire de carènes dorsales séparées par

une dépression et, sur chacune de ses faces laté¬

rales, une carène longitudinale située aux trois

quarts de sa hauteur et s’étendant sur les deux

ZOOSYSTEMA • 1999 • 21 (2)

347

Crosnier A.

Tableau 1. — Répartition des épines sur l'ischion, le mérus et le carpe des troisièmes, quatrièmes et cinquièmes péréiopodes chez

Heterocarpus calmani, H. woodmasoni et H. intermedius.

Ischion

Mérus

Carpe

Heterocarpus

calmani

2-3

6-9 (8-9)

8-9

2-4

3-4

2-3

0-7 (4-5)

woodmasoni

1-4 (2)

2-3 (2)

8 -18 (10-14)

6-18 (12-15)

3-12 (9-11)

0-7 (4-5)

1-7 (2-5)

intermedius

2-4 (2-3)

0-2

10-14(13-14)

11-15

9-12(10)

4-5

2-3

1-3

tiers de sa longueur, Les pleurons des quatre pre¬

miers segments sont arrondis ; celui du cin¬

quième porte une épine inférciposférieure. Le

sixième segment est de ] ,8 à 1,9 fois plus long

que haut. Le telsoti est environ 1,5 lois plus long

que le sixième segment et très effilé à son extré¬

mité ; il est armé de 1 deux paires seulement

d’épines dorsolarérales et d’une paire d'épines

subterminales ; ces dernières sont implantées sur

la face inférieure du relson et sont souvent éro¬

dées chez l’adulte, si bien qu elles ne sont plus,

alors, visibles en vue de dessus.

Taille. Le plus grand spécimen capturé esc une

femelle dont la carapace mesure 36 mm (LT voi¬

sin de 170 mm).

Coloration. L’ensemble de l'animal est d’un

rouge soutenu avec de nombreuses pairies lavées

de blanc (faces latérales des segments abdomi¬

naux, extrémités des scaphocérires, des pléopodcs

et des uropodes, larges portions des péréiopodes

3-5, flagelles antennulaires ci amennaires). Le

rostre présente des bandes transversales alternati¬

vement blanches et rouges, la taille de ces der¬

nières s’amenuisant au fur et à mesure que l’on

va vers l’extrémité du rostre, La forte épine dor¬

sale du troisième segment abdominal ne présente

pas de coloration particulière.

Remarques

Jusqu’à une époque récente, les Heterocarpus

dont la carapace possède une carène postanten-

naire (c’est-à-dire une carène dont l’extrémité

antérieure se confond avec l'épine antennaire ou

est située dans son prolongement immédiat) et

est sans carène cardiolatérale sur sa partie posté¬

rieure, dont tous les segments abdominaux sont

sans épine médiane sur leur bord postérieur et

dont la face dorsale du troisième segment abdo¬

minal porte une forte épine vers son milieu,

étaient identifiés à H woodmasoni Alcock, 1901.

En 1988, nous avons montré, suivant en cela ce

que Caïman (1939) puis Chace (1985) avaient

envisagé, que les spécimens provenant de l’océan

Indien occidental appartiennent à une espèce

nouvelle que nous avons nommée H. calmani

Crosnier, 1988.

H apparaît maintenant que les Heterocarpus de ce

mcine groupe, capturés: de l'est de l’Australie

jusqu’au banc Tuscarosa (178 r, W) appartiennent

à une troisième espece, que nous nommons

intermedius pour rappeler qu elle présente des

caractères appartenant tantôt à 1 une, tantôt à

l’autre des deux autres espèces du groupe.

En effet, H. intermedius partage avec H. calmani

un telson portant deux paires seulement d’épines

dorsolatéraies et avec H. woodmasoni l’absence

d une crête au niveau des trois premières dents

postrostralcs et rostrales.

H. intermedius se distingue essentiellement de

H '. calmani par :

— l’absence d'une crête postrostrale (comparez la

l igure la avec la figure la de Crosnier 1988) ;

sur le vivant, l’absence d’une tache touge vif,

zébrée de blanc, au niveau de la. grosse épine dor¬

sale du troisième segment abdominal (voir la

figure 1 b de Crosnier 1988).

On peut citer, en outre, quelques différences sou¬

vent minimes au niveau de la dentition (bord

ventral du rostre avec sept à dix dents - le plus

souvent huit - contre six ou sept chez H. calma¬

nt), de la spinulation des péréiopodes (notam¬

ment des mérus : dix à quatorze épines sut le

mérus des P3, onze à quinze sur celui des P4 et

neuf à douze sur celui des P5, contre respective¬

ment six à neuf, huit à neuf deux à quatre chez

H. calmani) et du nombre des articles du carpe

348

ZOOSYSTEMA • 1999 • 21 (2)

Un Heterocarpus nouveau

des P2 (onze à quatorze et vingt-quatre à vingt-

six contre huit à onze et vingt à vingt-trois chez

H. calmant),

H. intermcdius se distingue essentiellement de

H. woodmasoni par le telson qui ne porte que deux

paires d’épines dorsolarérales au lieu de quatre.

La dentition, la spinulation des péréiopodes

(cf. Tableau 1 ) et le nombre des articles du carpe

des deuxièmes péréiopodes ne présentent pas de

différences vraiment significatives entre ces deux

espèces. Il est possible que des différences de

coloration existent sur le vivant, mais nous

n’avons pas eu la possibilité de le vérifier.

Le Dr Davic du Queensland Muséum nous

ayant envoyé des Heterocarpus récoltés en

Australie, se trouvant dans la collection de son

Muséum, nous y avons trouvé, outre les' H. inter-

medius mentionnés plus haut :

— des H. woodmasoni [Côte nord-ouest. Sans

autre précision, 3 ex. (QM-W 11962). — Côte

nord-est. 1 1°35*S. I44°04’E, L.N.L. de l’île

Raine, 420 m, 12.11.1979, 38 ex. (QM-

W8235). — G-May, 9°51’S, 144°11’E, à l’est

des îles Murray, 460-464 m, 27.V. 1993, 44 <3,

33 2 (QM-W 11208)] ;

- et des H. calmant [Côte est. Southern lntruder,

23°59’S, 152“59’E, 380 m, 8.VIII.I983, 8 ex.

(QM-W 11460). — 23°46’S, I53°06’E, 550 m.

9.VM.1983, I ex. (QM-W 11417). — 23°54’S,

1 53°01’E, 465 ni, 29.XI.1983, 8 ex. (QM-

W 11315). — 23°52\S, 153°02’E, 650 m.

29.XI.1983, 3 ex. (QM-W 11293). — 23°22’S,

152°45’E, 350-310 m, 30.XI.1983, 5 ex. (QM-

W 11283). — 23°52’S, 153°02’E, 650 m,

29.X1.1983, 1 ex. (QM-W 11286). — 23°21’S,

153°23’E, 410 m, 30.XL 1983, 6 ex. (QM-

W 11308).

CRAICMIN SURVEY : srn 6, 23°28’S,

153“19’E, 562 m, 20.IX.1980, 1 ex. (QM-

W 10077) et 1 ex. (QM-W 10063). — Stn 24,

23 n 00,3’S, 152°12,5’E, 387 m, 3.X.1980, 5 ex.

(QM-W 10062)1.

Il est à noter que c’est la première fois, à notre

connaissance, que H. calmant est mentionné en

dehors de l'océan indien.

Une question que l’on peut se poser est, bien

entendu, de savoir si l’espèce décrite par Alcock,

en 1901, de la mer des Andamans, par 485 m de

profondeur, est bien celle dont le telson porte

quatre paires d’épines dorsolarérales. Alcock ne

mentionne pas ce caractère dans sa description

et, par ailleurs, sur le dessin de l’espèce qui a été

publié (Alcock & McArdle 1901 : pl. 51, fig. 2),

les épines dorsolatérales du telson ne sont pas

marquées. Les types d Alcock devraient se trou¬

ver au Zoological Survcy of India, à Calcutta,

sous le numéro 3124/9. Malhcuseuscmcnt,

toutes nos correspondances avec cet organisme

sont demeurées, comme à l’habitude, sans répon¬

se. Si l’on considère toutefois les répartitions géo¬

graphiques des deux espèces telles qu elles sont

connues actuellement, il semble logique

d’admettre que l’espèce d’Alcock est celle dont le

telson porte quatre paires d épines,

Dans la clé des espèces du genre Heterocarpus que

nous avons publiée en 1988, la séparation de

H, woodmasoni et //. calmant se fait au point

quatre. Compte tenu de la description de

H. intermcdius , ce point 4 doit se présenter

maintenant sous la forme :

4. Telson portant quatre paires d’épines dorsolatérales (sans compter celles de l’extrémi¬

té) . H. woodmasoni Alcock, 1901

(Nier des Andamans, côtes nord et nord-est de l’Australie, Indonésie, Philippines, mer

de Chine méridionale, Japon, dont nous avons examiné un exemplaire récolté en baie

de Tosa ; 291-655 m)

— Telson portant deux paires d'épines dorsolatérales...4bis

4bis. Trois premières dents postrosrrales et rostrales supérieures implantées sur une

légère crête. Epines du troisième segment abdominal colorée, ainsi que le voisinage de

sa base, en rouge foncé avec des zébrures blanches .. H. calmant Crosnier, 1988

(Océan Indien occidental sud : côte est-africaine, Madagascar. Côte est de l’Australie ;

250-650 m)

ZOOSYSTEMA • 1999 • 21 (2)

349

Crosnier A.

— Trois premières dents postrost raies et roseraies supérieures non implantées sur une

crête. Épine du troisième segment abdominal sans coloration particulière .

........... H. intermedius n. sp.

(Côte est de l’Australie, îles Chesterfield, Nouvelle-Calédonie, Vanuatu, bancs

Tuscarora et Combe : I l°47’.S, 178°20’Wet 12°27’S, 177°28’W ; 500-750 m)

Lorsque cette clé a été établie, elle n’a pu prendre

en compte Heterocarpus agassizi Allen et Butler

qui n’a été décrit qu’en 1994 (Allen & Butler

1994 : 431, figs 6-9).

Cette espèce semble très proche de H. longirostris

MacGilchrisc, 1905, et ne lui a malheureusement

pas été comparée par Allen & Butler. Par ailleurs

Tholotype et les 119 para types de H, agassizi ne

semblent pas avoir été renvoyés dans les collec¬

tions de la Scripps Institution of Oceanography

et demeurent introuvables.

Dans ces conditions, on ne peut sc fonder que

sur la description et les dessins publiés de cette

espèce pour la comparer à H longirostris dont

elle semble se distinguer par :

- un rostre nettement plus court, d’une longueur

voisine de celle de la carapace (au lieu d’être de

1,5 à 1,75 fois plus long que la carapace) ;

- la présence d’un très court exopode sur les troi¬

sièmes maxillipèdes (au lieu d'un simple bour¬

geon) ;

- peut-être des carènes dorsales abdominales un

peu moins larges (mais les dessins de H. agassizi

publiés sont loin d'être excellents et nous parais¬

sent un peu sujet à caution).

Dans une lettre, le Dr Allen nous a mentionné

les autres différences suivantes :

- le nombre de dents rosrrales plus faible ( 10-11

au lieu de 12-14 chez H. longirostris) ;

- la présence d une carène dorsale médiane,

arrondie, sur les segments 2-5 (au lieu de 3-6

chez H. longirostris ), la face dorsale du segment 6

chez. H. agassizi étant aplatie avec seulement

l’esquisse d’une courte carène centrale ;

- le scapbocérite dont la longueur est intérieure à

la moitié de celle de la carapace (au lieu d'être

égale aux trois quarts chez H, longirostris),

Le caractère fourni par le rostre est le plus specta¬

culaire, mais il est souvent assez variable chez une

même espèce en fonction de la taille et des prove¬

nances géographiques (voir Crosnier 1988 : 62, à

propos de H. dorsalis). Il serait donc souhaitable

de pouvoir comparer des séries suffisantes des

deux espèces afin de confirmer leurs caractères

distinctifs.

Avec nos connaissances actuelles, la clé que nous

avons publiée en 1988 peut être complétée, au

moins provisoirement, de la manière suivante :

24. Rostre à peu près aussi long que la carapace.25

— Rostre plus de 1,5 fois plus long que la carapace.

...................................................... H. longirostris Ma cG il christ, 1905

(Iles Maldives, mer des Andamans, ouest de Sumatra, îles Mariannes ; 1 000-1 756 m)

25. Rostre sans dents sur la moitié distale de son bord dorsal. Epine antennaire beau¬

coup plus longue que l’épine ptérygostomienne. Troisièmes maxillipèdes avec un exo¬

pode réduit à un bourgeon.. H. grimaldii À. Milnc Edwards & Bouvier, 1900

(Atlantique oriental, du Portugal aux Açores et à l'Angola ; 914-2 834 m)

— Rostre denté sur route la longueur de son bord dorsal. Epine antennaire de même

taille que I épine ptérygostomienne. Troisièmes maxillipèdes avec un exopode court mais

bien distinct. H, agassizi .Allen & Butler, 1994

(Pacifique central, 22°07.7’N, 171°36,0’E ; 1 281 m)

350

ZOOSYSTEMA • 1999 • 21 (2)

Un Heterocarpus nouveau

La conclusion que ces quelques pages nous amè¬

nent à formuler est que la systématique des

Heterocarpus réserve certainement encore bien

des surprises. Il semble s'avérer que des groupes

formés d’espèces extrêmement proches les unes

des autres existent, la coloration fournissant

alors, souvent, un élément distinctif précieux

lorsqu'elle peut être observée sur des animaux

très frais. Un exemple récent a été fourni par le

groupe d’espèces proches de H. emifer A. Milne

Edwards, 1881 (voir Crosnier 1988), et nous en

donnons ici un autre exemple.

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Alcock A. & McArdle A. F. 1901, — Illustrations of

the Zoology of the Royal Indian Marine Survey Ship

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Soumis le 12 mars 1998 ;

accepté le 3 juillet 1998.

ZOOSYSTEMA • 1999 • 21 (2)

351

The identity of Chlorodius miliaris A. Milne

Edwards, 1873, and the establishment

of a new genus of Chlorodiinae (Crustacea,

Decapoda, Brachyura, Xanthoidea, Xanthidae)

from New Caledonia

Paul F. CLARK

Department of Zoology, The Natural History Muséum,

Cromwell Road, London SW7 5BD (England)

pfc@nhm.ac.uk

Peter K. L. NG

School of Biological Sciences, National University of Singapore,

Kent Ridge, Singapore 119260 (Republic of Singapore)

dbsngk!@leonis. nus.edu.sg

Clark P. F. & Ng P, K. L. 1999. — The identity of Chlorodius miliaris A, Milne Edwards,

1873, and the establishment ot a new genus ot Chlorodiinae (Crustacea, Decapoda,

Brachyura, Xanthoidea, Xanthidae) from New Caledonia. Zoosystema 21 (2) : 353-365.

KEYWORDS

Xanthidae,

Chlorodiinae,

taxonomy,

new genus,

Indo-West Pacific.

ABSTRACT

The identity of the poorly known xanthid crab Chlorodius mtlidris A. Milne

Edwards, 1873, front New Caledonia is clarified, and a new genus of

Chlorodiinae is dcscribed foi lire species. Sulcodius n. gcn. possesses well-

defined carapace régions; distinctly granulated cheliped surfaces, an tntero

external angle of the basal anrennal segment rhat exiends ituo the orbital hia¬

tus, filling it entirely; and a cheliped merus wirh a strongly grooved dorsal

surface. The new genus Is allied lo Chlorodielln Rarhbun, 1897, Pilodius

Dana, 1851 and Phymodius A. Milne Edsvards, 1863 and the relationships bet-

ween the four généra are discussed, and distinguishing characters tabulated.

MOTS CLÉS

Xanthidae,

Chlorodiinae,

taxonomie,

genre nouveau.

Pacifique indo-ouest

RÉSUMÉ

Identité de Chlorodius miliaris A. Milne Edwards, 1873 et établissement d'un

nouveau genre de Chlorodiinae (Crustacea, Decapoda, Brachyura, Xanthoidea,

Xanthidae) de Nouvelle-Calédonie.

L’identité de Chlorodius miliaris A, Milne Edwards, 1873, crabe mal connu

de la famille des Xanrhidac, trouvé en Nouvelle-Calédonie, est clarifiée et un

nouveau genre île Chlorodiinae est décrit pour cette espèce, Sulcodius n, gen.

présente une carapace aux régions bien délimitées, des chélipèdes granuleux,

l'angle antéro-externe de l’article basal antennaire qui pénètre dans le hiatus

orbitaire et le remplit entièrement, et le mérus des chélipèdes a une surface

dorsale avec un profond sillon. Ce nouveau genre est proche de Chloroclielta

Rathbun, 1897, Pilodius Dana, 1851 et Phymodius A. Milne Edwards, 1863.

Les relations existant entre ces genres sont discutées. Un tableau récapitulant

leurs caractères distinctifs est proposé.

ZOOSYSTEMA • 1999 • 21 (2)

353

Clark P. F. & Ng P. K. L.

INTRODUCTION

The type sériés of Chlorodius miliaris A. Milne

Edwards, 1873, is extant and deposited in the

Muséum national d’Histoire naturelle, Paris. It

comprises three dried damaged syntype females

from New Caledonia which were only bricfly

described and schematically illustratcd. In a

report on the zoological collections of H MS

Alert, Miers (1884: 531) recorded some spécifie

variation when lie idemifled Chlorodius miliaris

from Seychelles and stated thaï the two adult

males “nierely differed from M.-Edwards’s des¬

cription based on specimens troni New

Caledonia”. A redescription of Chlorodius milia-

ris was given by De Man (1887: 280-281) when

he examined material colJected from the

Malaysia n Archipelago, liowever, h is identifica¬

tion was questioned by Ralss (1938: 38), Forest

& Guinot (1961: 96), Guinot (1964: 23) and

Serène (1984: 257). The phorographs ofa lemale

syntype from New Caledonia by Guinot (1964:

pl. X), clarifiée! the rrtorphological features of the

right cheliped (fig. 1) and the carapace (fig. 2).

Serène (1984: pl. XLIID) later reproduced the

carapace figure of Guinot (1964: pl. X, fig. 2),

puhlîshed a ltey separatïng the species attributed

to Chlorodiella (Guinot 1964: 254-255, 257),

but did not redescribe C. miliaris. To date a full

description, induding the structure of the sternal

plastron and male pleopods of Chlorodius milia¬

ris , is not available.

The taxonomy of Chlorodius miliaris is confused.

Rathbun (1897) consîdered Chlorodius H. Milne

Edwards, 1834 invalid and proposcd a new

replacement narne Chlorodiella. l'hc generic pla¬

cement of C. miliaris was further complicated

when Ralss (1938: 38) revlscd Zoozymodes (now

Zozymodrs) Heller, 1861 and stated that to this

genus belong the “C miliaris A.M.E.” lorm des¬

cribed by De Man. According to Balss

(1938; 52) the types of Chlorodius miliaris were

not seen because they were no longer extant in

the Paris Muséum, however he did examine the

material of De Man which w'as considered to be

identical ro Leptodius efferens Rathbun, 1907.

Ralss was uncertain about the derermination of

the Malaysian Archipelago material as C. miliaris

by De Man because he commented thaï this

could only be canfîrmed by the re-examination

of the A. Milne Edwards’ types (1873) sittee his

illustration (pl. 8, fig. 3) obviously referred to

Fig. 1. — Sulcodius miliaris (A. Milne Edwards, 1873); 3 14.5 mm, New Caledonia (MNHN MP-B 24818); dorsal view.

354

ZOOSYSTEMA ■ 1999 ■ 21 (2)

Identity of Chlorodius miliaris A. Milne Edwards, 1873

another species. But Balss (1938) appeared to

contradict apparent misgivings in the figure

legends ro lafel 11, fig. 2 (unpaginated), “Fig. 2.

Zoozywodes miliaris (A.M.F.) 2: I. De Mans

Exemplare von Nordwachter lslandThis cap-

don appeared to introduce a ncw combination

for Chlorodius miliaris and suggcsted that the

material of De Man was synonymous with the

spécimens front New Caledonia. l.ater, Focest &

Guinot (1961: 96) stated that the threc female

types of Chlorodius miliaris were still présent in

the collections of the Muséum national

d’Histoirc naturelle: they had examined the

ornamentation of the carapace and chelipeds,

and considered this species of A. Milne Edwards

to be vafid but not to bclong to Chlorodiella.

They agreed with Balss (1938: 38) rhat this spe¬

cies should not be attributed to Chlorodiella , but

disagreed with its placement in Zozymodes (as

Zoozymodes). Whilc reviewing Zozymodes,

Guinot (1964; 22-23) commented upon the

work of Balss (1938: 38). She also considered

Chlorodius milia H s A . Milne Edwards, 1873 as a

valid species, provided photographs of a female

syittype (1964: pl X, figs 1; 2) and regarder! its

placement by Balss in Zozymodes to be incorrect.

Further, Guinot considered the description of

Chlorodius miliaris by De Man ( 1888: 280) to be

poor. Although she had not seen the material

that Balss (1938: 38 Taf. II. Ptg. 2) had attributed

to Leptodius tfferem Rathbun, 1907 (p. 39, pl. 1,

fig. 11; pl. 7, lig. 6, 6a) and was not cotisequent-

ly in a position to place il in a genus, Guinot

concluded that the assignment of this Rathbun

species to Zozymodes by Balss was not justified.

In a recent révision of the Indo-Pacific Océan

and Red Sea Brachyura, Serène (1984: 257-238,

pl. XI.III D) referred the Chlorodius miliaris of

A. Milne F.dwards to Chlorodiella but did not

résolve the taxonomie position of this species

Fig. 2. — Sulcodius miliaris (A. Milne Edwards, 1873); chelipeds: A, <S 25.7 x 14.5 mm, New Caledonia (MNHN MP-B 24818): B, S

22.6 X 13.4 mm, New Caledonia (MNHN MP-B 24819). Scale bar: 2 mm.

ZOOSYSTEMA • 1999 • 21 (2)

355

Clark P. F. & Ng P. K. L.

Table 1.

— Generic différences between Phymodius. Pilodius,

Chlorodiella and Sulcodius n

. gen.

Characters

Sulcodius

Pilodius

Phymodius

Chlorodiella

Basal antennal

anteroexternal

angle

prolonged completely

filling orbital hiatus

(Fig. 6A)

Prolonged into orbital

hiatus (Fig. 6B)

not prolonged into

orbital hiatus.

(Fig. 6C)

not prolonged into

orbital hiatus

(Fig. 6D)

Antenna flagellum

does not enter orbital

hiatus (Fig. 6A)

does not enter orbital

hiatus (Fig. 6B)

enters orbital hiatus

(Fig. 6C)

enters orbital hiatus

(Fig. 6D)

Merus of the

cheliped

dorsal surface deeply

concave, strongly

grooved medlally

(Fig. t)

dorsal surface gently

concave, not deeply

grooved

dorsal surface gently

concave, not deeply

grooved

dorsal surface gently

concave not deeply

grooved

Ambulatory

dactylus

subterrninal spine

distmctly larger than

terminal spine

(Fig. 7A)

subterminal spine

smaller than terminal

spine (Fig. 7B)

subterminal spine

smaller than terminal

spine (Fig. 7C)

subterminal spine

subequal to equal

size to terminal spine

(Fig. 7D)

Anterior sternal

plastron

broad. male abdo¬

men reaching only

one-third length of

sternite 4

(Fig. 8A, B)

broad, male abdo¬

men reaching mid-

length of sternite 4

(Fig. 8C)

narrow, male abdo¬

men reaching mid-

length of sternite 4

(Fig. 8D)

broad, male abdo¬

men reaching one-

quarter length of

sternite 4 (Fig. 8E)

identified by De Man from Nordwachter Island

(1984: 181, 257).

Xanrhid crab material collecred by ORSTOM

(France) from New Caledonia was identified as

Chlorodiella miliaris. As the spécifie and generîc

starus of this species was questioned by a number

ofauthors, its rediscovc-ry in New Caledonia may

now help résolve à number of taxonomie pro-

blems. Examination of the présent sériés of

C. miliaris spécimens shows that this species can-

not be retained in Chlorodiella becausc it has a

very different carapace and antcnnal structure. In

the form of the carapace, C. miliaris resembles

members of Phymodius, but in the form of the

antennae, C. miliaris seems doser to Pilodius.

These différences suggest that a new genus

should be cstablished for C. miliaris in rhe

Chlorodiinae.

The description of the carapace régions tollowed

that proposed by Dana (1853). Mcasurcmcnts

provided are the carapace width.

Spécimens are deposited in the collection of the

Muséum national d’Histoire naturelle (MNHN),

Paris, France; The Natural Ilistory Muséum

(NHM), London, England, and the Zoological

Référencé Collection (ZRC), Department of

Zoology, National Unîversîty of Singapore.

SYSTEMATICS

Family XANTHIDAE

MacLeay, 1838, sensu Serène, 1984

Subfamily Cl ILOROD1INAE Alcock, 1898

Sulcodius n. gen.

ChlorodiusA. Milite Edwards, 1873: 212 (part).

Chlorodiella - Serène 1984: 254 (part),

Type species. — Chlorodius miliaris A. Milne

Edwards, 1873, by présent désignation. Monotypic.

ETYMOUX.Y, — The naine Sulcodius is dérivée! from

an arbltrary combination ol the Latin "suIntrus", allud-

Ing ro the distinctive fttrrow on the merus of the chelip-

ed, and the sufïix o( the related généra Pilodius and

Phymodius. Gender masculine.

DlAGNOSlS

Carapace approximately hexagonal, most régions

well-defined. Basal ancennal segment very large,

356

ZOOSYSTEMA • 1999 - 21 (2)

Identity of Chlorodius miliaris A. Milne Edwards, 1873

Fig, 3. — Sulcodius miliaris (A. Milne Edwards, 1873): anterior thoracic slernites and abdomen: A, i 25.7 x 14.5 mm, New

Caledonia (MNHN MP-B 24818); B, ï 15.5 x 9.4 mm. New Caledonia (MNHN MP-B 24817). Scale bars: 3 mm.

anteroexternal angle strongly produced, lilling up

entire space of orbital hiatus; flagellum excluded

frotn orbit, lodged berwcen cleft fbrntcd by outer

lobe of frontal margin and internai supraorbital

toorh. Chelipeds long; nierus very elongate,

extending bcyond edge ol carapace; dorsal sur¬

face of merus deeply grooved longitudinally, wirh

anterior and posterior margins folding medially.

Ambulatory dactylus witb large subdistal ventral

spine which is disrinctly larger than terminal

spine. Sternum relatively broad, stérilités one and

two fused, no suture diÿccrniblc; suture bervveen

sternites two and three very deep: sternite three

narrow, suture betvveen sternites three and four

shalknv but discernible: sternite four wirh dis¬

tinct but shallow médian longitudinal suture;

male abdomen reaching to about one-third length

of sternite four; posterior part of episternite

seven flap-like, overlapping outer anterior edge

of male abdominal segment three when closed.

Telson scmicircular.

Remarks

Sulcodius n. gen. belongs to the Chlorodiinae

because of its relatively broad carapace, non-

projecting front, rounded and spoon-tippcd fin-

gers of the chelipcds, well-developed dactylo-

propodal lock on the ambulatory legs and the

bifid termination of the ambulatory propodus

(Serène 1984).

Forest & Guinot (1961) briefly discussed the

taxonomie position of Chlorodius milidris, noting

that it was easily séparable Irom ail Other

Chlorodiella species by the distinct ornament¬

ation of the carapace and chelipcds. They colise-

quenrly agreed wirh Balss (1938) that C. miliaris

should nor rerained in Chlorodiella, but did not

support the view that the species mighr lie affiliat-

ed with the genus /jtzymodes Heller, 1861. The

genus Zosymodes is presemly classiFied in the

subfamily Zosiminae Alcock, 1898 (fide Serène

1989). Chlorodius miliaris is cettainly not a /osî-

mine as the tips ol the fingers ol its chelipeds are

spooned, the tips of the ambulatory dactyli are

bilurcated and the legs possess rhe dactylo-

propodal lock.

The extcrnal resemblance between Sulcodius

n. gen. and Chlorodiella is superficial, with the

ZOOSYSTEMA • 1999 • 21 (2)

357

Clark P. F. & Ng P. K. L.

Fig. 4. — Sulcodius miliaris (A. Milne Edwards, 1873), left first

pleopod; i 25.7 x 14.5 mm, New Caledonia (MNHN

MP-B 24818); A, ventral view; B. dorsal view. Scale bar;

0.2 mm.

carapace régions of C. miliaris being well-defined

(Fig. 1) (poorjy so in Chlorodiella). and the sur¬

faces of the chelipeds dîstinctly granulated (Fig. 2)

(smooth in Chlorodiella). The general form of

their sternal plastron, abdomen and ambulatory

dactylus however, is radier similar. la the form of

the carapace, Sulcodius n. gen. resembles

Phyrnodius but the anteroexcernal angle of the

basal ancennal segment in Sulcodius n. gen.

extends into the orbital hiatus, filling it entirely.

The sternal plastron of Sulcodius n. gen. is also

distinctly wider than thaï of Phyrnodius. In the

structure of the basal aniennal segment,

Sulcodius n. gen. resembles Pilodius, but the ante-

roexternal angle in Sulcodius n. gen., is more

strongly produced than in any known Pilodius

species (Clark èc Galil 1993; Ng & Yang 1998)

and the form of their sternal plastron differs mar-

kedly (Table 1). The différences are consistent

for a good sériés of C. miliaris examined from

New Caledonia and the lndian Océan, and are

valid for both males and females as well as srnall

Fig. 5. — Sulcodius miliaris (A, Milne Edwards, 1873), left first

pleopod; 7 21.2 x 13.7 mm, Seychelles (NHM 1882.24); A. ven¬

tral view; B. dorsal view. Scale bar: 0.2 mm.

specimens. These features vindicate the establish¬

ment of a separate gen us for Chlorodius miliaris.

I lie extensive prolongation of the basal antennal

segment into the orbital hiatus (Fig. 6) in

Sulcodius n. gen. requires further comment. In

his subfamilial key, Serène (1984: 14, 17) stated

that the basal segment of the antenna in

Chlorodiinae is charactcrised by the absence of an

“external antero-lareral lobule” which is feeble in

Pilodius. But this angle (= lobule of Serène) is

Strongly dcveloped in Sulcodius n. gen. and fi Ils

the entire orbital hiatus. Alrhougli (his charactcr

dues not comply complctely witli the key and

définition of the Chlorodiinae by Serène (1984),

there seerns no justification for establishing a

new subfamily to aceommodate this new genus

and perhaps Pilodius. However, Serènc’s (1984:

17) subfamilial key is confusing concerning the

358

ZOOSYSTEMA • 1999 • 21 (2)

Identity of Chlorodius miliaris A. Milne Edwards, 1873

Fig. 6. — 6 basal anlennal segment and orbit; A, Sulcodius miliaris (A. Milne Edwards. 1873) 22.6 x 13.4 mm. New Caledonia

(MNHN MP-B 24819); B. Pilodius granulatus Stimpson, 1859 11.5 x 6,9 mm, Singapore (ZRC 1999-0073); C, Phymodius ungulatus

(A. Milne Edwards. 1873) 22.7 x 15.3 mm, Cocos-Keeling Islands (ZRC 1965.11.11.16): D, Chlorodiella nigra (Forskâl. 1775)

17.3 xll.O mm. Singapore (ZRC 1999-0074). Scale bars: 1.0 mm.

exclusion of îhe antennal flagellum from the

orbit. In couplet I I, Chlorodiinae, there is a sug¬

gestion tliat the normal situation for tltc généra

other than Pilodius , is tor the antennal flagellum

to be excluded froni the orbit. This opposes the

First couplet to the key to the généra of the

Chlorodiinae, where Serène (1984; 232-233)

uses the antennal flagellum to sépara te Pilodius

Dana, 1851, from Phymodius h, Milne Edwards,

1S63, Tweedieia Ward, 1934, Chlorodiella

Rathbun, 1897, and Liocarpilodes Kluimnger,

1913. He notes that in Pilodius the basal seg¬

ment of the antenna has the antérolatéral exiern-

al angle more or less prolonged and lodged in the

ZOOSYSTEMA • 1999 • 21 (2)

359

Clark P. F. & Ng P. K. L.

Fig. 7. — S fourth left ambulatory dactylus; A, Sulcodius miliaris (A. Milne Edwards, 1873) $ 21.2 x 13.7 mm. Seychelles

(NHM 1882.24); B, Pilodius granulalus Stimpson, 1859 11.5 x 6.9 mm, Sîngapore (ZRC 1999-0073); C, Phymodius ungulatus

(A. Milne Edwards. 1873) 22.7 x 15.3 mm, Cocos- Keeling Islande (ZRC 1965.11 11.16): D. Chlorodiella nigra (Forskâl, 1775) 17.3 x

11.0 mm, Singapote (ZRC 1999 0074). Dactyli for B-D laterally transposed for comparîson wlth left dactylus of A. Scale bars: 1.0

orbital hiatus so lhat this opening can often be

completely closed. Consequently, rhe antennal

flageilum is cxcluded from the orbital hiatus,

l'he remaining four chlorodiine gênera possess

an antennal basal segment wirhout a prolonged

antérolatéral external angle and the orbital hiatus

is open, through whith the antennal flageilum

can be treely directed backvvards into the orbit.

After examination of spécimens of various

Phymodius , Tweedieiu, Chlorodiella and Liocar-

piIodes species, it was this lattet description, as

given in the key to the généra (Serène 1984:

232-233), which proved to be correct.

Sulcodius miliaris

(A. Milne Edwards, 1873) n. comb.

(Figs 1-5; 6A; 7A; 8A, B)

Chlorodius miliaris A. Milne Edwards, 1873: 216,

pl. 8, fig. 3. - Miers 1884: 531. — Guinot 1964:

23, pl. 10, figs 1-2.

Chlorodiella miliaris*. — Forest & Guinot 1961: 96.

Chlorodiella miliaris — Scrcne 1984: 255 (key), 257-

258, pl.XLIllD.

Zoozymodes miliaris — Balss 1938, unpaginated cap-

rion for pl. 2,11g. 2 (part).

non Chlorodius miliaris — De Man 1887: 280 =

Leptodiusoffrions (Rathbun, 1907) ?

non Zoozymodes miliaris — Balss 1938: pl. 2, fig. 2 =

/eptodius rffrirns (Rathbun. 1907) ?

MaTI RIAI. l-.XAMINI-.l). — Nouvelle-Calédonie. Coll.

M. Balansa. 3 9'9 syntypes 10.5-15.0 mm (MNHN-

B 20979). —Nouméa,4. VI. 1989, coll. DAWA, 1 d

22 6 mm (MNHN-B 24819). — Secteur de Nouméa,

stn 1.3, 22°20 S, I66°25’F, 20 ni, coll. B, Ricber de

Forges, 2 6 6 7.2-16.5 mm (MNHN-B 24836), I 9

9.4 mm (MNHN-B 24817). Stn 15. 22‘>20'S,

I66"3I'F., 27 m, 2 6 6 10.0-21.4 mm. I 9 13.3 mm

(MNHN-B 24827). — Stn 251, 22°I9'S, 166°25’E,

20 m. 3 9 9 13.0-17.0 mm (MNHN-B 24829). —

Sm 279, 20 l, 20'S, lôfaNT'E, 29 m, l d 16.5 mm

(MNHM-B 24833). — Baie de Prony, ile Ouen,

stn I 19, 22"28'S, |66"46"E, 20 m, 3 6 6

7.6-14.0 mm. 2 9 9 8.3-1 U.4 mm (MNHN-

B 24820). — Stn 124, 22“31’S, 166"44'F, 18 m,

3 d d 7.5-18.8 mm, I 9 15.4 mm (MNHN-

B 24822, nowZRC 1998,793). — Stn 155, 22 ft 31’S,

166°38’E, 23 in, 6 d d 8.0-12.5 mm, 9 9 9

9.0-15.0 mm (MNHN-B 24831). — Stn 232,

22°35'S, I66°43’E, 28 m. 1 d 14.0 mm (MNHN-

B 24824). — Baie de St Vincent, stn 178, 22"01 ’S,

I66"04'F-, 20 m, I d 25.7 mm (MNIIM-

B 24818). Stn 211, 21“55’S, 165 V 52'L, 12 m, 1 d

13.6 mm (MNHM-B 24825). — Atoll de Surprise,

360

ZOOSYSTEMA • 1999 • 21 (2)

Clark P. F. & Ng P. K. L.

stn 449, 18"22'S. 163°09’E, 21 m, 28.11.1985, I 3

11.9 mm (MNIIN-B 24823), — Grand Récif Sud,

stn 550, 22°59’S, 166"58'E, 24 m, 15.V11.1985,

2 3 3 8.0-16.0 mm, 1 9 8.0 mm (MN11N-

B 24828). — Lagon Nord-Ouest, Stn 920, 20°52.5'S.

164°28.0’E. 10-11 m, 26.IV. 1988. I î 13,5 mm

(MNHN-B 24834). — Stn DW 921, 20“51.2’S,

164°26.6’Li 12-13 m, 27.1V. 1988, 1 3 10.0 mm

(MNHN-B 24826). — Stn DW 922, 20°50.9’S,

164°24.4’E, 12-13 m, 27.1V. 1988, I 3 13.0 mm

(MNHN-B 24821). —.Stn DW 923, 20° 48.~’S,

164° 24.2’E, 9 m, 27.IV. 1988, I 9 8.5 mm

(MNHN-B 2<t832l. — Stn DW 9.39, 20° 36.7’S,

164° 15.5‘E, 12 m, 27.IV. 1988, 1 9 9.7 mm

(MNHN-B 248.30). — Passée de Boulari, 15 m,

20.111.1988, 1 6 8.5 mm (MNHN-B 24835).

Seychelles. 111,1882, c o1 1. H MS A/rrt, 2 3 3

16.2-21.2 mm INHM 1882.24), No. 194, 4-12 fins.

Description oi i argent (25.7 mm) male

SPECIMEN (MNHN-B 24818)

Carapace approximately hexagonal, broader than

long, most régions well-defined, grooves demar-

cating régions deep, not pililerous; 1F deflexed,

separated froni 2F by strongly granulated trans¬

verse rovv of small rounded granules; long soft

setae obscure frontal rnargin; 2F separated front

IM by transverse granulated ridge, which

becomes weak as it reaches orbital région; 2M

separated front 1 M by strong transverse granulat-

cd ridge which joins deep groove separaring ir

Iront 2L; 3M large, well-delined by distinct

grooves ail round, Separated front narrow 4M by

shallow tfa ns verse groove; IB sliglttly swollen; 2B

narrow, entire, but consrricted ntedially to form

waist-like structure; 2L région large, surface

convex; I L and .3L régions narrow, not clearly

demarcated; 4L large, clearly separated from 21.

and 5L by sballow groüves; 5L and 6L well-defi-

ned; I -3R poorly demarcated, surfaces stronglv

granulated, coveretl with long soit setae.

Subhepatic. suborbital, ptervgostomial and

sub-branchial régions granulated; pterygosto-

mial, sub-branchial and postérolatéral régions

densely covcrcd with long soit setae. Frontal

rnargin lamellilorm; projecring ourwards beyond

orbits, rnargin bilobed, initer lobe subtruncate,

outer lobe triangular, tip rounded; internai

supraorbital tooth large, level with frontal nt.tr-

gin, covercd with long soft setae which obscure

outline; supraorbital rnargin with deep médian

cleft; infraorbital rnargin with sharp granules,

entire; external orbital tooth acutely triangular,

well-developed; subheparic région with distinct

tooth visible dorsallv; antérolatéral rnargin gcntly

eonvex; llrst antérolatéral tooth broadly triangu-

Inr; second and thitd antérolatéral teeth acutely

triangular, rip rounded but turned sliglttly ante-

riorly, margins (cspecially anterior ntargins) lined

with granules; postérolatéral rnargin gcntly

sinuous, strongly converging tovvards posterior

carapace rnargin; posterior rnargin ol carapace

gcntly cmrvex posteriorly. Antennules lolding

almost tranversely, lowcr rnargin ol antennulary

fossa and imcrantennulary septum (proepistome)

granulated, Basal antennal segment very large,

granulated, external angle strongly produccd,

filling up entire space ol orbital hiatus; second

and subséquent segments attached to basal seg¬

ment rnedially; peduncle small, subovate; an ren¬

trai flagellant not cntcring orbits, but between

cleft formed by outer lobe ol Irontal rnargin and

internai supraorbital tooth. F.pistome broad,

médian part not depressed; posterior rnargin

strongly sinuous, with well-developed médian

lobe which is separated by a deep médian fissure

and two latéral lobes with convcx margins.

Tltird maxilliped quadrate, surfaces finely granu-

lose; ischium rectangular, separated front basis by

rnedially internipred suture, inner rnargin serra-

ted, médian oblique sulcus deep; merus squarish,

with sLightly produeed, subauficulilorm anterola-

tentl angle; exopod stout, relatively broad, reach-

ing distal érigé ol utérus, with long flagellant;

earpus rounded, dacrylus (wlten opposed against

merus) rcaching beyond lowcr edge of merus.

Chelipeds long, surfaces finely granulose; merus

very elongate, protruding vvell beyond edge of

carapace; merus unarmed, dorsal surface decply

grooved ntedially, lirtcd with dense short setae,

anterior and posterior margins folding ntedially;

earpus with weak tubercle on inner distal angle;

palm slender, elongate, Icit palm distinctlv larger

and more swollen titan right; fingers long, slender,

cutting edges ol larger chela with teeth on proxi¬

mal hall and small dcnticles on distal hall; cut¬

ting edges of smaller chela lined only with

dcnticles, tip distinctlv spoon-shapcd; fingers

pigmented black, pigmentation on pollex

extending somc distance into distal part ol palm

on both inner and outer surfaces.

362

ZOOSYSTEMA • 1999 • 21 (2)

Identity of Chlorodius miliaris A. Milne Hdwards, 1873

First ambulatory leg longest, outer surfaces gent-

ly granulated, margins of ail segments lined with

long soft setae which obscure the outline; merus

nor cristate, dorsal margin gcncly serrated; dorsal

margin of carpus gcntly serrated; dorsal and ven¬

tral margins ol propodus serrated; daoylopropo-

dal articulation very well-developed; dactylus

distinctly curved, outer margin lined with large,

sharp tubercles, inner margin with strtall sharp

granules; subdistal edge of ventral margin with

very large sharp returved spine which is distinct¬

ly larger than distal spine.

Anterior thoracic sternum relativcly broad, cover-

ed with numerous small granules, especially on

anterior half; stérilités one and two fttsed, no

suture disccrniblc; suture between sternitcs rwo

and three very deep, lined by rounded granules;

stérilité three narrow, suture between stcrnites

three and four shallovv but disccrniblc, lined with

irregularly spaced granules, eleft discernible Only

laterally; sternite foui with distinct but shallow

mecfian longitudinal suture; male abdomen

reaching to imaginât) line joining posterior part

of cheliped coxac; posterior part of episternite

seven lamellifprm, fiap-like, forming broad grouve

between it and test of sternum into which latéral

part of abdominal segment three lits.

Abdomen seven-segmented including telson, but

segments three ro five fttsed, immovable, sutures

not obvious except laterally (Fig. 3); segment one

broader laterally, posterior margin distinctly gra¬

nulated; segment two broader, latéral margins

granulated; segment three trapézoïdal, posterior

margin with broad dépréssion; segments four

and five progressively more rectangular; segment

six rectangular, latéral margins distinctly concave;

telson semicircular.

Gonopores coxal. Male first pleopod slender

(Figs 4; 5), broadly C-shapcd, dp directed out-

wards, dorsal part ot subdistal margin with long

plumose setae, test of margin with short, sharp

spines, distal part elongated, tip open. Male

second pleopod relatively short, basal segment

slender, well-developed, distal segment cup-like,

very short, not elongated.

Remarks

The three dried type spécimens in the MNHN

are ail female and dismembered, with only the

carapace and parts of the chelipeds (detached)

intact (l orest & Guinot 1961) The chelae and

meri of the chelipeds, however, are still intact.

These specimens were examincd by the présent

authors and tliere can be little doubt tliat the

présent specimens from New Galedonia and

Indian Océan are probably conspecific with

S. miliaris. The sélection of a lectotype is poiut-

less because ail the type material is in such poor

condition. The figure presented by A. Milne

Edwards (1873) of S. miliaris (as a Chlorodius) is

rather schematic. It does not show the sculpture

of the carapace accurately, nor the groove on the

merus of the cheliped. The figures provided by

Guinot ( 1964) are only of the carapace and palm

of the cheliped. Thercfore the species is redescri-

bed alu! illustrated here in detail on the basis of

the new specimens from the type locality.

Several characters vaty: the strength of the cara¬

pace granules, the degree of spinulation on the

infraorbital margin; and the shape of the distal

part of the male first pleopod. One of rhe two

largest males (MNHN-B 24818) is rather unu-

sual in having sharp granules lining the infra-

orbital margin whereas in most specimens, the

margin is lined with rounded or bluut granules.

This character is, however, not size-xelated as

sonie smaller specimens also have relativcly sharp

granules lining the infraorbital margin. The lar¬

ger chela of the largest male spécimen (MNHN-

B 24818, Fig. 2A) is also unusual in being

distinctly more slender than an cquivalcnc-sized

male (MNHN-B 24819, Fig. 2B) and with a

slightly different dentition on the cutring edges

of the fingers, 1rs chelipedal meri arc also relati¬

vely more slender and slightly longer. In almost

ail other specimens, including lemales and smal¬

ler males, the larger chela is more inflated and

rounded, the basal teeth on the cutting edges of

the fingers are stronger. There are sotne speci¬

mens, however, which have intermédiare chelae

(c.g. male 13.0 mm. MNFIN-B 24827). The

male first pleopod of the Iarge.st male (MNHN-

B 24818, Fig. 4) also has the distal part slightly

more rwisted.

The specimens from the Seychelles reported by

Miers (1884) (NHM 1882.24) are probably

conspecific with S. miliaris. The larger male

(13.7 mm) differs from ail the Pacific specimens

ZOOSYSTEMA • 1999 • 21 (2)

363

Clark P. F. & Ng P. K. L.

in having a more quadrangular and less crans-

versely ovoid carapace, the protûgasrric région is

more flactened, the antérolatéral teeth are more

spiniform, and the Angers of the largcr left chela

are more strongly gaping. Ihe smaller male .spé¬

cimen (9.9 mm), however. agréés well with the

Pacific spécimens in ail major charatrer States.

The distal part of the male firsr pleopod of both

Seychelles specimens appears to be relatively

more slender and sharply tapering compared to

specimens front New Caledonia of comparable

sizes, but this is regarded as intraspecifle varia¬

tion.

Note added in press. — Ng & Yang (1998)

recently described a new genus, Vellodius , for

Pilodius etisoides T.ikcda & Miyaké, 1968, a spe-

cies which was regarded as incerta sedis by Clark

& Galil (1993). Vellndius can be separated (rom

Sulcodius by the amceoexternaJ angle of its basal

anrennal segment not produced and not entering

the orbital hiatus, its proportionately more elon-

gate anterior sternal plastron and rhe absence of a

dorsal groove on the chelipedal merus.

Acknowledgements

We are gratcful to Alain Crosnier (ORSTOM)

for allowing us to study the material of Su/codius,

and providing us benchspace, coffee and valued

discussion in the attic of the MÜSORSTOM

laboratory. We also atknowledge the efforts of

Bertrand Richer de Forges (ORSTOM,

Nouméa) for collectiog and carefully preserving

the xanthoid specimens, as well as meticulously

documcnting the local ity data. Thanks arc also

due to Danièle Guinot (MNHN) for acce.ss to

the types. One of the authors (PPC) was working

under a visiting scientist grant from the MNHN,

Paris.

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364

ZOOSYSTEMA • 1999 • 21 (2)

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ZOOSYSTEMA • 1999 • 21 (2)

365

Description, écologie et chorologie de trois

espèces nouvelles de diplopodes (Myriapoda,

Diplopoda) des zones périphériques du

Massif central et du centre de la France

Jean-Paul MAURIÈS

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

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

collmlll@mnhn.tr

Richard Desmond KIME

Institut royal des Sciences naturelles (Entomologie),

29 rue Vautier, B-1000 Bruxelles (Belgique)

MOTS CLÉS

Diplopoda,

Faune,

taxonomie,

écologie,

France.

KEYWORDS

Diplopodj,

rauna,

taxonomy,

ecology,

France.

Mauriès J.-P. & Kime R. D. 1999 — Description, écologie et chorologie de trois espèces

nouvelles de diplopodes (Myriapoda, Diplopoda) des zones périphériques du Massif central

et du centre de la France. Zoosystema 21 (2) : 367-378.

RÉSUMÉ

Polydesmusgeoffroyi n. sp. (Polydesmida, Polydcsmidae) a été récolté dans des

chênaies du Cher, de la Lozère et de la Dordogne. Les deux autres espèces

sont des craspédosomidcs dont la distribution est nettement plus limitée :

Hylebainosoma (?) nontronensiS n. sp. (Haaseîda) se trouve dans des bois sur

sol acide aux confins du Périgord Vert et du Limousin ; les cinq espèces du

genre Hylebainosoma sont carpathiques ; Pyrgocyphosoma balazuci n. sp. n’est

connu que de deux localités (dont une grorte) de l’Ardèche ; sa découverte

accroît sensiblement vers l’ouest la distribution du genre Pyrgocyphosoma.

ABSTRACT

Description, ecology and distribution of three new species (Myriapoda,

Diplodopa) Jrom the péri pliera! areas of Massif Central and Central France.

Polydesmusgtoffroyi n. sp. (Polydesmida, Polydcsmidae) was coliected in oak-

woods situaced in lhe Cher, Lozère and Dordogne departments. The other

iwo species are craspedosomids with a much more iimired distribution:

Hylebainosoma (?) nontronensis n. sp. (Haaseida) occurs iu woods on acidic

soils in the border régions of tlie- Périgord Vert and the Limousin; the five

species of the genus Hylebainosoma are Carpathian. Pyrgocyphosoma balazuci

is onlv known from two localities, one of which is a cave, iu the Ardèche

department; its discovery sigmficantly extends the distributinnal area ot the

genus Pyrgocyphosoma in a westerly direction.

ZOOSYSTEMA • 1999 • 21 (2)

Mauriès J.-P. & Kime R. D.

INTRODUCTION

La liste des diplopodes de France, dont l’œuvre

posthume de Brolemann, 1935, consacrée aux

seuls craspedosomides/chordeumides, et l’ouvra¬

ge de Démangé (1981), constituent l’ossature, a

fait l’objet plus récemment de mises au point de

Geoffroy (1990a, b, 1991, 1996a, b) et Geoffroy

& Mauriès (1992). Actuellement, cette liste est

riche de 282 taxons, et malgré les lacunes encore

nombreuses concernant certains de nos départe¬

ments, on pouvait penser avoir une connaissance

assez complète de ce groupe sur le territoire fran¬

çais.

Or, il nous est donné de décrire ici trois espèces

nouvelles, qui sont toutes situées géographique¬

ment dans des zones assez peu prospectées de

part et d'autre du centre de la France et du pour¬

tour du Massif central ; elles appartiennent, l'une

aux Polydesmides, les deux autres aux craspédo-

somides {alias chordeumides).

MATÉRIEL ÉTUDIÉ

La première espèce est un polydesme récolté

voici plus de 20 ans dans le Cher par Jean-

Jacques Geoffroy. Sa description a été différée du

fait que le sexe mâle, étant représenté par un seul

individu, on pouvait supposer avoir affaire à un

cas de tératologie gonopodiale. C’est tout récem¬

ment qu'ont été identifiés plusieurs autres spéci¬

mens, dont des mâles, dans du matériel récolté

par l’un de nous (R. D. K) ces dernières années

en Dordogne et en 1981 en Lozère. Ces décou¬

vertes, effectuées en des lieux géographiquement

assez éloignés les uns des autres, confirment par¬

faitement la validité de l’espèce,

La deuxième, découverte par l'un de nous

(R. D. K.) à la faveur de prospections effectuées

dans le cadre de la création du Part régional du

Périgord Vert-Limousin, est une nouvelle espèce

de la famille des 1 iaaseida. L'examen de ses carac¬

tères gonopodiaux montre qu elle n'appartient

manifestement pas au seul genre représenté en

France par une espèce connue surtout de Suisse

et d’Autriche. Haasea fhwescem Latzel, 1884, qui

atteint sa limite occidentale dans les Vosges et le

Jura français. Curieusement, c’est avec les quatre

espèces d’un genre Romunosoma Ceuca, 1967

(non disponible), de Roumanie, que la nouvelle

espèce présente le plus d’affinités. Mais comme

ces Romanosoma peuvent être, sans difficulté,

rapprochés du genre monospécifique

liylebai)\osonul Verhoeff, 1899, c'est à ce dernier

genre que nous la rapportons ici, avec les doutes

afférents à la grande distance géographique.

La description de la troisième espèce, que nous

ne connaissions que par un seul mâle découvert

en 1984 au Mont Mezertç par le regretté Dr

J. Balazuc, est restée elle aussi en attente, pour

deux raisons. D’abord du fait de l’unicité du

mâle, qui a été dans le passé la source de descrip¬

tions abusives. Ensuite parce que l'un de nous

(J.-F. M.) soupçonnait, malgré les dénégations du

collecteur, quelque erreur d'étiquetage, cette

espèce étant alliée au genre majoritairement ita¬

lien Pyrgocyphosuma Verhoeff, 1910. Ht cela

jusqu’à ce que l'un de nous (J.-F. M.), récem¬

ment, découvre dans les collections du MNHN,

un autre mâle récolté naguère par J. Magné dans

une grotte de l’Ardèche. Cette découverte dans

une région géographique proche, efface tout

soupçon d’erreur d étiquetage et vient confirmer

la validité de l’espèce, qui prend place entre les

deux taxons génériques Helvetiosoma Verhoeff,

1910 et Pyryocyphosoma Verhoeff, 1910.

Aujourd’hui, grâce à la récente révision de Spelda

(sous presse), nous considérons qu elle doit

prendre place dans ce dernier genre.

SYSTÉMATIQUE

Ordre POLYDESMIDA Leach, 1815

Famille Poi.YDESMlDAE Leach, 1815

Genre Polydesmus Latreille, 1803

Polydesmus geoffroyi n. sp.

(Fig. lA-D)

MatCrii l.-TVTK. — France, Cher, commune de

Meillam à Samt-Rhomble (Bas-Berry-Baischaut),

forêt de Meillam (Carré UTM DM 67), 30.11 U 976,

coll. J. J. Geoffroy, I d holotype, 1 $ , 1 5 juvénile,

paratypes.

Autre MATÉRIEL. — France. Lozère, Saint-Laurent

de Trêves (4,5 km au sud de Florac), 04.XI.1981,

368

ZOOSYSTEMA • 1999 • 21 (2)

Nouveaux diplopodes de France

Fig. 1. — Polydesmus geoflroyi n. sp. et P. leslaceus Koch ; A, 6 holotype de P. geoffroyi, gonopode gauche, vue médiale : B, le

même, vue latérale externe ; C, gonopode gauche, vue médiale, d'un tf de P. geoffroyi de Lozère ; D, vue ventrale du troisième

anneau d'une ? paratype de P. geoffroyi montrant la base des P.2, les vulves en place, les incisures de l'arc pleurotergal, et la base

des P,3. ; E, incisures de l’arc pleurotergal du troisième anneau, vue ventrale, chez une 9 de P. testaceus ; tt, partie distale du tibio-

tarse ; pf, partie préfémorale, échelles : A-C, 0,5 mm ; D, E, 1 mm.

ZOOSYSTEMA • 1999 • 21 (2)

Mauriès J.-P. & Kime R. D.

Tableau 1 . — Mensurations en mm de Polydesmus geoffroyi n. sp.

6 holotype 6 â Lozère

6 Dordogne

îparatype

longueur

17 15,5-16,5

largeur de la tête

1,63

1,86

longueur antenne

3,20

3,37

largeur du collum

1,34

1,38

largeur du T.Il

1,62

1,88

largeur du T.lll

1.62

2,01

largeur du T.X

2,29

2,54

largeur du T. XVIII

1,32

1,71

longueur pattes moyennes

2,60

3 â <3.1 9, — Dordogne. Nantheuil, vallée de l’îsle,

chênaie sur sol sablonneux, 02.IV, 1997, coll. R. D.

Kime, 1 <3.2 9 9.

Tous ces matériaux sont en dépôt dans la Collection

Myriapodes du VIN H N, JC 008.

ÉTYMOLOGIE. — L'espèce est bien amicalement

dédiée à son collecteur.

Diagnose

Morphologie externe et caractères de la femelle

très proches de ceux de P testaceus C. L. Koch,

1847 mais gonopodes du mâle du type uniramé,

se rapprochant de ceux de P colhtris C. L. Koch,

1847 et de ceux de P. striganovae Golovatch,

1978.

Description

Caractères morphologiques externes

20 anneaux chez les adultes.

La coloration typique des polydesmes éda¬

phiques, d’un brun très clair au rosâtre, n'appa¬

raît que sur les échantillons de Dordogne. Tous

les autres sont d un blanc immaculé, décolorés

par un long séjour dans l'alcool (Tableau 1).

Habitus : sculpture dorsale et forme des carènes

très semblable â celles de P. testaceus. Collum en

ovale transverse. Deuxième tergite subrectangu¬

laire, aux bords latéraux avec seulement des

traces de deux ou trois denticulations. La largeur

des tergites s’accroît progressivement jusqu'au

dixième, puis diminue progressivement jusqu’au

telson. Tergites moyens à bords anréro-latéraux

très régulièrement arrondis, avec seulement des

traces de denticulations. Angle postérieur des

carènes obtus, légèrement saillant vers barrière à

partir du sixième tergite.

Telson sans particularité.

Pattes ambulatoires robustes, élargies surtout au

niveau des préfémurs.

Caractères sexuels du mâle

Gonopodes (Fig. IA-C) du type uniramé simple,

tel qu'il est représenté dans le genre Polydesmus

chez les espèces traditionnellement et artificielle¬

ment groupées sous le nom générique de

Brachydesmus (19 anneaux chez l’adulte) mais

aussi chez, certaines espèces, telles P colhiris , type

de Spanabracbium , qui est l’espèce la plus proche

de la nôtre par ses gonopodes, et dont elle se dis¬

tingue par la plus forte arcanire et la moindre

sveltesse d ensemble, par la plus grande robustesse

de la partie distale du tibiotarse (tt), par l’absence

de dent sur la convexité de l’arcature, et la

moindre épaisseur de la partie préfémorale (pf).

Caractères sexuels de la femelle

Il est impossible de distinguer les femelles de

celles de P. testaceus par les caractères externes.

Les incisures postvulvaires de l’arc ventral pleural

du troisième anneau (Fig. 1D) ressemblent beau¬

coup à celles de P testaceus (Fig. 1E) mais la dent

médiane est plus petite et les lobes latéraux sont

ici réduits à de simples saillies en angle obtus,

comme d’ailleurs chez beaucoup de P. testaceus.

Les coxae des P.2 sont, comme chez P testaceus,

faiblement renflées sur leur face caudale.

Les vulves ne présentent aucune particularité

notable.

370

ZOOSYSTEMA • 1999 • 21 (2)

Nouveaux diplopodes de France

Ordre CRASPEDOSOMATIDA (Gray, 1843)

Famille HaaSËIUAE Attems, 1899

Hylebainosoma (?) nontronensis n. sp.

(Fig. 2A-H)

Matériel-type. — France, Haure-Vienne, Pensol,

Les Simoulies, 19.XI. 1995, coll. R. D. Kime, 1 S

holotype, 2 S d et 1 9 paratvpes.

AUTRE MatéRIKI. — France. Dordogne, Milhac-de-

Nonrron. 02.X. 1995, I 9, 1 9 jeune. — Dordogne,

Saint-Romam-ci-Saint-Cléinent. 17.X.1996. coll.

R. D. Kimc-, 1 6.

Tous ces matériaux, récoltés par l'un des auteurs

(R. D. K.) sont en dépôt dans la Collection

Myriapodes du MNHN (DA 259).

ÉTYMOLOGIE. — Le nom de l’espèce désigne son aire

de distribution : le Nontronais, une région charnière

entre le Limousin et le Périgord Vert.

Diagnose

Aux P.9, colpocoxite robuste presque aussi long

que le télopodite ; aux P.8, pas de véritable « lame

en peigne ».

Description

Caractères morphologiques externes semblables à

ceux de H. flavescens

30 anneaux dans les deux sexes (Tableau 2).

Coloration brun jaunâtre claire.

Antennes grêles, mesurant 1,37 mm de long chez

le mâle holorypc ci 1,44 chez la femelle de

Milhac ; massue anrennaire 4,8 fois plus longue

que large.

Yeux pigmentés à ocelles peu nombreux (voir ci-

dessus) inscrits dans un champ subtriangulaire

ou en croissant, presque toujours disposés sur

sept rangs, même chez la femelle préadulte ; chez

le mâle de Saint-Romain-Saint-Clément, les

14 ocelles ne (orrnent pas de rangs évidents.

Métatergiies à bourrelets dorso-latéraux ovales

transverses, bien individualisés, peu bombés,

situés un peu plus haut que le milieu des lianes,

de part et d’autre d’une zone dorsale assez

arquée ; chaque bourrelet porte sur son pourtour

trois macrochètes rrtétatergaux longs er fins ; ces

derniers peuvent atteindre en longueur les deux

tiers du diamètre vertical d’un anneau moyen ;

l’angle qu’ils forment, de chaque côté, le macro-

chète antéro-externe étant pris comme

« sommet », est un angle obtus ( 120" environ).

La distance qui sépare le macrochère interne de

l’antéro-externe est un peu inférieure au tiers de

celle qui le sépare du sillon dorso-médian.

Caractères sexuels du mâle

Face plane au niveau du labre.

Gonopodes (P.8, Fig. 2A-C) se distinguant par

leur silhouette (en vue orale ou caudale) qui

montre une base élargie supportant deux piliers

(cheirjces) nettement plus grêles ; ces derniers se

caractérisent cri outre par l’absence du grand

« appendice lamellaire denté »parasagittal qui a

été si utilisé (sans doute abusivement) pour dis¬

tinguer les divers taxons décrits dans le genre

Haasea dans |a première moitié de ce siècle. Il est

remplacé ici par une lame beaucoup plus discrète

(d, Fig, 2C), plus ou moins fondue au chçiritc et

dont la partie apicale est acuminée et bordée de

fines spinulations ; le cheirite lui-même ne se

divise que distalement en trois branches d’impor¬

tance inégale, deux postérieures fines er subégales

(b et c) et une antérieure (a), plus longue er plus

épaisse, dirigée vers l’avant.

Aux paragonopodés (P.9, Fig. 2D), chaque télo¬

podite est constitué, comme chez les autres

espèces, par un grand préfémur (p) ici large et

haut, peu rétréci à la base, surmonté d’un moi¬

gnon lélopudial (r, ici nettement plus large que

long) et flanqué intérieurement d’un coxite (x).

Ce dernier, bien que plus petit que le préfémur,

est d’une taille plus grande que chez les autres

espèces du genre ; mais sa structure est plus

simple, en bâtonnet robuste et arqué différen¬

ciant subapicalement et caudalement une lamelle

rétroarquée.

Les P. 10 (Fig, 2E) ourla structure caractéristique

des autres espèces du genre, avec un coxite (x)

très volumineux abritant une grosse glande coxale

et un télopodite peu modifié de cinq articles.

Les P. 11 et autres pattes postgonopodiales ne pré¬

sentent aucune particularité (absence de glandes

coxales).

Caractères sexuels de la femelle

P.2 (Fig. 2F) à tarses garnis d’un rang de plus

d’une quinzaine de soies rigides subégales.

Vulves (Fig. 2F, H) ne se montrant pas très diffé-

ZOOSYSTEMA • 1999 • ZI (Z)

371

Nouveaux diplopodes de France

Tableau 2. - Mensurations en mm, nombre d'anneaux et d'ocelles d 'Hylebainosoma (?) nontronensi n. sp- ’ Anneaux anormale¬

ment télescopés par la tixatlon. St R, et St Cl.. Saint Romam-et-Saînt-Clément.

Nombre

d’anneaux

Longueur

Largeur

Diamètre

vertical

Longueur

antenne

L. patte

moyenne

Nombre d’ocelles :

total et (par rang)

<S holotype

8,4

0,70

1,37

1,2

13 (1123321)

d paratype

8,5

0,91

0.69

14(1122332)

d idem

7,8

0,78

0.66

12(1123221)

9 idem

7.3*

0,89

0.72

13 (1123321)

2 Milhac

9,6

1,07

0,86

1,44

0,96

13(1123321)

2 juv. idem

0,77

0,64

13 (1123321)

o* St R. et St Cl.

6.3*

0,77

0,66

14 ( rangs?)

rentes, par leur forme générale, de celles obser¬

vées sur quelques-unes des autres espèces du

genre (Kurnik 1986) ; l’opercule (op) est bien

développé, mais les valves sont plus symétriques

et le lien caudal entre les deux bourses est ici

réduit à un simple bandeau.

P.3 (Fig. 2G) constituées d’un sternite un peu

diminué en hauteur portant une paire de

membres réduits constitués chacun d’un préfé-

mur troncônique surmonté d’un reste de télopo-

dite en petit bourgeon.

Remarque

Si les caractères externes sont semblables, pat

contre, tant par la structure simplifiée des gono-

podes (et notamment I absence complète de la

lame dentée, <■ kératite » des auteurs germano¬

phones) que par le relatif grand développement

du coxite des paragonopodes, notre espèce se

sépare radicalement du genre Haaseû., seul genre

de la famille représenté en France. Ces caractères

gonopodiaux la rapprochent étrangement des

quatre espèces de Roumanie qui constituent le

genre (non valide par défaut de désignation

d’une espèce-type) Romanosoma Ceuca, 1967.

Or, par la structure de leurs gonopodes anté¬

rieurs, ces dernières espèces sont manifestement

affines du genre monospécifique Hylebainosoma

Verhoeff, 1899, confiné à la Slovaquie (Schmecks)

et le Nord de la Hongrie (Velsic, Komitat Bars

Josvajo, Kom. Borsod). C’est donc manifeste¬

ment au voisinage de ce dernier genre, qui cons¬

titue, avec Xylopbageuma Verhoeff, 1911, la tribu

des Hylebainosomini, que se place la nouvelle

espèce française, que nous classons ici, avec

quelques doutes, dans le genre Hylebainosoma ,

qu’une révision sérieuse de la famille devrait enri¬

chir des quatre espèces roumaines du genre à

valider.

Famille CraSPEDOSOMATIDAE Gray, 1843

Genre Pyrgocyphosoma Verhoeff, 1910

Pyrgocyphosoma balazuci n. sp.

(Fig. 3A-E)

MATÉRIEL- TYPE. — France, Ardèche, Borée, Mont

Mézenc, flanc est, 01.VI. 1984, coll. J. Balazuc, 1 â

holotype.

AUTRE MATERIEL. — France, Ardèche, Saint-Paul-le-

Jeune (canton des Vans), Goule de Sauvas,

06.VI11.1965,coll. Magné. 1 d, I 2.

Ces matériaux sont eu dépôt dans la Collection

Myriapodes du MNHN (DA 258).

ÉTYMOLOGIE, — l 'espèce est dédiée à la mémoire de

son collecteur, le regretté Dr Jean Balazuc, spécialiste

bien connu de la biospéoiogie du département de

l’Ardèche.

Diagnose

Diffère de tous ses congénères par la présence,

sur la lace interne des cheirites des gonopodes du

mâle, en position subapicale, d’un long appendi¬

ce rérroarqué plus large distalement que basale-

ment.

Description

Caractères morphologiques externes

30 anneaux dans les deux sexes.

ZOOSYSTEMA • 1999 • 21 (2)

373

Mauriès J.-P. & Kime R. D.

Fig. 3. — Pyrgocyphosoma balazuci n. sp. ; A. moitié gauche du septième anneau du S holotype, vue ventrale, montrant le décou¬

page du bord ventral du pleurotergite : B, P,3 (gonopodes antérieurs) du même d, vue caudale (le chelrite droit a été extrait) ;

C, cheirite droit isolé, extrait du précédent ; □, P.9 (paragonopodes ou podosternite) du d holotype, vue orale . E, vue distale de la

vulve droite d'une i paratype. Échelles : A, 0,5 mm , B-E. 0.2 mm.

Longueur du mâle holotype : 15 mm ; diamètre

vertical tnax. ; 1,2 mm ; largeur max. : 1,4 mm.

Coloration blanchâtre, du fait de l'absence

presque complète de pigmentation, excepté

quelques traces entre les yeux (qui sont noirs),

sur les antennes et sur les flancs.

Antennes plurôt grêles, un peu plus longues que

la largeur maximale du corps (1,75 mm chez le

mâle holotype).

Yeux triangulaires pigmentés, composés d une

vingtaine d’ocelles disposés en neul rangs (1,2,

3, 4, 5, 3, 2).

Habitus de CritspedosomU , avec boursouflures

mératergales médiocrement développées et non

coupées par un quelconque sillon. Les 3 + 3 soies

mctatergales sont très courtes et fines ; leur lon¬

gueur n'excède pas 0,15 mm sur les anneaux

moyens et postérieurs, 0,20 mm sur les anneaux

374

ZOOSYSTEMA • 1999 • 21 (2)

Nouveaux diplopodes de France

antérieurs ; I angle qu elles forment, de chaque

côté, la soie antéro-externe étant prise comme

«sommet », est un angle obtus (120° environ).

La soie interne est un peu plus prés du sillon

dorso-médian que de la soie antéro-externe,

Pattes assez longues et robustes mesurant en

moyenne 1,8 mm.

Caractères sexuels du mâle

Face concave.

Septième anneau (Fig 3A) à peine plus renflé

que les autres anneaux, présentant de part et

d'autre un lobe pleural en triangle émoussé plus

proche du type Pyrgocyphosorna que du type

Helvetiosoma.

Gonopodes (Fig. 3B, C) : cheirite (K) présentant

la particularité de posséda en même temps ;

- à la place de la grande dent caudale subapicale

qui existe chez plusieurs taxons italiens du genre

Pyrgocyphosorna (tels P. armigerum Verhoeff, 1925

et P dentatmn Broelemaun, 1892 mais aussi

P. dalmazzense Verhoeff, 1930, P. brunatense

Verhoeff, 1910 et P. fhrentinum Silvestri, 1903),

se trouve un long appendice interne rétroarqué

(k) qui s’élargit disralement et dont l'apex est

découpé en 3 + 2 dents ;

- dans la concavité de l’arcarure du bord antéro-

interne, au lieu d’une forte déni triangulaire

orale qui existe chez trois taxons du nord de

l’Italie (P. renanum Verhoeff, 1932, P. roccavio-

nense Verhoeff, 1937 et P serravallense Verhoeff,

1936), mais aussi en plus petit chez P brunatense

Verhoeff, 1910 et serpentinum Verhoeff, 1932 se

trouve un lobule subquadrangulaire (j) bidenté ;

- plus basalement, se trouve une dent simple (i),

située sur le bord postérieur interne, qui n'a

d'équivalent chez aucun autre Laxon du genre.

Coxites (X) du type tout à fait classique de la

famille. Les tigelles paramédianes sont ici courtes

et fines, érigées, acuminées, leur longueur d'envi¬

ron le tiers de la longueur du coxite lui-même.

Paragonopodes (Fig. 3D) présentant les deux

lobes latéraux postérieurs subquadrangulaires (L)

des Helvetiosoma. mais pouvant être comparés à

ceux de Pyrgocyphosorna dentatmn par leur forme

générale. Ils se distinguent de ces derniers par le

processus oral médian M, il surface granuleuse,

qui est nettement plus long et plus anguleux api-

calement, dépassant très nettement les ressauts

(m) (supposés abriter les débouchés de glandes

coxales) qui se trouvent sur sa face caudale.

Caractères sexuels de la femelle

La seule femelle connue ne montre aucune parti¬

cularité morphologique.

Les vulves (Fig. 3E) sont pratiquement iden¬

tiques à celles de Pyrgocyphosorna dorme, figurées

par Brolcmann, 1935 ; elles ont la même forme

générale que chez P dentatmn, P dalmazzense et

P. savonense, dont elles ne se distinguent que par

d’infimes détails (Kurnik 1986).

ÉCOLOGIE DES TROIS ESPÈCES

L’apparente rareté de ces trois espèces est due en

partie au manque de prospections, mais peut-être

aussi à leur écologie.

Nous ne savons rien de Pyrgocyphosorna balazuci,

sinon qu'elle a été rencontrée seulement en

Ardèche, dans deux localités, une cavernicole et

une édaphique, Pour cetre dernière, nous ne

savons pas si le Dr Balazuc l’avait récoltée sur les

parties somitales, découvertes, ou dans les parties

basses, boisées, du flanc est du Mont Mézenc,

Hylebatnosoma nontronensis , dont nous avons

souligné plus haut l’originalité de sa situation du

point de vue taxonomique a, comme la précé¬

dente, une aire de répartition restreinte, à cheval

sur les deux départements de la Dordogne et de

la Haute-Vienne, là où se rencontrent le Périgord

Vert et le Limousin. Du point de vue écologique,

elle semble inféodée aux bois et forets sur sol

acide de la zone des terrains métamorphiques et

volcaniques du Massif central, région considérée

généralement comme continentale plutôt

qu’atlantique. Les localités connues présentent les

caractéristiques suivantes :

- Pensol : bois de châtaignier ( Castanea saliva),

chêne (Qutrcus), bouleau ( Betula ) et pin (Pinus),

sur sol acide à nioder (substrat cristallin : embré-

chites à deux micas) ;

- Milhac de Nomroti : bois de pin et châtaignier

avec fougères (Pteridium), sur sol acide à moder

(substrat : altérités colluvionnécs) ;

- Saint-Romain-ct-Saint-Clémcnt : même végé¬

tation que la précédente, sur sol acide (substrat :

granité et micaschiste) ;

ZOOSYSTEMA • 1999 • 21 (2)

375

Mauriès J.-P. & Kime R. D.

Ces caractéristiques écologiques entrent curieuse¬

ment en corrélation avec la taxonomie, suggérant

des ressemblances, voire des liens, avec certains

éléments de la faune alpine.

Polydesmus gcojjroyi a été récoltée en trois locali¬

tés, toujours en forêt, dans la litière et/ou le bois

mort des chênaies.

La forêt de Meillant, près de Saint-Amand-

Montrond dans le Cher, est une chênaie (Quercus

robur) mélangée à du hêtre (Fagus siluatica ), du

charme (Carpinus becula) et des genévriers

(Juniperus commutiis) ; elle est située dans une

zone faiblement karstique, sur calcaire Bajocien-

Bathonien et présente un sol brun à mull

(Geoffroy, ni lin).

La chênaie de S’aiut-Laurenr-de-Trèves, au sud de

Florac en Lozère, est située sur une forte pente

dans la vallée du Tarnon, sur la bordure orientale

du Causse Méjean (calcaire jurassique karstique).

Celle de la commune de Nantheuil, près de

Thiviers en Dordogne est, au contraire, située sur

un substrat métamorphique, dans la zone vulca-

no-détritique du Bas-Limousin ; le sol à mull est

léger, bien drainé et situé bien au-dessus de la

nappe phréatique.

CHOROLOG1E

La faune diplopodologique des zones périphé¬

riques du Massif central et du Centre de la

France est très mal connue, du fait du manque

de prospections, qui affecte aussi le Massif cen¬

tral lui-même. Il n’est donc pas très surprenant

d’avoir découvert trois espèces nouvelles dans

cette région. L’avenir nous dira si elles ont une

aire de répartition plus vaste que celle que nous

connaissons aujourd’hui.

Concernant Polydesmus geoffi-oyi, qui présente la

distribution la plus étendue et la plus disjointe

des trois, on peut prévoir que sa répartition est

certainement plus vaste, compte tenu également

de sa plus grande tolérance écologique.

Quant aux deux autres espèces, elles semblent

avoir une répartition plus restreinte, probable¬

ment endémiques dans leurs zones, ce que corro¬

bore leur moindre tolérance écologique et en

outre, elles sont éloignées géographiquement des

autres espèces de leurs genres respectifs, qui sont

l'un sud-alpin, l’autre carpathiquc.

Le genre Pyrgocyphosoma est centré sur l’Italie

septentrionale ; il est riche en espèces et chacune

d’elle a une très petite aire de répartition. Le seul

représentant français connu jusqu'à ce jour se

trouvant à la frontière franco-italienne, dans les

Alpes-Maritimes, la nouvelle espèce, sans doute

témoin d’une aire autrefois plus vaste, agrandit

notablement f aire de distribution du genre.

Le cas de Hy le b a i no sam a rwntronemis est plus

intéressant encore, ses plus proches congénères

ayant une distribution carpathique. Son existence

dans le futur Parc régional du Périgord Vert-

Limousin est pour le moins remarquable. Si l’on

note que le substrat est granitique ou cristallin et

que par ailleurs Tajovsky (1997) a capturé

Hylebainosonui tatmnum Verhoeff, 1899 sur sub¬

strat granitique et siliceux en forêt de conifères

en Slovaquie, on ne peut qu’être frappé par la

convergence des données écologiques et taxono¬

miques. D ailleurs, dans le cadre de prospections

effectuées par l’un de nous (R. D. K.) en

Périgord et Limousin et suite à deux transects

effectués dans le Massif central en automne

1997, il est apparu avec netteté que le Massif

centra! abrite d’autres espèces liées à la faune

alpine/périalpine. Ainsi, Orthorbordeumell/ipalli-

dti Rothenbiihler, 1899 a été signalé des départe¬

ments du Puy-de-Dôme, de Corrèze et de

Haute-Vienne, tandis qu’ Hdvetiosom/i nrvemum

(Ribauc in Brolemann, 1932) parvient jusqu’en

Dordogne. Cette région de France sc trouve à la

limite occidentale de l’une des grandes zones bio¬

géographiques du continent européen, la Zone

centrale qui couvre (Scanners & Bourdeau 1995)

le Massif central, le nord et le nord-ouest des

Alpes (Ardennes, Vosges, Jura, Allemagne,

Pologne) puis, par la Tchéquie et la Slovaquie,

descend vers le sud, couvrant la péninsule Balka¬

nique (Roumanie, Hongrie, ex Yougoslavie,

Bulgarie) jusqu’à I extrême nord de la Grèce.

Les auteurs de cette modeste contribution à la

connaissance de la faune de France espèrent que

ces découvertes seront un encouragement à pros¬

pecter dans les zones réputées o priori er à tort

non intéressantes, et qui sont encore très nom¬

breuses sur l’Hexagone.

376

ZOOSYSTEMA • 1999 • 21 (2)

Nouveaux diplopodes de France

Remerciements

À Mme Michèle Bertoncini (CNRS, MNHN),

auteur patient et minutieux de l'iconographie de

cette note. A Jean-Jacques Geoffroy pour les ren¬

seignements d'ordre géographique et écologique

qu’il nous a obligeamment fournis.

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lion de la biodiversité des diplopodes de France.

Biosystema 14 : 65-72.

Geoffroy |. |. & Mauriés J.-P. 1992.— Les diplo¬

podes édaphiques et souterrains de France : don¬

nées récentes, répartition des espèces nouvellement

décrites et peu connues. Mémoires de Biospéologie

19: 127-133.

Golovatch S. 1. 1978. — New and poorly known spe-

cies of the Diplopoda front the European part of

lhe USSR. Zoolngitcheskii ZhttmalS.S.S.R., Moscou

57 (3) : 453-457.

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in The Cvelopedia of Anatomy and Physiology,

Volume 3. Todd Ed., London.

Kurnik 1. 1986. — Zitr Taxonomie ostalpiner

Chordeumatida: vulvennwrphologte and idenlifrka-

tion (1er Wethchen. Dissertation rut 1 rlangung des

Dnktorgrades an der Naturwissenschaftlichcn

Fakultat der Universirât Iniisbruck, Innsbruck

(Austria), 105 p.

Latzel R. 1884. — Die Myriopoden der

Ôslerrciçhisçh- Ungartstben Monarchie. //. Die

Symphykn, Panropoden and Diplopoden. Alfred

Kbldcr, VVicn, 413 p.

Loksa I, 1962. — E litige noue und wenig belcj tinte

Diplopoden aux Ung-arn Annales Unioersitatis

Sdentarium Budapestensis de Rolando Eôtvôs nomi¬

nale, Sectio Biologica 5 -.157-170.

Rorhertbühler I L 1899. — Ein Bcitrag zur Kenntnis

der Myriapodenfauna der Schweiz. Revue suisse de

Zoologie, Genève6 : 199-271.

Sîlvestri F. 1903 in Berlese A. - Acitrl, Myriapoda et

Scorpioncs. Hurusque in ltalia Repenti, fasc. 100,

n° 9, Paduva.

Spelda f. sous presse. — The gémis Pyrgocyphosoma

Vcrhocff, 1910 (Diplopoda, Chordeumatida,

L raspedosomandae): new aspects on systenuuics,

distribution and ecology. Arthropode) Selecta,

Moscou.

Stanncrs D, & Bourdeau P. 1995. — Furopçs envi¬

ronment. the Dobris A su, s me nr. F.uropean

Environment Agency, Copenhague, 676 p.

Tajovsky K. 1997. Distribution ofthe tnillipedes

(Diplopoda) nlong an altirudinal gradient in three

mountain régions (Czech Republie and Slnvak

Republic), in Many legged animais — A collection

of papeis on Myriapoda and Onychophoia.

F.nghoff F.d.. Proceeding of dte IOth international

Congress ol Myriapodology, Copcnhagen, 29 Jul.-

2 Aug. 1996. Fntomo/ogica Scandinuvica suppl. 51:

225-234.

Verhoeff K. W. 1899. — Beitrage zur Kenntnis

palaarkttsclier Myriopoden. VIII. Aufsatz. Archiv

fitr Naturgeschlchtc 1 (2): 95-154.

Verhoefl K. W 1910- — Über Diplopoden, 11-15.

(31-35) Aufsatz: Beitrage zur Kenntnis der

Clomeriden, luliden, Ascospermorpha und

I vsiopetaliden, sowie zur Fauna Siziliens. Nova

Àrta, Halle 92 (2) : 139-448.

Verhoeff K. W. 1911. — Xylophagetima, eitte lieue

Gattung der Orohainosomidae. Liber Diplopoden,

45. Aufsatz. Zuologhcher Anzeiger, Leipzig 38 (7/8):

193-208.

VerhoefFK. W- 1925. — Ncue Diplopoden-Beitrâge.

95. Diplopnden-Aufsatz, Zoologishe Jahrbücher,

Jena 50: 61-122.

Verhoeff K. W. 1930. — Über Diplopoden aus

Italien, namentlich Piémont. 114. Diplopoden-

ZOOSYSTEMA • 1999 • 21 (2)

377

Mauriès J.-P. & Kime R. D.

Aufsatz. Zoologische Jahrbücher, Jena 59: 387-446

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den ôstlichen Apenninen. 125. Diplopoden-

Aufsatz. Zoologische Jahrbücher, Jena 63 (3):

300-351.

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Nordwestitalien und über einige andere mediter-

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Zoologische Jahrbücher, Jena 68 (4/5): 353-444.

Verhoeff K. W. 1937. — Studien in Nordwestitalien

1929-1933. 129. Diplopoden-Aufsatz. Zoologische

Jahrbücher, Jena 70 (1/2): 87-170.

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Akademie der Wissenschaften 60: 225-242.

Soumis le 8 juin 1998 ;

accepté le 14 septembre 1998.

378

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux des collections du

Muséum national d’Histoire naturelle de Paris.

6. Coucous (Cuculidae)

Claire VOISIN & Jean-François VOISIN

Laboratoire de Zoologie (Mammifères et Oiseaux), Muséum national d'Histoire naturelle

55 rue de Buffon, F-75321 Paris cedex 05 (France)

Voisin C. & Voisin J.-F. 1999. — Liste des types d’oiseaux des collections du Muséum

national d'Histoire naturelle de Paris. 6. Coucous (Cuculidae). Zoosystema 21 (2) : 379-

402.

MOTS CLÉS

oiseaux,

r VS

Cuculidae,

Muséum national d’Histoire

naturelle.

RÉSUMÉ

Les types de quarante et un taxons de Coucous (famille des Cuculidae) sont

passés en revue de façon critique. Nous avons désigné des lecrotvpes pour les

taxons suivants : Cumins pyrrbophanus Vieillot, 1817 ; Cuculus nigrorufits

Cuvier, 1817 ; Cuculus variegatus Vieillot, 1817 ; Melias dinrdi Lesson, 1831;

Cultrides pucheramt Deville, 1851; Carpococcyx renauldi Oustalet:, 1896;

Bubutus isidori Lesson, 1831 ; Bubutus duvaucelit Lesson, 1831 ; CocCyccus

erythrorhynchus Cuvier in Lesson, 1831 ; Coua cursor Grandidier, 1867 ;

Coiut verreauxi Grandidier, 1867 ; Coua coquireli Grandidier, 1867; Coua

cristata pyropyga Grandidier, 1867 ; Coua reynaudii Pucheran, 1845 ; Coua

serriand Pucheran, 1 845 ; Centre pus aterdlbus Lesson, 1 826 ; Cen trapus cele-

bensis Quoy & Gaimard, 1830 ; Centrapus savorgnani Oustalet, 1886 ;

Centropus lafresnayanus Verreaux in Vinson, 1865 ; Centropus pumilus

Lesson, 1831 ; Crotopbaga casasii Lesson, 1828. Vingt spécimens, portant la

mention « type » mais qui n’en sont pas au sens du Code de Nomenclature

Zoologique, édition de 1985, ont aussi été examinés. Cuculus lepidus

S. Mülier, 1845, est un synonyme plus récent de Cuculus linealus Lesson,

1831.

ZOOSYSTEMA • 1999 • 21 (2)

379

Voisin C. & Voisin J.-F.

KEYWORDS

birds.

types,

Cuculidae,

Muséum national d’Histoire

naturelle.

ABSTRACT

List of the types uf birds in tbe collections of the Muséum national il’Histoire

naturelle of Paris. 6. Cuckoos (fitmily Cuculidae)

The types of fourty one cuckoo (fini il y Cuculidae) taxa are reviewed cricical-

ly. Lectotvpes are destgnated for the following taxa : Cuculus pyrrhophdnus

Vieillot. 1817; Cuculus nigrantfits Cuvier, 1817; Cuculus mriegatus Vieillot,

1817; Melias diardi Lesson, 1831; Cuit rida pucheranti Deville, 1851;

Carpococcyx renauldt Ouscalet, 1896; Bubutus tsidori Lesson, 18.31; Htibutus

duvaueelii Lesson, 1831; Cmcyccus erythrurhynchus Cuvier in Lesson, 1831;

Coua cursor Gr.indtdier, 1867; Cotta venrauxi Ctandidier, 1867; Coua coque-

reli Ctandidier, 1867; (loua cristata pyrvpyga Crandidier, 1867; Coua reynau-

dii Pucheran. 184 5; Coua serriaua Pucheran, 1835: Centra pu s atentlbiis

Lesson, 1820; Ce)t trapus celebetlsis Quoy Gaimard, 1830; Cmtropns sttvor-

gnani Oustalet. 1886; Centropus la/resnayantts Verreaux in Vinson, 1865;

Centropm pumîlm Lesson, 1831; Crotophagp casasii Lesson. 1828. 1 wcnty

additional spécimens, labcllcd as types ", but which tliey are mit according to

the meaning of type in the Code of Zoological Nomenclature, 1985 édition,

are also exantined. Cuculus lepidus S. Müller, 1845, is a more recent synonym

ol Cuculus linedtus Lesson, 1831.

INTRODUCTION

Ce travail fait suite à cinq articles précédents

concernant les types d'oiseaux du Muséum natio¬

nal d’Histoire naturelle (J.-F. Voisin 1992, 1995 ;

C. Voisin 1993 ; Voisin & Voisin 1996 ; Voisin

et al. 1997) et suit les mêmes conventions.

Pour chaque spécimen, nous avons noté successi¬

vement :

— le nom sous lequel il a été décrit originalement

(en caractères gras) ;

— la catégorie de type à laquelle il appartient ;

— la référence de la description ;

— le nom qu’il porte usuellement dans la nomen¬

clature actuelle (en italiques ordinaires) ;

— le numéro du spécimen au Catalogue général,

qui seul fait foi actuellement :

— les numéros du spécimen dans I Ancien et le

Nouveau Catalogues (A.C. ; N.C.).

Les mentions qui se trouvent sous le socle et sur

les étiquettes ont toutes été recopiées en respec¬

tant l’orthographe d’origine. Les retours à la

ligne sont indiqués par le symbole /, le passage

du recto au verso d une étiquette par le signe III.

Un grand nombre de socles ont été refaits à la fin

du XIX e siècle, une rénovation qui a particulière¬

ment touché les Cuculiformes. Les mentions des

anciens socles ont alors été recopiées sur les nou¬

veaux, sans naturellement que nous ayions de

garantie qu'il n'y ait eu ni omission ni erreur. Ces

socles refaits se reconnaissent à la meilleure qualité

de leur bois, à la disposition assez homogène de

leurs inscriptions, ainsi qu'au fait que ces der¬

nières sont toutes de la même main sur le même

socle. Contrairement à ce que nous avons conti¬

nué de faire pour les anciens, nous avons recopié

les inscriptions de ces nouveaux socles à la file, en

partant du haut, sans nous soucier d'y distinguer

des groupes, fous les spécimens étudiés ont été

munis d’une étiquette rouge de type qui reprend

les principales mentions des autres étiquettes

et/ou du socle. Sauf mention contraire, toutes les

autres étiquettes sont blanches.

On trouvera une note sur les différents cata¬

logues autrefois ou toujours en usage au

Laboratoire des Mammifères et Oiseaux du

Muséum dans Voisin et al. (1997). Le Nouveau

Catalogue des « Coucous » a été établi à la fin du

siècle dernier, sans esprit critique, en particulier

pour les indications de ty pes. Quelques annota¬

tions et ajouts y ont été portés ultérieurement par

le Professeur J. Berlioz. Pour garder un lien entre

380

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

les spécimens démontés et leur socle, le taxider¬

miste qui a procédé aux opérations de démontage

a attaché à l’une de leur pattes une étiquette por¬

tant un numéro qui correspond normalement à

leur numéro du Nouveau Catalogue.

Lorsque nous parlerons du « Code » dans les

lignes qui suivent, il s’agira uniquement du Code

International de Nomenclature Zoologique, a ter

(1985).

Les auteurs du XIX e siècle décrivaient dans leurs

ouvrages toutes les espèces connues de leur

temps, en plus de celles qu'ils considéraient

comme nouvelles. Lorsqu'ils traitaient d'une

espèce déjà connue, ils la décrivaient souvent

d’après des spécimens qu’ils avaient sous les yeux,

et en donnaient une liste des descriptions anté¬

rieures. Même s’ils portent la mention « type »,

ces spécimens n'en sont évidemment pas au sens

du Code, et ils seront examinés à la fin de cet

article. De son côté. G. Cuvier avait l’habitude

de nommer et de décrire oralement des spéci

mens d’espèces qu’il estimait nouvelles, entre

autres pendant ses cours. Ses disciples notaient

ces descriptions et les reprenaient dans leurs

écrits ultérieurs. Afin de respecter la paternité de

ces dénominations, les élèves de Cuvier les fai¬

saient alors suivre de la mention « Cuv. » ou

« Cuvier ». On peut donc leur appliquer les dis¬

positions de l'article 50a du Code en les faisant

suivre de mentions telles que : » Cuvier in

Lesson » ou » Cuvier in Dumont ». Dans un but

de simplification, nous parlerons dans les lignes

qui suivent de « désignation implicite » lorsque

fauteur d'un raxon indique qu’il a examiné, en

plus de l’holorype, un certain nombre de spéci¬

mens, sans toutefois les désigner comme para-

types, ce qu’ils sont néanmoins (articles 72a iii et

72b v du Code).

CATALOGUE

Types de taxons dont la dénomination

EST ENCORE EN VIGUEUR

Cucui.us Sonneratii Latham, 1790. Holotype

PAR MONOTYPIE.

Latham, Index Ornithologicus sive Systema

Ornithologiac I : 215.

Penthoceryx sonneratii sonneratii (Latham,

1790).

C.G. ; 1997-1213.

Autres numéros de catalogues : A.C. n° 1961.

Inscriptions sous le socle :

Çacomantis / sonneratî / (Laili.) / Sonnerat / Inde

/ 5 [même écriture].

Le bec ainsi que les rémiges primaires et secon¬

daires du côté gauche manquent sur ce spécimen

par ailleurs en bon état Sonnerat (1782), qui tra¬

vaillait au Jardin du Roi, ancêtre du Muséum à

l’époque de Buffon, décrivit cette espèce le pre¬

mier et la nomma «« le Petit Coucou des Indes ».

Latham (1790) lui a donné son nom latin en se

référant à Sonnerat comme le montre la mention

« Petit Coucou des Indes, Sonn, voy. » qui se

trouve après la description latine. Latham a donc

utilisé l’ouvrage de Sonnerat pour sa description

et le type de Latham est le même que le spéci¬

men décrit par Sonnerat.

Cuculus pyrrhophanus Vieillot, 1817.

Lectotype, présente désignation.

Vieillot, Nouveau Dictionnaire d’Histoire

Naturelle 8 : 234.

Çacomantis pyrrhophanus pyrrhophanus (Vieillot,

1817 ).

C.G. : 1997-1225.

Autres numéros de catalogues : A.C. n° 1964,

N.C. n° 98.

Inscriptions sous le socle :

1. N' 1, Calédonie / M. Labillardière.

2. Only type of pyrrholoplius / 30/9/25 MA I .

3. 1964 / Çacomantis pyrrophanes / V / C. bronzi¬

ons Gr. / Type [même écriture ancienne que le I.].

4. wrong / Gr. M. 30.9.25 [même écriture mal¬

adroite que le 2, mais au crayon].

Étiquette du socle :

Çacomantis pyrrholoplius / (V.) / C. bronzinus

(Gr.) / Labillardière / N. Calédonie.

VieiIlot (1817) décrit successivement les plu¬

mages du mâle, de la femelle et du jeune. JJ avait

donc au moins trois spécimens de cette espèce

devant lui ; seul celui-ci existe encore dans nos

collections. Le taxon Çacomantis pyrrholophus

porté sur l’étiquette du socle n'existe pas. Cette

mention a probablement abusé le scripteur

ZOOSYSTEMA • 1999 • 21 (2)

381

Voisin C. & Voisin J.-F.

anglophone qui l’a reportée sous le socle et a cor¬

rigé cette erreur le jour même. Ce scripteur, qui

signe MAT ou bien Gr. M, doit être Gregory M.

Mathews, qui était en Europe en 1925, à en

juger par les comptes rendus des séances du

British Ornithologists Club de cette année-là.

Labillardièrc, de son vrai nom J. J. de La

Billardière, était médecin et un des naturalistes

de l'expédition d'Entrccasteaux, partie de 1791 à

1794 à la recherche de l’expédition de La Pérouse

disparue en mer. Ce spécimen a été collecté au

havre de Balab, en Nouvelle Calédonie, et non

en « Nouvelle Hollande » (Australie) comme le

pense Vieillot (Srresemann 1953). Les erreurs

sont d’ailleurs fréquentes dans les localités d’ori¬

gine attribuées aux spécimens rapportés par

l’expédition d’Entrccasteaux (Stresemann 1953 ;

Voisin et al. 1997).

Melias Diardi Lesson. 1831.

Lesson, Traitéd'Omithologie : 132.

Rhopodytes diardi diardi (Lesson, 1831).

Premier spécimen. Lectotvpe, présente désigna¬

tion.

C.G. : 1997-1186.

Autres numéros de catalogues : A.C. n° 1752,

N C. n° 237.

Inscriptions sous le socle :

Java / M. Diard / 1821 / 1752 / Rhopodytes

Diardi Less. [même écriture].

Étiquette du socle :

Rhopodytes Diardi / (Less.) / Type / M. Diard /

Java.

Deuxième spécimen. Paralectotype.

C.G. r 1997-1185.

Autres numéros de catalogues : A.C. n° 1754,

N.C. n° 238.

Inscriptions sous le socle :

Java / M. Diard / 1820 / 1754 / Rhopodytes

Diardi / Less. I [même écriture, encre pâle] / 12

[au crayon |.

Étiquette :

Rhopodytes Diardi / (Less.) / Type / M. Diard /

Java.

Troisième spécimen, n’appartient pas à la série-

type.

C.G. : 1997-1202

Autres numéros de catalogues : A.C. n° 1753,

N.C. n° 236, 12.

Inscriptions sous le socle :

Malacca [surchargé, taché] / Souleyet / 1753 /

Rhopodytes Diardi / Less. / Type [même écriture]

/ 12 [au crayon].

Étiquette :

Rhopodvtes Diardi / (Less.) / Type / La Bonite /

Malacca.

La mention v Java » qui se trouve sur le socle des

deux premiers spécimens est certainement erronée,

car l’espèce vit dans la péninsule Malaise et à

Sumatra. Qu’ils aienr été envoyés de Java ne veut

pas forcément dire qu’ils y fussent récoltés, à

moins que l'espèce n’ait disparu de cette île

depuis le début du XIX e siècle. Au numéro 1753

de l’Ancien Catalogue, il est mentionné que le

troisième spécimen fut rapporté par Eydoux et

Souleyet au retour du voyage de La Bonite

(1836-1837). Il n'a donc pas pu être examiné

pour décrire l’espèce par Lesson (1830) qui pré¬

cise de plus « M. Diard a envoyé deux individus

de Java. (Mus. de Paris) ». C’est donc d’après les

deux premiers spécimens que le Melias diardi a

été décrit.

Melias tristis Lesson, 1831. Holotype par

monotypie.

Lesson, Traité d’Omithologie : 132.

Rhopodytes tristis tristis (Lesson, 1831).

C.G. 1997-1197.

Autres numéros de catalogues : A.C. n° 1749,

N.C, n" 233.

Inscriptions sous le socle :

Bengale / M. Duvaucel / 1825 / 1749 Rhopodytes

tiistis / Less. I Type / [même écriture] 13 [au

crayon].

Étiquette du socle :

Rhopodytes tristis / (Less.) / Type / M. Duvaucel /

Bengale.

Taccocua Lescbenaultii Lesson, 1831.

Lesson, Traité d’Omithologie : 144.

Taccocua lescbenaultii lescbenaultii Lesson, 1831.

Premier spécimen. Llolotype par monotypie.

C.G. : 1997-1200.

Autres numéros de catalogues : A.C. n° 1812,

N.C. n° 232.

382

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

Inscriptions sous le socle :

1. de Linde [sic] / par M Lechenault [sic] / juillet

1822 [écriture ancienne, plus ou moins effacée].

2. type du Tacoceun de Lechenault |sic] / 1812.

3. Taccocua sirkee (Gr.) [autre écriture ancienne].

4. Grand Taaoïde de / M. Lesson.

Etiquette du socle :

Taccocua sirkee I (Gr.) / T. Leschenaulti (Less.) T.

/ Leschenault / Inde.

Deuxième spccintcn, n’appartient pas à la série-

type.

C.G. : 1997- 1263.

Autres numéros de catalogues : A.C. n° 1814,

N.C. n° 231 •

Inscriptions sous le socle :

Bengale / M. Duvaucel / 1825 n°118 / Taccocua

sirkie I Grav / T. Leschenaulti I Lesson / n° 71 c? /

1814.

Étiquette du socle :

Taccocua sirkie / (Gr.) / T. Leschenaulti (Less.) T. /

â / M. Duvaucel / Bengale.

Lesson précise que le spécimen d’après lequel il

décrit le taxon Taccocua leschenaultii lui été

donné par Leschenault, ce qui est bien le cas du

premier spécimen, tuais non celui du second, qui

a été rapporté par Duvaucel et porte la mention

« type » non sous son socle, mais seulement sur

l’étiquette de ce dernier, probablement à la suite

d’une erreur. Le premier spécimen a également

servi à décrire le genre Taccocua Lesson, 1831.

Phaenicophaeus calyorhynchus Temminck,

1825. Syntype.

Temminck, Nouveau Recueil de Planches colo¬

riées 59 ; pl. 349.

Rhamphococcyx calyorhynchus calyorhynchus

(Temminck, 1825).

C.G. : 1997-1201.

Autre numéro de catalogue : A.C. n° 1764.

Inscriptions sous le socle :

Célèbes / Ech. eoll. Temminck / 1823 /

Phoenicophacus / calorhynchus / Tenu / 1764

[même ccriturc|.

Étiquette du socle :

Phoenicophacus / calorhynchus / (Tenu) / M.

Temminck / Célèbes.

L’orthographe originale du nom d’espèce est bien

calyorhynchus et non calorhynchus. Dans sa dia¬

gnose Temminck indique que les spécimens ont

été rapportés des MoJuques par « Monsieur

Reinwardt », et termine en écrivant : « On a

trouvé cette espèce à l’île de Célèbes. Deux indi¬

vidus font partie du Musée des Pays-Bas, un troi¬

sième est déposé dans le Musée de Paris ». Les

deux spécimens déposés au Musée de Leide y

sont conservés sous les numéros RMNLl 88262

et 88263 (Van den Hoek Ostende et al. 1997).

En vertu de la Recommandation 74 D du Code,

c’est parmi eux qu’il conviendrait de désigner un

lectotype.

Coccyzus erythropyga Lesson, 1842. Holotype

par monotypie.

Lesson, Revue de Zoologie 1842 : 210.

Morococcyx erythropyga erythropyga (Lesson,

1842).

C.G. 1847-1810.

Autres numéros de catalogues : A.C. n° 1883,

N.C. n° 173.

Inscriptions sous le socle :

Amérique Centrale / donné par M. Lesson / 1847 -

1810 / Moroccyx / crythropygia / Type de

(l.csson) / 1883 [même écriture!.

Étiquette du socle :

Morococcyx eryrhropygia / (Less.) / Type / M.

Lesson / Amérique cent.

Le nom d'espèce erythropyga, du grec Ttuya

« croupion », est un substantif, et doit donc res¬

ter invariable.

Cultrides Pucheranii Deville, 1851.

Deville, Revue et Magasin de Zoologie (sér. 2) 3 :

211 .

Neomorphus pucheranii pucheranii (Deville,

1851).

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. 1847-869.

Autres numéros de catalogues : A.C. n° 1831,

N.C, n° 209.

Inscriptions sous le socle :

Pérou / Sta Maria / M. Castelnau et Deville / 1847

n° 869 / 1831 / $ / Neomorpha Pucheranii /

Deville / Type |même écriture).

Étiquette du socle :

Neomorphus / femelle / Pucherani (Dev.) T. / M.

Deville / Pérou.

ZOOSYSTEMA • 1999 • 21 (2)

383

Voisin C. & Voisin J.-F.

Deuxième spécimen, Paralectotype.

C.G. 1847-870.

Autres numéros de catalogues : A.C. n° 1832,

N.C. n° 208.

Inscriptions sous le socle :

1. Brésil / P. [le tour maculé par une grosse tache

d’encre] Pérou ? / M.M. Castelnau er Deville /

1847 n° 870/ 1832.

2. Neomorphus Pucheranii / Deville / Type.

3. fig. dans T Allas Z V de M. Castelnau et Deville.

Etiquette du socle ;

Neomorphus / Pucheranii (Dcv.j T. / M. Deville /

Pérou.

Ces deux spécimens ont été représentés par Des

Murs, in Castelnau (1855). Lorsqu'il décrivait

Neomorphus pucheranii, Deville (1851) pensait

que le n° 209 représentait le plumage « du jeune

âge », alors que le n° 208 était l'adulte. En fait, il

s’agissait de différences de sous-espèces et non

d’âge, comme le fîr remarquer lodd (1925). Ce

dernier nomma Neomorphus pucheranii lepido-

phanes lodd, 1925 la forme qui vit au Pérou et

au Brésil au sud de l’Amazone, de la rivière

Ucayali à la rive gauche du Punis- Elle corres¬

pond au spécimen attribué à la forme adulte par

Deville et représenté pl. 6 par Des Murs (1855),

c'est-à-dire au n° 208. Le type de cette sous-espècc

est conservé dans les collections du Carnegie

Muséum, aux F.rats-Unis, sous le n" 92465

(Todd 1925). Dans ce cas. le lectotvpe de

Neomorphus pucheranii pucheranii ne peut être

que le spécimen n° 209, censé représenter le

jeune âge, et dont Neomorphus napensis

Chapman (1928) esr synonyme (Peters 1940). La

sous-espèce Neomorphus pucheranii pucheranii vit

au nord de l’Amazone, de l’Équateur au Pérou

ainsi qu a l’ouest du Brésil jusqu’à la rive nord de

la rivière Solimoes. Le spécimen n a 208 reste

néanmoins le paralectotype de l’espèce Neo¬

morphus pucheranii.

Carpococcyx Renauldi Oustalct, 1896.

Oustalet. Bulletin du Muséum d’Histoire natu¬

relle de Paris 1896 (2) : 314.

Carpococcyx renauldi Oustalet, 1896.

Premier spécimen. Lectotvpe, présente désigna¬

tion.

C.G. 1896-2364.

Autre numéro de catalogue : N.C. n° 38 C.

Etiquette :

Carpococcyx Renauldi / Oustalet / Type / R.P.

Renaulr / Annam / 1896 - iT 2364 / Catal. des

Gai. 38 C.

Deuxième spécimen. Paralectotype.

C.G. : 1896-2363.

Autre numéro de catalogue : N.C. 38 B [sur le

socle].

Inscriptions sous le socle :

1. Annam / Acq du R.P. Renauld. / 1896 -

n“ 2363 [inscription illisible-e.e. ?]

2. Carpococcyx Renauldi / Oust (autre écriture],

3. Inscription illisible car recouverte d’encre.

Etiquette du socle :

Carpococcyx Renauldi / (Ousr.) / R.P. Renauld /

Annam./ 38 B.

A la fin de sa description de l’espèce, Oustalet

écrit : « La nouvelle espèce que je viens de carac¬

tériser, d’après trois individus... ». Seuls les deux

spécimens ci-dessus (1896-2363 et 2364) sont

inscrits au Catalogue général. Dans ce même

registre il est noté qu’un troisième spécimen a

bien été reçu du R. P. Renauld mais que, en très-

mauvais état, il n'a pas été « pris en charge ». Le

numéro de Nouveau Catalogue sur l’étiquet te du

lectotvpe indique qu’il s'agit d'un spécimen

autrefois monté. Il lut remis en « peau plate » il y

a longtemps, et son socle a disparu.

Coua cristata dumonti Delacour, 1931.

Delacour, L'Oiseau et la Revue française

dOntithoïogie I : 475.

Coua cristata dumonti Delacour, 1931.

Premier spécimen. 1 lolorype par désignation ori¬

ginale.

C.G. 1932-1085.

Etiquette verre :

Coua / crisrala / dumonti / Delacour / a : 142 /

Type [en travers et encadré de rouge] III Madag

[en travers! / 1009 c? 00 / Tsiandro / Muséum

Paris / 11.7. 29 / C.G. 1932 N° 1085.

Deuxième spécimen. Paratype.

C.G. 1932-1086.

Étiquette verte :

Coua / cristara / iris brun / dumonti / Delacour /

a : 142 III Madag [en travers] S 00 / 936 /

384

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

Muséum de Paris / Tsiandro / C.G. 1932-1086 /

8.7.29 .

Troisième spécimen. Paratype.

C.G. 1932-1087.

Etiquette verte :

Coua / crisrata / dumonti Delacour / S / a : 137

/// Madag [en travers) 1104 / Tsiandro / Muséum

de Paris / 16.7.29 / C.G. 1932- N" 1087.

Quatrième spécimen. Paratype.

C.G. 1932-1088.

Etiquette verte :

Coua / cristata / dumonti / Delac SI a : 135 ///

Madag [en travers] 1132 / Tsiandro / Muséum de

Paris / 17.7.29 / C.G. 1932- N°1088.

Cinquième spécimen. Paratype.

C.G. 1932-1089.

Étiquette :

Mission F.A.A. / à Madagascar / J. Delacour [ces

trois lignes en travers] / le 5 March 1931 /

Namoroka 1 day south of Soalala / Coua cristata

dumonti / sexe : cî Delac. / N" 9873 [en travers]

III Muséum de Paris / C.G. 1932 - N°1089 /

Aile 137 / Rand et Archbold coll. / & Dumont /

N° 9873.

Sixième spécimen. Paratype.

C.G. 1932-1090.

Étiquette :

Mission l-.A.A. / à Madagascar [en travers) /

J. Delacour / le 27 feb, 1931 / Soalala / Sexe : d /

N° 9956 [en travers] III Coua cristata / dumonti

Delac. / Muséum de Paris / C.G. 1932-N°1090 /

Aile 142 / Rand et Archbold coll. / & Dumont.

Septième spécimen. Paratype.

C.G. 1932-1091.

Étiquette :

Mission l-.A.A. / à Madagascar / J. Delacour [en

travers] / le 5 Match 1931 / Namoroka 1 day

south of Soalala / Coua cristata dumonti / Sexe : V

/ Delac. /// N“ 9871 [en travers! / Muséum de

Paris / C.G. 1932- N°I091 / Aile 1.38 / Rand et

Archbold coll. / & Dumont.

Huitième spécimen. Paratype.

C.G. 1932-1092.

Étiquette :

Mission F.A.A. / Muséum de Paris / 22.5.1930 /

C.G. 1932 - N° 1092 / R. Decary coll. III « Coua »

19/ Coua cristata dumonti / Delac. / Maeva-

tanana / Madag.

Neuvième spécimen. Paratype.

C.G. 1932-1093.

Étiquette verte :

Coua / cristata / dumonti / ? Delac. O / a : 136

/// 1031 / Tsiandro / 12.7.29 / Muséum de Paris /

C.G. 1932- N°1093.

Delacour précise qu i] a établi sa description sur

29 spécimens, sans indiquer où ils les a déposés.

Seuls les 9 que nous venons d’examiner figurent

dans les collections du Muséum national

d’Histoire naturelle, et nous n avons pas réussi à

retrouver la trace des 20 autres.

Coua cristata maxima Milon, 1950. Holotype

par désignation originale.

Milon. Bulletin du Muséum national d’Histoire

naturelle (2) XXII ( 1 ) : 65.

Coua cristata maxima Milon, 1950.

C.G. 1950-392.

Étiquette blanche :

Don du Colonel Ph. Milon / Loc : Madagascar S. /

C.G. 1950 N°392 III Type [en travers, encadré de

rouge] / Coua cristata maxima Milon / Loc. : Port

Dauphin / 18 février 1948 / 6.

Coua cristata variété pyropyga Grandidier,

1867.

Grandidier, Revue et Magasin de Zoologie (sér. 2)

19:86,392.

Coua cristata pyropyga Grandidier, 1867.

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. 1867-753.

Autres numéros de catalogues : A.C. n° 1822 C,

N.C. n u 36 [non reporté sur le socle].

Inscriptions sous le socle :

1. de la côte Sud Ouest / de Madagascar / donné

par M. A. Grandidier / 1867 / 753.

2. Jne â.

3. Coua cristata (Puch.) [sic] [cette dernière men¬

tion barrée au crayon, un L. inscrit au-dessus] / Bp

Consp. av. t. I. p. 62 / Var. pyropyga.

4. paraphe entre parenthèses, souligné et rappelant

un A majuscule,

Étiquette du socle :

Coua cristata (L.) 9 / Var. pyropyga (Grand.) /

M, Grandidier / Madagascar.

Deuxième spécimen. Paralectotype.

C.G. 1867-754.

ZOOSYSTEMA • 1999 - 21 (2)

385

Voisin C. & Voisin J.-F.

Autres numéros de catalogues : A.C. n° 1822 A,

N.C. n°38 [non reporté sur le socle].

Inscriptions sous le socle :

1. de la côte Sud Ouest de / Madagascar / donné

par M. A. Grandidier / 1867 / 754.

2. 6 3.

3. Coua cristara (Puch.) [sic] [cette dernière men¬

tion barrée au crayon, un L. inscrit au-dessus] / Bp

Consp. av. t. I, p. 62 / Var. pyrnpyga.

4. même paraphe que le précédent.

Etiquette du socle :

Coua cris tara (L) ? I Var. pyropyga (Grand.) /

M. Grandidier / Madagascar.

Troisième spécimen. Paralectotype.

Conservé au Musée de Zoologie de Nancy.

C.G. 1867-755.

Autre numéro de catalogue : A.C. n° 1822 B.

Inscriptions sous' le socle :

1. de la cote sud ouest / de Madagascar / donné par

M. A. Granclidiet.

2. Coua cris tau (Puch.) [sic| [cette dernière men¬

tion barrée, un L. inscrit au-dessus] / Bp. Consp.

av. t. 1. p. 62 / Var. pyropyga.

3. même paraphe que les précédents.

Le troisième spécimen a été donné en 1931 au

Musée zoologique de l’Universiré et de la Ville de

Nancy, et s’y trouve toujours (A Philippot,

comm. pers). Les écritures et les paraphes sont les

mêmes sur les trois socles. De même que pour les

taxons précédents, ces spécimens sont trop sem¬

blables pour que l’on puisse décider auquel se

rapportent les mesures données par Grandidier

dans sa description, La référence au Conspectus

Generum Aviurn de Charles Bonaparte est

curieuse. En effet, la parution de cet ouvrage s’est

échelonnée de 1850 à 1857, et il ne peut donc

mentionner un taxon décrit en 1867, On ne

trouve même pas trace de ce dernier dans les très

nombreuses notes manuscrites portées dans

l’exemplaire personnel de Grandidier, auparavant

propriété de J. Vcrrcaux, lequel lavait fait relier

avec des feuilles blanches réservées à cet usage.

L'hypothèse la plus probable est que cette ins¬

cription renvoie à la mention de Coua cristata

(Linné), page 109 et non 62, où ce sont des

Caprimulgidcs qui sont traités.

Coua serriana Pucheran, 1845.

Pucheran. Revue de Zoologie 1845 : 51.

Coua serriana Pucheran, 1845.

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G.: 1997-1188.

Autres numéros de catalogues : A.C. n° 1819,

N.C. tT9,

Inscriptions sous le socle :

1. de Magadascar / par M. Bernier / 1835 [année]

/ 1819

2. Nom de pays / Coua Semang

3. Coccyzus [ccvzus barré pour transformer

Coccyzus en Coua, ua étant écrit au-dessus] serria¬

na Puch. [presque effacé] (T).

Etiquette du socle :

Coua serriana / (Puch.) / Type / M. Bernier /

Madagascar.

Second spécimen. Paralectotype.

C.G. : 1997-1187.

Autres numéros de catalogues : A.C. n° 1820,

N.C. n° 8.

Inscriptions sous le socle :

Madagascar / M. Bernier / 1834 [année] / 1820 /

Coua serriana / Puch. Type [même écriture].

Etiquette du socle :

Coua serriana / Type / (Puch.) / M. Bernier /

Madagascar.

Troisième spécimen. Paralectotype.

C.G. : 1997-1189.

Autres numéros de catalogues : A.C. n° 1821,

N.C. n° 10.

Inscriptions sous le socle :

Madagascar / M. Sganzin / 1832 [année] / Coua

serriana P Puch, Type [.toutes ces mentions de la

même main, sauf les numéros de catalogues].

Étiquette du socle :

Coua serriana / (Puch.) I Type / M. Sganzin /

Madagascar.

Coua cursor Grandidier, 1867.

Grandidier, Revue et Magasin de Zoologie (sér. 2)

19:86,391.

Coua cursor Grandidier, 1867.

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. 1867-759.

Autres numéros de catalogues : A.C. n° 1818 B,

N.C. n° 30.

Inscriptions sous le socle :

386

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

1. de Madagascar / donné par M r A. Grandidier :

â / Coua cursor / 1 vpe / paraphe.

2. 1818 b [barré] 18]8 B [autre main, au crayon].

Etiquette du socle :

Coua cursor / (Grand.) 161 Type / M. Grandidier /

Madagascar.

Deuxième spécimen. Paralectotype.

C.G : 1867-758.

Autres numéros de catalogues : A.C. n° 1818 A,

N.C. n° 31.

Inscriptions sous le socle :

1. de Madagascar / envoyé par M 1 Grandidier /

Coua cursor. A. Grandidier / paraphe.

2. 1818 A- 31.

Étiquette du socle :

Coua cursor / (Grand.) / Type / M. Grandidier /

Madagascar.

Les inscriptions et le paraphe compliqué sont de

la même main sous les deux socles. Les spéci¬

mens que nous avons sous les yeux sont trop

semblables pour que l'on puisse déterminer

auquel des deux s’appliquent les mesures que

Grandidier donne dans sa description, d'autant

plus qu'il ne précise pas de quelle manière il les a

prises. Dans ces circonstances, nous avons choisi

comme lcctotype le spécimen déjà inscrit comme

« type » dans le Nouveau Catalogue.

Coua Verreauxi Grandidier, 1867.

Grandidier, Revue et Magasin de Zoologie (sér. 2)

19:86,417.

Coua verreauxi Grandidier, 1867.

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. : 1867-757.

Autres numéros de catalogues : A.C. n° 1826 B,

N.C. n°32.

Inscriptions sous le socle :

1. de Madagascar. / donné par M r A. Grandidier;

1867/757/ 1826 B.

2. Coua Vetrauxii [rayé] Verreauxii [écrit dessous]

A. G randidier / Paraphe.

Étiquette du socle :

Coua verreauxii / (Grand.) / Type / M. Grandidier

/ Madagascar.

Deuxième spécimen. Paralectotype.

C.G. 1867-756.

Autres numéros de catalogues : A.C. n° 1826 A,

N.C. n u 33.

Inscriptions sous le socle :

1. de Madagascar / donne par M r Grandidier /

1858 / 756/ 1826 A.

2. Coua Verrâuxii [sic]. A. Grandidier / Paraphe.

3. F [au crayon].

Étiquette du socle :

Coua verreauxii / (Grand.) 16 1 M. Grandidier /

Type / Madagascar.

Les écritures des inscriptions ainsi que les

paraphes rappelant un H sont les mêmes sous les

deux socles, mais diffèrent de ceux des socles des

types de Coua cursor. Comme pour cetcc dernière

espèce, ces deux spécimens de Coua verreauxi

sont trop semblables pour que l'on puisse déter¬

miner auquel s’appliquent les mesures données

par Grandidier dans sa description.

Coua Coquereli Grandidier, 1867.

Grandidier, Revue et Magasin de Zoologie (sér. 2)

19:86,391.

Coua coquereli Grandidier, 1867.

Premier spccimcn. Lectotype, présente désigna¬

tion.

C.G. 1867-75L

Autres numéros de catalogues : A.C. n° 1828 D,

N.C. n°28.

Inscriptions sous le socle :

1. de Madagascar / donné par M. A. Grandidier /

1867 751 / 6 / Coua Coquerellii [sic], Alf.

Grandidier / Revue et Mag. Zoolo. / 1867 p. [la

suite détruite par un trou, toutes ces mentions delà

même écriture].

2. Peau nue d’un bleu clair / avec du violet rosé

à l'angle postérieur [autre écriture, au crayon) /

1828 D.

3. Paraphe.

Étiquette du socle :

Coua Coquerelii 6 / (Grand.) Type /

M. Grandidier / Madagascar / 28.

Deuxième spécimen. Paralectotype.

C.G. 1867-752.

Autres numéros de catalogues : A.C. n° 1828 C,

N.C. n° 29.

Inscriptions sous le socle :

1, de Madagascar / donné par M r A. Grandidier /

1867 752 Cat. Gen. / Coua Coquerellii [sic]. Alf.

Grandidier / Rev. et mag. Zool (1867 p. [la suite

illisible] / 6.

ZOOSYSTEMA • 1999 • 21 (2)

387

Voisin C. & Voisin J.-F.

2. Paraphe.

3. 1828 C [au crayon].

Étiquette du socle :

Coua Coquerelii (emelle / (Grand.) Type /

M. Grandidier / Madagascar / 29.

Sous les deux socles, l'écriture des mentions 1 et

le paraphe sont les mêmes que sous les socles des

types de Coua verreauxii. Comme pour les deux

taxons précédents, les spécimens sont trop sem¬

blables pour que l'on puisse décider auquel se

rapportent les mesures données par Grandidier

dans sa description. Le bec du second spécimen

est cassé.

Coua Reynaudii Pucheran, 1845.

Pucheran, Revue de Zoologie 1845 : 51.

Coua reynaudii Pucheran, 1845.

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. : 1997-1218.

Autres numéros de catalogues : A.C. n° 1815,

N.C. n” 13.

Inscriptions sous le socle :

Madagascar / M. Bernier / 1834 / 1815 / 6 /

Coua Reynaudii / Puch. / Type.

Étiquette du socle :

Coua Reynaudii / (Puch.) / 6 / Type / M. Bernier

/ Madagascar.

Deuxième spécimen. Paralectotvpe.

C.G. : 1997-1216.

Autres numéros de catalogues : A.C. n° 1816,

N.C. n°l4.

Inscriptions sous le socle :

Madagascar / M. Bernier / 1834 / 1816 9 / Coua

reynaudi / Puch / Type / 12 1/2 23.

Étiquette du socle :

Coua Reynaudii / (Puch.) 16 1 Type / M. Bernier

/ Madagascar.

Troisième spécimen. Paralectotype.

C.G. : 1997-1217.

Autres numéros de catalogues : A.C. n° 1817,

N.C. n° 15.

Inscriptions sous le socle :

Madagascar / M. Bernier / 1834 / 1817 / Coua

Reynaudii / Puch / Type.

Étiquette du socle :

Coua Reynaudii / (Puch.) / Type / M. Bernier /

Madagascar.

Quatrième spécimen. Paralectotype.

C.G. ; 1997-1220.

Autres numéros de catalogues : A.C. n° 1818,

N.C. n° 16.

Inscriptions sous le socle :

Madagascar I M. Bernier / 1818 / Coua Reynaudii

I Puch. / Type

Étiquette du socle :

Coua Reynaudii / (Puch.) 16 1 Type / M. Bernier /

Madagascar.

Cinquième spécimen. Paralectotype.

C.G, : 1997-1219.

Autres numéros de catalogues : A.C. n° 1821 A,

N.C. n° 17.

Inscriptions sous le socle :

Madagascar / M. Bernier / 1834 / 1821 A 6 /

Coua Reynaudii / Type [encre pâlie].

Étiquette du socle :

Coua Reynaudii / (Puch.) 16 1 Type / M. Bernier

/ Madagascar.

Ces cinq spécimens, tous mâles, ont été envoyés

en même temps au Muséum par M. Bernier, et

Pucheran les a donc tous eus à sa disposition

pour rédiger sa description, fort succincte car elle

tient en quatre lignes et concerne effectivement

le mâle adulte.

Coccycus [sic] Delalandei Temminck, 1827.

Holotype par monotypie.

Temminck, Nouveau Recueil de Planches colo¬

riées 74, pl, 440.

Coua delalandei (Temminck, 1827).

C.G. : 1997-1190.

Autres numéros de catalogues : A.C. n° 1827

[même numéro d’ordre que celui de l’année de

description du taxon], N.C. n” 6.

Jeton fixé à la patte : 6 / 2514.

Inscriptions sous le socle :

1. Madagascar / M. Delaiande / 1 820.

2. Colin Delalandei / Pucher [peu lisible, le bois a

•• bu . l’encre] / Coccy/.tis “ [signe de ditto en-des¬

sous de Delalandei| Tein. / Type / 1827.

Étiquette du socle :

Cou.i de Dcl.tlande / Coua Delalandei / ad. Type

(Teinnt.) / Voy. de Delalande Madagascar.

Le socle semble bien d origine, même si toutes

les inscriptions sont de la même écriture

ancienne. Temminck précise qu’il décrit cette

388

ZOOSYSTEMA • 1999 ■ 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

espèce d'après un « individu unique .... déposé

dans les galeries du Muséum de Paris ». Outre ce

dernier, le Muséum de Paris possède un second

spécimen de cette espèce éteinte et (ort rare dans

les collections, rapporté par Bernier en 1834 et

qui semble être le dernier collecté (Jouanin

1962).

Centropus violacens Quoy & Gaimard, 1830.

Holotype par monotypie.

Quoy Sc Gaimard, Voyage de découvertes de

L’Astrolabe, Zool. 1 : 229 (1830) et Atlas Ois.,

pl. 19 (1833).

Centropus violaceus Quoy & Gaimard, 1830.

C.G. : 1997-1195.

Autres numéros de catalogues : A.C. n° 1801,

N.C. n° 315.

Inscriptions sous le socle :

N‘ !U Irlande / M.M. Quoy et Gaimard /

L’Astrolabe 1829 / 1801 / Nesocentor violaceus /

Q et G / t ype.

Étiquette du socle :

Nesocentor violaceus / Type (Q & G) /

L’Astrolabe / N. Irlande.

Les mentions sous le socle sont toutes de la

même main et postérieures à la création du genre

Nesocentor Cabanis & Pleine, 1863, ce qui

montre que ce spécimen Fait bien partie de ceux

dont on a changé le socle à la fin du siècle der¬

nier. 11 Fut le seul de son espèce dans nos collec¬

tions jusqu’à l’arrivée d’un second en 1921.

Centropus Menebiki Garnot in Lesson ôi

Garnot, 1828.

Garnot in Lesson & Garnot, Voyage autour du

Monde exécuté par ordre du Roi sur la Corvette de

Sa Maj. La Coquille pendant les années 1822,

1823,' 1824 et 1825. Zoologie 1 : 600 (1828),

Atlas, pl. 33 (1838).

Centropus menebiki menebiki Garnot, 1828.

(= Centropus menbeki menbeki auct.).

Premier spécimen. Holotype par monotypie.

C.G./ 1997-1196.

Autres numéros de catalogues : A.C. n° 1805,

N.C. n° 309.

Inscriptions sous le socle :

N c,lc Guinée / par M.M. Garnot et Lesson / Exp.

de la Coquille / n” 1 54 / Nesocentor Menebekii

|sic| / (Less et Gar) / 1805 Type [même écriture].

Étiquette du socle :

Nesocentor Menebekii / Type / (Less.& Garn.) /

La Coquille / N. Guinée.

Second spécimen, n’appartient pas à la série-type.

C.G. : 1997-1194.

Autre numéro de catalogue : N.C. n° 310 (pas

de numéro d Ancien Catalogue).

Inscriptions sous le socle :

Astrolabe / Voyage Pôle Sud / N' 11 " Guinée /

Nesocentor Menebekii [sic] / (1. et G) / iris jaune

|tout de la même écriture et difficilement lisible] /

17 [crayon].

Etiquette du socle :

Nesocentor Menebekii [sic] / Type (Less. & Garn.)

/ Voy. Pôle Sud / N. Guinée.

Le second spécimen de Centropus menebiki a été

rapporté par l’expédition de L’Astrolabe, com¬

mandée par Jules Dumont d Urville et qui eut

lieu de 1826 à 1829. Il ne peur donc erre l’un des

types de ce taxon qui fut décrit en 1828, avant

même le retour de ce navire. La mention « type »

n’est d’ailleurs portée que par l’étiquette du socle.

Dans la légende de la figure parue dans l Atlas on

lit : menbeki. Il F agir certainement d’une erreur

d’inadvertance, car dans la table des matières de

l’Atlas, on trouve menebikii et en tête de la des¬

cription menebiki. D’après Davies, Shcrborn &

Woodward (1906) la parution de l’Atlas se serait

étalée sur de nombreuses années et la date 1826

portée sur la couverture ne correspondrait qu’à

celle des premières livraisons. Lu planche 33, qui

nous intéresse ne serait parue qu’en 1838, soit

dix ans après la description. Dans ces conditions

l’orthographe correcte est menebiki.

Centropus ateralbus Lesson, 1826.

Lesson, Bulletin Universel des Sciences et de

l'Industrie 8 (2) : 113.

Centropus ateralbus Lesson, 1826.

Premier spécimen. Lectotype présente désigna¬

tion.

C.G. : 1997-1191.

Autres numéros de catalogues : A.C. n° 1806,

N.C n° 305.

Inscriptions sous le socle :

N' ik Irlande / M. Garnot et Lesson / La Coquille

ZOOSYSTEMA • 1999 • 21 (2)

389

Voisin C. & Voisin J.-F.

Nesoccntor uteralbus [on a d’abord écrit « atteral-

lus », puis corrige] / Less. et Garnot / Type / 1806

[même écriture].

Etiquette du socle :

Nesoccntor aicralbus / (Less. & Garn.) / Type / La

Coquille / N. Irlande.

Deuxième spécimen. Paralectotype.

C.G. : 1997-1192.

Autre numéros de catalogues: A.C. n° 1807,

N.C. n° 306.

Inscriptions sous le socle :

1. Centropus airalbiui [? - peu lisible et rayé, écri¬

ture ancienne].

2. [6 lettres majuscules effacées, plus récentes. On

reconnaît P, M, et N],

3. Lesson et Garnot [écriture ancienne] / 1807

[autre écriture].

4. Nesocentor / ateralbus / (Less et Garn) / ateral-

bus [écriture plus récente. La fin du premier ateral¬

bus corrigée au crayon, le second entièrement au

crayon],

5. 151 / Expédition du perrey 437 / Nlle Irlande

[écriture ancienne].

Étiquette du socle :

Nesoccntor ateralbus / (Less. & Garn.) / Type / La

Coquille / N. Irlande.

Ces deux spécimens ont été rapportés au

Muséum en même temps, car L. I. Duperrey,

mentionné sous le socle du second, était le

Commandant de La Coquille , et Lesson les a eus

certainement à sa disposition tous les deux lors¬

qu’il a décrit Centropus ateralbus. Le (ait que la

mention » Type », présente dans l’Ancien

Catalogue, n ait pas été reportée sous le socle du

second doit procéder d un oubli.

Centropus celebensis Quov & Gaimard, 1830.

Lectotype présente désignation.

Quoy & Gaimard, Voyage de découvertes de

L’Astrolabe, Zool, 1 : 230. Atlas des Oiseaux,

pl. 20.

Centropus celebensis celebensis Quoy & Gaimard,

1830.

C.G. : 1997-1193.

Autres numéros de catalogues : A.C. n” 1802,

N.C. n°303.

Inscriptions sous le socle :

1. Pyrrhocentor* / = Celebensis / (Q. et G.) [écri¬

ture ancienne].

2. Coucal des Célèbes par M Quoy et Gaim [écrit

en suivant le bord du socle] / Célèbes.

3. Expédition de lastrolabe [sic] / 1829 / 1802

[autre écriture, comme les mentions 2],

Etiquette du socle :

Pyrrhocentor / celebensis (Q. & G.) T. /

1 Astrolabe / Célèbes.

Deux autres spécimens appartenant à la série-

type se trouvent au Nalionaal Natuurhistorisch

Muséum de Leiden (Pays-Bas), sous les numéros

RMNH 88168 et 88169 (Hoek Ostcnde et al.

1997). Ils deviennent dont des paralectotypes.

Cuculus nigrorufus Cuvier, 1816. Lectotype,

présente désignation.

Cuvier, Le Régne animal i : 426 (note).

Centropus nigrurufus (Cuvier, 1817).

C.G. : 1997-1 147.

Autres numéros de catalogues : A.C. n° 1776,

N.C. n°270.

Inscriptions sous le socle :

Gap. B. Espérance / 1776 ! Centropus nigrorufus /

Cuvier / Type / Expd. Vaillant / 1776 / Cap. B.

Espérance |méme écrirurej.

Étiquette du socle :

Centropus nigrorufus / (Cuv.) / Type / Cap de

B.E.

La description de Cuvier est explicitement fon¬

dée sut telle du « Coucal noiruu » de l.evaillant

(1808) qui décrit un mâle et une femelle, tués

alors qu'ils étaient en couple. Nous ignorons ce

qu’est devenu le second spécimen. La mention

« Vaillant » sous le socle indique bien que c’est

Levaillant qui a rapporté ce spécimen.

Centropus melanops Lesson, 1831. Holotype

par monotypie.

Lesson, Traité d'Ornithologie : 137.

Centropus melanops melanops Lesson, 1831.

C.G. : 1997-1248.

Autres numéros de catalogues : A.C. n° 1797,

N.C. n” 307, 14.

Inscriptions sous le socle :

[paraphe illisible] / Pliillippines [sic] / M. Capit.

Dubuisson / 1821 / 1797 14 / Pyrrhocentor mela¬

nops / Cuv. / Type / 1797 [même écriture].

Etiquette du socle :

Pyrrhocentor / melanops (Cuv.) Type /

M. Dubuisson / Philippines.

Lesson termine sa description du Centropus

390

ZOOSYSTEMA • 1999 ■ 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

melanops par la mention « Mus. de Paris ; M. le

capitaine de vaisseau Dubuisson », ce qui indique

qu’il travaillait bien sur ce spécimen. D’autre

part. Cuvier ria pas décrit ce taxon et la mention

de son nom sous le socle est erronnéc. Il n’est pas

non plus l’auteur du genre Pyrrhocentor, qu’il faut

attribuera Cabanis & Heine (1863).

Taxons mis en synonymie

Cuculus edolius CUVIER, 1817.

Cuvier, Le Régné animal 1 : 425.

Clamator serratus (Sparrman, 1786).

Premier spécimen. Holotype par monotypie.

C.G. : 1997-1214.

Autres numéros de catalogues : A.C. et N.C. :

néant. 1 10-2,

Inscriptions sous le socle :

Coucou edolio I Vaill: 207. / C. ater / Type, [écri¬

ture ancienne sur une très vieille étiquette collée

sous un socle qui n'a pas été refait. La mention

« type » a été repassée, le T étant moderne].

Etiquette du socle :

Coucou edolio d Vaill. 207 / Cuculus edolius

Cuv. / C. serratus, C. ater Gm. /

du Cap de Bonne Espérance.

Deuxième spécimen, n’appartient pas à la série-

type.

C.G. : 1997-1215.

Autres numéros de catalogues : A.C. n° 1927,

N.C. n° 144.

Inscriptions sous le socle :

1. Cap de B. Espérance / M. Delalande / 1822 /

1927/ 9.

2. Coccystes serratus / Sparm. / C. edolius Cuv. /

Type.

Étiquette du socle :

Coccystes serratus 9 / C. edolius (Cuv.) Type /

Delalande Cap de B.E.

Conformément à une habitude répandue à

l’époque, le numéro 207 marqué sous le socle du

premier spécimen est celui de la planche de

v l’Histoire naturelle des Oiseaux d’Afrique » de

Levaillant (1806) sur laquelle est représenté cet

oiseau. Cet exemplaire est bien celui dont

Levaillant s’est servi pour décrire son n Coucou

edolio ». La description, très brève, de Cuculus

edolius par Cuvier renvoie à celle du « Coucou

edolio ». Le type de Cuvier est donc le même que

l’exemplaire de Levaillant. Le second spécimen a

été donné au Muséum par Delalande en 1822 et

ne peut donc pas erre celui auquel se réfère

Cuvier pour sa description, parue en 1817.

Coccystes brazzae Oustalet, 1886. Holotype par

monotypie.

Oustalet, Le Naturaliste 8 (38) : 299.

Paebycoccyx audeberti validus (Reichenow,

1879).

C.G. 1885-247.

Autre numéro de catalogue : N.C. n° 161.

Inscriptions sous le socle :

Congo / M. Brazza / 1885 n 0 247 / â / Coccystes

Brazzae / Oust. / Type [même écriture].

Etiquette du socle :

Coccystes Brazzae 9 / (Oust.) Type : M. de Brazza /

Congo.

Oustalet (1886) a décrit Coccystes brazzae sur ce

seul spécimen.

Cuculus lineatus Lesson, 1831. Holotype par

monotypie.

Lesson, Traité d'Ornithologie : 152.

Cuculus saturnins lepidus S. Millier, 1845.

C.G, : 1997-1235.

Autres numéros de catalogues : A.C. n° 1941,

N.C, n u 50.

Inscriptions sous le socle :

CochinchinE / M. Diard / 1821 / 1941 / Cuculus

poliocephalus / Lath. / C. lineatus / Less. / Type

(même écriture].

Étiquette du socle :

Cuculus poliocephalus / C. lineatus (Less.) Type /

M. Diard / Cochinchine.

La description, fort succincte, de Lesson,

s’applique bien à cet oiseau et se termine par :

« Habite les îles indiennes de l’Est. (Mus. de

Paris) ». Ceci confirme que le spécimen-type se

trouve bien au Muséum de Paris, mais est en

contradiction avec la mention » Cochinchine »

qui laisse suspecter une erreur de transcription,

d'ailleurs ancienne car elle existe déjà dans

l’Ancien Catalogue. Aucune des livres de spéci¬

mens envoyés de Cochinchine au Muséum par

Diard ne fait état de « Coucou à ventre rayé » ou

ZOOSYSTEMA • 1999 • 21 (2)

391

Voisin C. & Voisin J.-F.

de « Coucou rayé », ou y trouve seulement la

mention d’un « Coucou noir à gros bec » et d'un

« Coucou noir (petit) ■> (numéros 12 et 13, juin

1822). Au contraire, parmi les oiseaux de Java et

Sumatra reçus de ce voyageur, on trouve mention

de trois « Coucous rayés » (numéro 86, 6 mai

1821), ainsi que d’un « Coucou jaune rayé >«

(numéro 132, 27 décembre 1821), Il semble

donc logique de penser que notre exemplaire

provient non pas de Cochinchine, d’où Diard n’a

envoyé que peu de matériel, mais bien de Java ou

de Sumatra. Ceci est confirmé par i examen de ce

spécimen, qui, par tous ses caractères, et entre

autres ses mensurations (cttlmen : 17 mm, bec de

l'extrémité distale de la narine à l’apex :

13,1 mm, aile : 152 mm, tarse ; 17 mm), est un

Cuculus su tannus le pi dus S. Millier, 1845, dont

Faire de distribution s’étend de la péninsule

Malaise à Timor, et non un C poliocephalus

poliocephalus Latham, 1790, qui ne se rencontre

qu’en tant que migrateur en Cochinchine

(Delacour & Jabouille 1931 ; David-Beaulieu

1932 ; Peters 1940), où sa présence est d’ailleurs

mise en doute par Wells ik Becking (1975).

Nous avons donc affaire à l’holotype de Cuculus

lineatus, Contrairement à ce que le determina-

teur, probablement abusé par l’erreur de prove¬

nance, a inscrit sous le socle, C. lineatus n’est pas

synonyme de C. poliocephalus poliocephalus, mais

est au contraire un synonyme plus ancien de

C. saturatus lepidus :

Cuculus lineatus I.esson, 1831

= Cuculus lepidus S. Muller 1845, syn. nov.

Cette synonymie semble avoir échappé aux

auteurs et, entre autres, ni Shelley (1891), ni

Peters (1940), ni Becking (1975) n’y font allu¬

sion. Cependant, le nom de lineatus Lesson,

1831 n'a pas été employé depuis des décennies

pour désigner ce raxon, et la localité d’origine de

l'holotype n’est pas connue avec précision Afin

de préserver la stabilité de la nomenclature, il

conviendrait de demander à la Commission

internationale de Nomenclature zoologique de

supprimer l'épithète subspécifique lineatus

Lesson au profit de lepidus S. Millier, en vertu

des dispositions de l’article 79 du Code.

Cuculus variegatus Vieillot, 1817. Lectotype,

présente désignation.

Vieillot, Nouveau Dictionnaire d'Histoire natu¬

relle 8 : 224.

Cuculuspal/iduspallidus (Latham, 1801).

C.C. : 1997-1224.

Autres numéros de catalogues : A.C. n° 1954,

N.C. n° 80.

Inscriptions sous le socle :

l’imor / M. Maugé / 1954 / Cacomantis pallidus. /

Lath / C. Variegatus / Vieil. / Type [même écri-

turel.

Etiquette du socle :

Cacomantis pallidus. / (Lath.) / M. Maugé /

Timor.

Ce spécimen est en excellent état conformément

il la réputation de R. Maugé, zoologue et taxider¬

miste de l’expédition Baudin, en 1800-1804.

Cuculusp. pallidus est répandu dans l'est et le sud

de l'Australie continentale ainsi qu’en Tasmanie.

L’erreur de localité sur le socle provient peut-être

du lait que l’expédition Baudin fit escale à Timor

(Brosse 1983). R. Maugé étant décédé en

Tasmanie en février 1802, l'origine do tous ses

spécimens n'a pas du être possible à établir avec

certitude. Dans sa diagnose, Vieillot écrit,

« originaire d’Australasie ». Vieillot fait d’abord la

description de l’adulte, puis celle du jeune. Bien

qu’il ait eu au moins deux spécimens à sa disposi¬

tion, seul celui-ci demeure dans nos collections.

Cuculus melanorhynchus Cuvier in I.esson,

1831.

Cuvier in Lesson, Traité d’Omithologie : 141.

Coccyzus melacoryphus Vieillot, 1817.

Premier spécimen. Holotypc par monorypie.

C.G. : 1997-1234.

Autres numéros de catalogues : A.C. n° 1882,

N.C. n°168.

Inscriptions sous le socle :

Brésil / M. St. Hilaire / 1822 / 1882 / Coccyzus

melanocoryphus [sic] V. / C. melanorhynchus

Cuv. / Type [même écriture].

Etiquette :

Coccyzus melanocorhyphus [sic] / C. melanorhyn¬

chus (Cuv.) I. / Brésil.

Deuxième spécimen, n’appartient pas à la série-

type.

392

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

C.G. / 1997-1232.

Autres numéros de catalogues : A.C. n° 1881,

N.C. n° 167.

Inscriptions sous le socle :

Brésil / M.Salle / 1821 / 1881 10 [superposés] /

Coccyzus / mclanorhynchus / V. [.sic] / C. melano-

rhynchus / Cuv. / Type [même écriture].

Étiquette du socle :

Coccyzus melanocorhyphus [sic] / C. melanorhyn-

chus (Cuv.) / T. / M. Sallé / Brésil.

Troisième spécimen, n’appartient pas à la série-

type.

C.G. : 1863-114.

Autres numéros de catalogues : A.C. n° 1882 A,

N.C. n° 166.

Inscriptions sous le socle :

Brésil / M. Richard / 1863 n“ 114 / 1882 A / 1881

/ Coccyzus melanocorhynchus [sic] / melanorhyn-

chus / Cuv, / Type [même écriture].

Étiquette du socle :

Coccyzus melanocorhyphus [sic] / C. mdanorhyn-

chus (Cuv.) / T./ M. Richard / Brésil.

Dans sa description, Lesson indique que le spéci¬

men décrit est originaire du Brésil et qu’il a été

donné par Auguste de Saint I lilaire. Il s’agit donc

du premier spécimen. Rien ne permet d’affirmer

que Lesson se soit aussi servi du second spéci¬

men, rapporté un an plus tôt par un autre voya¬

geur, Sallé. Le troisième spécimen enfin n’a été

rapporté au Muséum qu’en 1863, longtemps

après la publication du travail de Lesson, par

Richard. La mention « type ■> sous les socles du

second et du troisième spécimen a dû être ajoutée

par erreur lorsque lesdits socles ont été changés.

L’orthographe originale du nom donné à cette

espèce par Vieillot (1817) est bien meldcorypbus ,

et non melanocoryphus. Il s’agit probablement

d’une erreur d’inadvertance.

Piaya braebyptera Lesson, 1831.

Lesson, Traité d’Ornithologie : 140.

Piaya melanogaster (Vieillot, 1817).

Premier spécimen. Holotype par monotypie.

C.G. : 1997-1227.

Autres numéros de catalogues : A.C. n° 1891,

N.C. n° 185.

Inscriptions sous le socle :

Cayenne / Leblond / 1891 / Piaya mclanogastra /

V / P. braebyptera / Less. Type [même écriture].

Étiquette du socle :

Piaya mclanogastra / (V.) / P. brachyprera (Less.)

T. / M. Leblond / Cayenne [même écriture].

Deuxième spécimen, n’appartient pas à la série-

type.

C.G. : 1997-1226.

Autres numéros de catalogues : A.C. n° 1890,

N.C. n° 184.

Inscriptions sous le socle :

1. Sur rectangle de papier colle : Brésil [écriture

plus récente) / Castelnau / et Deville.

2. 1890 / Piaya / melanogastra / (Type). (V) [écri¬

ture ancienne, encre pâlie et assez effacée, le V

recouvre un l.[.

3. melanogastra / Vieill. [au crayon).

4. P. Braebyptera / Less. ! [recouvre en partie l’ins¬

cription précédente, même écriture que 2],

Étiquette du socle :

Piaya nielanogustrra / (V.) / P. brachvptera (Less.)

T. de Castelnau / Brésil.

Lesson Indique que le spécimen d’après lequel il

décrit son Piaya braebyptera est originaire de

Cayenne et a été procuré par Leblond. 11 s'agit

donc du premier spécimen, Au contraire, les

incriptions sous le socle indiquent que le second

spécimen a été rapporté par Castelnau. Il ne peut

donc pas être un des types de Piaya braebyptera

Lesson car 1 expédition de Castelnau en

Amérique du Sud eut lieu de 1843 à 1847 alors

que la description de Lesson est parue en 1831.

Le type de Cuculus melanogaster Vieillot, 1817 se

trouve au Nationaal Natuurhistorisch Muséum

de Leiden (Hoek Ostende et al. 1997)-

Coccycua monaebus Lesson, 1831. Holotype par

monotypie.

Lesson, Traité d’Ornithologte : 142.

Piaya minuta minuta (Vieillot, 1817).

C.G. : 1997-1240.

Autres numéros de catalogues : A.C. n° 1894,

N.C. n" 175.

Inscriptions sous le socle :

Cayenne / M. Brocheton / 1894 / Piaya minuta /

V. / C. monachus. / Less. Typ. / [même écriture].

Étiquette du socle :

ZOOSYSTEMA • 1999 • 21 (2)

393

Voisin C. & Voisin J.-F.

Piaya minuta (Vieil!.) / C. monachus (Less.) Type

/ M. Brocheton Cayenne.

Dans son travail, Lesson indique que le spccimen

qu’il décrit se trouve à la Galerie du Muséum.

Bubutus isidori Lesson, 1831.

Lesson, Traitéd Ornithologie : 143.

Rhinortba chlorophaea chlorophaea (Raffles,

1822).

Premier spécimen. Lectotype présente désigna¬

tion.

C.G. : 1997-1233.

Autres numéros de catalogues : A.C. n° 1745,

N.C. n° 244, 73.

Inscriptions sous le socle :

Sumatra / par M Duvaucel / 1821 n° 73 / Bubutus

/ Isidori / (Lesson type) / Rhinortha / chlorophaea

I (Raffl) / 1745 [même écriture].

Etiquette du socle :

Rhinortha chlorophaea (Raffl.) / C. Isidori! (Less.)

Type / Duvaucel 1 Sumatra.

Deuxième spécimen, l’ara lectotype.

C.G. : 1997-1237.

Autres numéros de catalogues : A.C. n° 1744,

N.C. n° 245, 73.

Inscriptions sous le socle :

Sumatra / M. Duvaucel / 1821 n° 73 / 1744 [encre

pâlie] / Rhinortha chlorophaea / Raffl. / B. isidori.

Less / J ype [même écriture].

Etiquette du Socle :

Rhinortha chlorophaea (Raffl.) / C. Isidorii (Less.)

Type / Duvaucel / Sumatra.

Les deux spécimens portent le numéro 73.

L’espèce est dédiée à Isidore Geoffroy Saint-

Hilaire.

Bubutus Duvaucelii Lesson, 1831.

Lesson, Traité d’Ornitbôlogie : 143.

Rhinortba cbloropbaea chlorophaea (Raffles,

1822).

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. : 1997-1231.

Autres numéros de catalogues : A.C. n° 1742

bis, N.C. n° 246.

Inscriptions sous le socle :

Sumatra / M. Duvaucel 1821 / 1742 bis /

Rhinortha chlorophaea / B. Duvaucellii [sic] Less. /

Type [même écriture].

Etiquette du socle :

Rhinortha chlorophaea (Raffl.) / C. Duvaucelii

(Less.) Type / Duvaucel / Sumatra.

Deuxième spécimen. Paralectotype.

C.G. : 1997-1221.

Autres numéros de catalogues : A.C. n° 1743,

N.C. n°247.

Inscriptions sous le socle :

Sumatra / M. Duvaucel / 1821 / n°73 / 1743 /

Rhinortha chlorophaea / Raffl. / B. Duvaucelii /

Les. Type [mêmeécriture].

Etiquette du socle :

Rhinortha chlorophaea (Raffl.) / C. Duvaucelii

(Less.) Type / Duvaucel / Sumatra.

Ce spécimen est numéroté 73, ce qui indique

peut-être une certaine hésitation quant à la dis¬

tinction spécifique entre Bubutus isidori et ce

taxon. Dans sa description, Lesson indique que

B. duvaucelii habite Sumatra et que ces spéci¬

mens ont été envoyés par Duvaucel.

Coccyccus erythrorhynchus Cuvier in Lesson,

1831.

Cuvier in Lesson, Traité d’Omitbologie : 141.

Zanclostomus javanicus javanicus (Horsfield,

1821).

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. : 1997-1242.

Autres numéros de catalogues : A.C. n° 1756,

N.C. n°240.

Inscriptions sous le socle :

Java / M. Diard / 1821 / 1756 / Zanclostomus

javanicus / Horsf. / P. erythrorynchus Cuv. / Type

[même écriture).

Étiquette du socle :

Zanclostomus javanicus / (Hor.) / P. crythrorhyn-

chus (Cuv.) T. / M. Diard Java.

Deuxième spécimen. Paralectotype.

C.G. : 1997-1241.

Autres numéros de catalogues : A.C. n° 1755,

N.C. n°239.

Inscriptions sous le socle :

394

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

Bengale / M. Diard et / Duvaucel / 1821 n° 156/

1755 / Zanclostomus javanicus / H. / P. erythro-

rhyncha (peu lisible car repassé] Cuv. / Type

[même écriture].

Etiquette du socle :

Zanclostomus javanicus / (Hor.) / P. erythrorhyn-

chus (Cuv.) T. / Duvaucel Bengale.

Seul le premier spécimen porte une mention

d’origine exacte (Java).

Piaya chrysogaster Lesson, 1831. Holotype par

monotypie.

Lesson, Traité d'Qtnithologie : 140.

Zanclostomus javanicus javanicus (Horsfield,

1821).

C.G. : 1997-1243.

Autres numéros de catalogues : A.C. n° 1757,

N.C. n° 241.

Inscriptions sous le socle :

Cayenne / 1757 / Zanclostomus javanicus / Horsf.

/ P. chrysogaster / Cuv. et Less. / Type [même écri¬

ture].

Etiquette du socle :

Zanclostomus javanicus / P. chrysogastra (Cuv. &

Less.) T. / Cayenne.

Ce spécimen est bien un Zanclostomus javanicus

(Horsfield, 1821) comme les deux précédents.

C’est certainement l'erreur sur sa localité d’origine

(Cayenne au lieu de Java) qui a conduit Lesson

dans sa description à le considérer comme une

espèce à part, comme le montrent les mentions

» Ressemble étonnamenr par scs couleurs ec par

sa taille au piâye à bec rouge f- Zanclostomus

javanicus | » et « habite la Gujane [sicj ». Lesson

ne mentionne pas Cuvier comme auteur du

taxon P. chrysogaster., et ce doit être à la suite

d’une erreur que son nom figure sous le socle.

Pboenicophaus barrotii Eydoux & Souleyet,

1841. Holotype par monotypie.

Eydoux A Souleyet, Voyage autour du monde

exécuté pendant les années 1836 et 1837 sur la

Corvette La Bonite, Zoologie : 89, Atlas pl. 6.

Lepidogrammus cumingi (Fraser, 1839).

C.G: 1997-1199.

Autres numéros de catalogues : A.C. n° 1767,

N.C. n° 224,

Inscriptions sous le socle :

Philippines / M. Barrot / 1839 / 1767 Dasylophus

Cutningîi / I ras . / Ph. Barroti. Eyd. & Soûl. /

Type, [même écriture, encre très pâlie par

endroits].

Étiquette du socle :

Dasylophus Cumingii / (Fras.) / Ph. Barroti (E. &

S.) T. / M. Barrot / Philippines / 224.

Le genre Pha.enicopha.eus Stephens, 1815, est

orthographié Pboenicophaus par Vieillot (1816)

et plusieurs auteurs subséquents. 11 ressort du

texte de la description de Ph. barrotii quelle a été

faite sur un seul spécimen.

Crotopbaga Casasii Lesson in Lesson & Garnot,

1828. Lectotype, présente désignation.

Lesson ht Lesson & Garnot, Voyage autour du

Monde exécuté par ordre du Roi sur la Corvette de

Sa Maj. La Coquille pendant les années 1822,

1823, 1824 et 1825, Zoologie 1 (2) : 619.

Crotophagn su Ici ras tris sulcirostris Swainson,

1827.

C.G. : 1997-1230

Autres numéros de catalogues : A.C. n° 1857,

N.C. n° 369,

Inscriptions sous le socle :

1. 10 Dr Coquille |au crayon],

2. Crorophaga / sulcirostris / (Sw) / C. Casasii /

(Less. et Gain) / T. / Lesson et Garnot / Pérou /

1 857 (autre écriture].

Étiquette du socle :

Croiophaga sulcirostris / C. Casasii (Less. &

Gain.) T. / I.a Coquille / Pérou.

Le tome I, Zoologie , du « Voyage de La Coquille »

comprend deux parties dont la première est

parue en 1826 er la seconde en 1828. La descrip¬

tion du Crotopbaga Casasii se trouve dans certe

dernière, et est donc parue après celle du C. sul¬

cirostris de Swainson. La description de Lesson se

termine par « nous en tuâmes plusieurs

individus », qu’il a donc eu à sa disposirion pour

faire son travail.

Dromococcyx mexicana Bonaparte, 1856.

Holotype par monotypie.

Bonaparte, Comptes Rendus hebdomadaires des

Séances de l'Académie des Sciences de Paris XI II :

957.

Dromococcyx phasianellus phdsianellus (Spix,

1824).

C.G. 1856-1025.

ZOOSYSTEMA • 1999 • 21 (2)

395

Voisin C. & Voisin J.-F.

Autres numéros de catalogues ; A,C. n° 1872,

N.C. n° 197.

Inscriptions sous le socle :

1. o Mexique / Acq à M. Sallée [le deuxième e

barré] / 1856 n° 1025 Cat. Gen.

2. B. ntexicana / (Bp type).

3. Dromococcvx / phasianellus / (Spix) / Type /

1872 - 12 .

Etiquette du socle :

Dromococcvx / phasianellus (Spix.) / M. Salle /

Mexique.

S’il n’est pas un lapsus calami , le B devant ntexi-

cana est peut-être une référence à Bonaparte, qui

n’a d’ailleurs décrit ce taxon que très succincte¬

ment.

Centropus savorgnani Oustalct, 1886.

Oustalet, Le Naturaliste 8 (38) : 299.

Centropus anselli Sharpe, 1874.

Premier spécimen. Lectotype présente désigna¬

tion.

C.G. 1886-256.

Autre numéro de catalogue : N.C. n° 272.

Inscriptions sous le socle :

Gabon / Franceville / M.S.de Brazza / C.G. 1886

n°256 là/ Centropus savorgnani / Oust. / Type /

Paraphe.

Étiquette du socle :

Centropus savorgnani / (Oust.) / Type / M. de

Brazza / Gabon.

Deuxième spécimen. Paralectotype.

C.G. 1886-257.

Autres numéros de catalogues : N.C. n° 273.

Inscriptions sous le socle :

Congo / M. S. d. Brazza / 1886 n°257 / juv. /

Centropus savorgnani / Oust. / Congo / M. S. d.

Brazza / 1886 fi 0 257 / Paraphe.

Etiquette du socle :

Centropus savorgnani / (Oust.) / Type / M. de

Brazza / Gabon.

Les inscriptions sous les deux socles sont entière¬

ment de la même main et signées du même

paraphe en forme de T et de J accolés. Le pre¬

mier spécimen est un adulte et le second un juvé¬

nile avec encore du duvet, conformément à la

description d’Oustalet.

Centropus Lafresnayanns , Verreaux in Vinson,

1865. Lectotype. présente désignation.

Verreaux in Vinson, Voyage à Madagascar au

couronnement de Radanta II 1865, Annexe B :

3, 5.

Centropus toulou toulou (P. L. S. Millier, 1776).

C.G. : 1997-1244.

Autres numéros de catalogues : A.C. n° 1792,

N.C. n u 263.

Inscriptions sous le socle :

L Madagascar / par M. Sganzin / 1832.28.

2. Cenrropus Madagascariensis / (Bris.) / C.

Lalresnayus (Ver) / Type.

3. Décrit sous le / nom de Cucttlus toulou dès

1776.

Étiquerte du socle :

Centropus / madagascariensis (Briss.) /

C. T afresnaytis (Verr.) Type / M. Sganzin /

Madagascar.

J. Verreaux (1865) a établi sa description sur ce

spécimen mâle qui se trouvait à l’époque dans le

magasin de son frère, E. Verreaux, et de deux

spécimens femelles de la collection de Lafresnaye,

actuellement disparus. En ce qui concerne la

mention Centropus madagascariensis , voir plus

bas à ce taxon.

Centropus pianilus Lesson, 1831.

Lesson, Traité a Ornithologie : 136.

Centropus bengalensis javanensis (Dumont,

1818),

Premier spécimen. Lectotype, présente désigna¬

tion.

C.G. : 1997-1228.

Autres numéros de catalogues : A.C. n° 1782,

N.C. n° 294.

Inscriptions sous le socle :

Java i M. Diard / 1821 / 1782 / Centrococcyx

javanensis / Dum. / C. pumilus Less. / Type

[meme écriture].

Étiquette du socle :

Centrococcyx javanensis / Dum. / C. pumilus

(I.ess.) P. / M.Diard / Java.

Deuxième spécimen. Paralectotype.

C.G. 1997-1229.

Autres numéros de catalogues : A.C. n° 1783,

N.C. n‘* 295.

Inscriptions sous le socle :

396

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

Java / M. Leschenault / 1807 / 1783 / Centro-

coccyx javanensis / Dum. / C. pumilus Less. /

Type [môme écriture].

Etiquette du socle :

Centrococcyx javanensis / Dum. / C. pumilus

(Less.) T. / M. Leschenault / Java.

Sous les deux socles, les mentions sont de la

même main et ont la même disposition. Ces

deux spécimens, rapportés de Java par

Leschenault et Diard, ont servi à Lesson pour sa

description du mâle. Au contraire, nous n’avons

pas pu identifier le ou les spécimens qui lui ont

servi à décrire la femelle, et qui provenaient de

Sumatra.

Centropus affinis Lesson, 1831. Holotype par

monotypie

(= Centropus affinis Horsfield, 1821).

Lesson, Traité (l'Ornithologie : 136.

Centropus bengalensis javanensis (Dumont,

1818).

C.G. : 1997-1236.

Autres numéros de catalogues : A.C. n u 1780,

N.C. n° 292.

Inscriptions sous le socle :

Java / M. Diard 1821 / 1780 / Centrococcyx. /

Centropus. affinis / Less. / Type [même écriture].

Étiquette du socle :

Centrococcyx affinis / (Horsf.) / C. affinis (Less.)

T. / M. Diard / Java.

I^esson (1831) ne donne aucune référence avant

sa description de Centropus affinis , comme si la

description de Horsfield lui était restée inconnue.

Il s’agit sans doute d'un oubli, mais la stricte

application des dispositions du Code aboutit à

considérer C. affinis Lesson, 1831 à la fois

comme un synonyme et un homonyme plus

récent de C. affinis Horsfield, 1822.

Spécimen dont la qualité de type ne peut

ÊTRE ÉTABLIE AVEC CERTITUDE

Cuculus tenuirostris Lesson, 1831.

Lesson, Traité (/ ‘Ornithologie : 146.

Cuculus varias Vahl, 1 “97.

C.G. /: 1997-1212.

Autres numéros de catalogues : A.C. n° 1951,

N.C. n° 74.

Inscriptions sous le socle :

1. Bengale / %r 1822 / Hierococcys varius / Vahl.

/ 1951.

2. est-ce le type / de Cuculus tenuirostris / Lesson ?

/ J. Berlioz, [écriture et signature de J. Berlioz]

Étiquette du socle :

flierococcyx varius / (Vahl.) / Bengale.

Le bec de ce spécimen est endommagé. Le socle,

refait à la fin du siècle dernier, ne porte aucune

mention précisant qu’il s’agit du type de Lesson,

et le nom du collecteur n'est pas non plus indi¬

qué. Ces données se sont elles perdues ? Dans sa

description, Lesson indique que les spécimens du

Bengale qu’il a examinés sont au Muséum de

Paris et qu’ils ont été récoltés par Duvâucel. La

date de collecte ou d arrivée de cet exemplaire

(nov. 1822) correspond en effet à la période pen¬

dant laquelle Duvâucel faisait parvenir au

Muséum des collections du Bengale. Ce spéci¬

men est donc probablement un dés types de

Cuculus tenuirostris Lesson, niais les mentions

sous le socle sont trop incomplètes pour que l’on

puisse en avoir la certitude, conclusion à laquelle

J. Berlioz était visiblement déjà arrivé avant nous.

Lesson disposait certainement de plusieurs spéci¬

mens pour son travail, car il décrit successive¬

ment le plumage de l’adulte et du « jeune âge »,

pour terminer par une •< variété de Timor » qui

serait <« peut-être distincte comme espèce ».

Spécimens considérés à tort comme des

TYPES

Centropus menebiki Garnot in Lesson &

Garnot, 1828.

C.G. 1997-1194.

Piaya braebyptera Lesson, 1831.

C.G. : 1997-1226.

Piaya melanorbynchus Lesson, 1831.

C.G. : 1868-114 et 1997-1232.

Cuculus edolius Cuvier, 1817.

C.G. : 1997-1215.

Melias Diarcli Lesson, 1831.

C.G. : 1997-1202.

ZOOSYSTEMA • 1999 • 21 (2)

397

Voisin C. & Voisin J.-F.

Taccocua Lescbenaulti Lesson, 1831.

C.G.: 1997-1263

Les six spécimens ci-dessus ont déjà été traités à

propos des séries-types auxquelles ils n’appartien¬

nent pas.

Cuculus Levaillantii Lesson, 1831.

Lesson, Traité (/‘Ornithologie : 148.

(= Coccyzus Levaillantii Swainson, 1829).

Clama tôt cafer (Lichtenstein, 1793).

C.G. : 1997-1245.

Autres numéros de catalogues : A.C. n° 1923,

N.C. n° 153.

Inscriptions sous le socle :

Sénégal / M Sarvigny [le r caché par une tache

d’encre noire plus récente] / 1824 / 1923 /

Coccyzus cafer. Llcht. / Cucii. Levaillantii / Less.

Type [même écriture].

Étiquette du socle :

Coccyte* caler (Licht.) / C. Levaillanti (Less.) T /

M. Savigny.

Comme pour les taxons précédents, Lesson

(1931) cite la description de Swainson en préam¬

bule à la sienne, ce qui prouve qu’il la connais¬

sait.

Surniculus Itigubris Lesson, 1831.

Lesson, Traitéd'Ornithologie : 151.

Surniculus lugubris lugubris (Horsfield, 1821).

Premier spécimen.

C.G, : 1997-1222.

Autres numéros de catalogues : A.C. n° 1987,

N.C. n° 139.

Inscriptions sous le socle :

Java / M. Diard / 1821 / 1987 / Surniculus

lugubris / Horsf. / T. / 1987 [même écriture].

Étiquette du socle :

Surniculus lugubris / (Horsf.) / Type /

M. Diard / Java.

Deuxième spécimen.

Numéro de catalogue général : 1997-1223.

Autres numéros de catalogues : A.C. n° 1985,

N.C. n“ 14(1.

Inscriptions sous le socle :

Java / M. Diard / 1821 / 1985 / Surniculus lugu¬

bris / Horsf. [même écriture].

Étiquette du socle :

Surniculus lugubris / (Horsf.) / Type / M. Diard /

Java,

Conformément à son habitude, Lesson (1931)

cite la publication de Horsfield avant de com¬

mencer sa propre description de Surniculus lugu¬

bris, Les deux spécimens ci-dessus n'ont d’autre

titre que d’avoir servi à Lesson pour celle-ci, ainsi

que pour définir le genre Surniculus. Ils ne sont

pas indiqués comme types dans l'Ancien

Catalogue.

Piaya minor Lesson, 1831.

Lesson, Traité d’Ornithologie : 141.

Coccyzus minor (Gmelin, 1788).

C.G. : 1997-1209.

Autres numéros de catalogues : A.C. n° 1878,

N.C. n" 171.

Inscriptions sous le socle :

Porto Rico / M. Maugé / 18~8 / Coccyzus minor /

Grn. I Piaya minor / Lesson / Typ. [même écri¬

ture].

Étiquette du socle :

Coccyzus minor / (Gm.) / M. Maugé / Porto Rico.

Lesson (1831) cite Gmelin en préambule à sa

description de Piaya minor. ce qui prouve bien

qu’il était au courant de la description qu’en avait

faite cet auteur. Ce spécimen n’a d’autre mérite

que d'avoir servi à Lesson pour établir une seconde

description de l'espèce.

Piaya atnericana Lesson, 1831.

Lesson, Traité d'Ornithologie : 142.

Coccyzus arnericanus arnericanus (Linné, 1758).

C.G. : 1997-1238.

Autre? numéros de catalogues : A.C. n° 1873,

N.C. n°162.

Inscriptions sous le socle :

Antilles / en 1810 / Coccyzus Américanus [sic] /

(L) / 1873 / P. arnericanus / (Less type) [même

écriture].

Étiquette du socle :

Coccyzus arnericanus / P. arnericanus (Less.) Type

I Antilles.

En préambule à sa description de l’espèce, Lesson

(1831) donne les références de plusieurs descrip¬

tions autres, faites avant la sienne sous les nom

à’arnericanus ou de carolinensis. Ce spécimen

398

ZOOSYSTEMA ■ 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

n'est donc pas un type, et n'est d'ailleurs pas

mentionné comme tel dans l'Ancien Catalogue.

Phoenicophaetis curvirostris Shaw, 1810.

(= Cuculus curvirostris Shaw et Nodder, 1810).

Shaw &c Nodder, The Naturalisés Miscellany

21:905.

Rhamphococcyx curvirostris curvirostris (Shaw,

1810).

Premier spécimen.

C.G. : 1997-1210.

Autres numéros de catalogues : A.C. n° 1760,

N.C. n°2l4.

Inscriptions sous le socle :

Java / par M. Diard / 27 X bre 1821 / Phoeni-

cophaeus / curvirostris / (Shaw) / Type 1760

[même écriture].

Etiquette du socle :

Phoenicophaeus / curvirostris (Shaw) / T. / M.

Diard / Java.

Deuxième spécimen.

C.G. : 1997-1211.

Autres numéros de catalogues : A.C. n° 1760,

N.C, n° 215.

Inscriptions sous le socle :

Java / par M. Diard / X bre 1821 / Phoeni-

cophaeus / curvirostris / (Shaw) / Type / 1761

[même écriture).

Étiquette du socle :

Phoenicophaeus / curvirostris (Shaw) / T. /

M. Diard / Java.

Les deux spécimens ci-dessus ne peuvent être les

types de Cuculus curvirostris , car ce taxon a été

décrit par Shaw &t Nodder en 1810, alors qu’ils

ont été collectés en octobre 1821. Peut-être a-t-

on oublié de recopier certaines données lorsque

l’on a refait leurs socles à la fin du siècle dernier,

ou bien ont-ils tout simplement été examinés

ultérieurement par Shaw ?

Melias pyrrhocephaltis Lesson, 1831.

Lesson, Traité (/'Ornithologie : 131.

Phaenicophaeuspyrrhocephaltis (Pennant, 1769).

Premier spécimen.

C.G. : 1997-1204.

Autres numéros de catalogues : A.C. n° 1759,

N.C. n° 222.

Inscriptions sous le socle :

Ceylan / 1822 / 1759 / Phoenicophaeneus [sic] /

pyrrhocephalus / Less. / Type [même écriture].

Étiquette du socle :

Phoenicophaeus / pyrrhocephalus (Less.) / T. /

Leschenault / Ceylan.

Deuxième spécimen.

C.G. : 1997-1203.

Autres numéros de catalogues : A.C. n° 1758,

N.C. n D 221.

Inscriptions sous le socle :

Ceylan / Type / Phoenicophaeus Pyrrhocephalus /

(Less.) / 1758 [même écriture].

Étiquette sur le socle :

Phoenicophaeus / pyrrhocephalus (Less.) / T. /

Ceylan.

D’après les catalogues, le premier spécimen a été

rapporté par Lcschenault en 1822. Lesson (1831)

cite plusieurs descriptions antérieures en préam¬

bule à la sienne, ce qui montre qu'il savait que

son Melias pyrrhocephalus avait déjà été décrit.

Phaenicophaus superciliosus Cuvier in

Dumont, 1823.

Dumont in Cuvier (ed.), Dictionnaire des

Sciences naturelles 28:451.

Dasylophus superciliosus (Cuvier in Dumont,

1823).

C.G. : 1997-1205.

Autres numéros de catalogue : A.C. n° 1765,

N.C. n°228.

Inscriptions sous le socle :

Philippines / par M. Philibert / Novembre 1822 /

1765 / Dasylophus / superciliosus / (Cuv) / Type /

12 [même écriture].

Étiquette du socle :

Dasylophus / superciliosus (Cuv.) T. / M. Philibert

/ Philippines / 228.

Dumont, qui attribue ce taxon à Cuvier, l’a

décrit dans le Dictionnaire des Sciences naturelles

dont ce même Cuvier était le rédacteur (1823).

On y lit : a Phoenicophatts superciliosus Cuv. Cet

oiseau des Philippines, dont il existe au Muséum

d’Histoire naturelle de Paris deux individus don¬

nés par M. Dussumier [,..] » (suit une descrip¬

tion précise de l'espèce). Le spécimen ci-dessus

n’a pas été collecté par Dussumier, mais par

Philibert en novembre 1822, ce qui, compte

ZOOSYSTEMA • 1999 • 21 (2)

399

Voisin C. & Voisin J.-F.

tenu des délais de transport de l'époque, laisse

douter qu’il fût arrivé à temps des Philippines à

Paris pour que Dumont puisse le décrire dans un

ouvrage paru dès 1823. De plus, aucun des

divers catalogues du Laboratoire des Mammifères

et Oiseaux ne mentionne de spécimen Pb. super-

ciliosus rapporté par Dussumier. Dans ces condi¬

tions, les éléments manquent pour conclure qu'il

s’agit de l’un des deux spécimens utilisés par

Dumont.

Cultrides geoffroyi Pucheran, 1845.

Pucheran, Revue de Zoologie : 51.

Neomorphusgeoffroyi (Temminck, 1820).

C.G. : 1997-1198.

Autres numéros de catalogues : A.C. n° 1829,

N.C. n° 211.

Inscriptions sous le socle :

Brésil / M. St Hilaire / 1822 / 1829 / Neomorphus

geollroyî / Pucheran / Type.

Étiquette du socle :

Neomorphus / Geoffroyi (Tem.) T. /

M.A.S. Hilaire / Brésil.

Temminck a décrit Coccyzus geoffroyi dès 1820,

donc avant que ce spécimen ne soit rapporté du

Brésil par A. Geoffroy Saint-Hilaire en 1822. Il

ne peut donc être un des types de ce taxon, mais

il a cependant servi à Pucheran pour décrire le

genre Cultrides.

Centropus philippensis l esson in Tesson &

Garnot, 1828.

Lesson ht Tesson & Garnot, Voyage autour du

Monde exécuté par ordre du Roi sur la Corvette de

Sa Maj. La Coquille pendant les années 1822,

1823, 1824 et 1825, Zoologie I (2) : 619.

Centropus sinensis eurycercus Blyth, 1845.

Premier spécimen.

C.G. : 1997-1208.

Autres numéros de catalogues : A.C. n° 1795,

N.C. n° 281.

Inscriptions sous le socle :

Bengale / M. Duvaucet et Diard / 1819 / 1795 /

Centrncoccyx eiirvcercus / Hay / C. philippensis /

Cuv. Type (même écriture].

Étiquette du socle :

Centrococcyx / eurycercus (Hay.) / C. philippensis

(Cuv.) Type / M. Diard / Java.

Deuxième spécimen.

C.G. : 1997-1207.

Autres numéros de catalogues : A.C. n° 1794,

N.C. u 0 282.

Inscriptions sous le socle :

Sumatra / M. Duvaucel / 1821 / 1794 /

Centrococcyx eurycercus / Hay / C. philippensis /

Typ. d. Cuvier [même écriture].

Etiquette du socle :

Centrococcyx / eurycercus (Hay.) / C. philippensis

(Cuv.) Type / Duvaucel / Sumatra.

Les tomes Zoologie du Voyage de La Coquille,

parus en 1826 et 1828, regroupent les études,

par différents auteurs, de l’important matériel

zoologique rapporté du voyage de cette corvette

autour du monde de 1822 à 1825. Lesson et

Garnot, qui avaient participé à l’expédition, en

rédigèrent la partie sur les oiseaux et n om visi¬

blement travaillé que sur ce matériel, dont les

deux spécimens ci-dessus ne font pas partie. Rien

non plus dans les inscriptions sous les socles ni

dans les catalogues ne permet d’affirmer que

Lesson les a étudiés. Cuvier n’a jamais non plus

décrit ce taxon, et les deux spécimens ci-dessus

n’ont probablement d’autre mérite que de lui

avoir peut-être servi lors de ses cours.

La synonymie et les homonymies de C. philip¬

pensis Lesson sont complexes et mériteraient une

étude particulière.

Centropus madagascariensis Scblegel, 1 864.

Schlegel, Revue méthodique et critique des collec¬

tions déposées dans cet établissement. Muséum

d’Histoire naturelle des Pays-Bas 1 (25) ; 65.

Centropus toutou undou (P.I..S. Muller, 1776).

C.G, : 1997-1206.

Aurres numéros de catalogues : A.C. n° 1793,

N.C. n" 264.

Inscriptions sous le socle :

Madagascar / M, Fouloy 1820 / 1793 / S /

Centropus madagascariensis / Briss. / Cucu. tolu.

Gin. / Type [même écriture).

Étiquette du socle :

Centropus madagascariensis / (Briss.) / â / C.

tolu. (Gm.) / M. Foulov / Madagascar.

Ce spécimen rapporté en 1820 ne peut être celui

sur lequel Schlegel (1864) fonde la diagnose de

400

ZOOSYSTEMA • 1999 • 21 (2)

Liste des types d’oiseaux du MNHN, Cuculidae

son Cm trop ils mtidagascariensis , car et dernier a été

acquis des frères Verrcaux en 1863 par le 's Rijlcs

Muséum van Natuurlijke Historié (aujourd’hui

Nationaal Natuuthistorisch Muséum, à Leiden). Il

ne peut pas non plus avoir été examine ni par

Gmclin en 1788 ni par [frisson en 1760. La men¬

tion « type » n’a d’ailleurs pas été reportée sur l’éti¬

quette, ni dans l’Ancien Catalogue. Le nom donné

par Brisson (1760) était « Coucou de Mada¬

gascar », en français.

Remerciements

Nous tenons à remercier ici le

Dr R. W. R. J. Dekker, du Nationaal

Natuurhistorisch Muséum, de Leiden (Pays-Bas),

le Dr R. Prys-Jones, le Dr M. Adams et le

Dr M. Walters, du Natural History Muséum,

Tring (Grande-Bretagne), M. A. Philippot, du

Musée de Zoologie de Nancy (France), le

Dr F. Vuilleumier, de l'American Muséum of

Natural History, New York (Etats-Unis), ainsi que

le Dr Chr. Jouanin et le Dr A, Ohler, du Muséum

national d’Histoire naturelle, Paris (France) pour

l'aide qu'ils nous ont apportée, tant sous la forme

de commentaires constructifs, de renseignements,

que de prêt de spécimens de collection.

RÉFÉRENCES

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affinity of Cuculus lepidus Millier. Ibis 1 17:

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Bonaparte Ch. 1856. — Notice sur les Tableaux des

Gallinacés. Complet Rendus hebdomadaires des

Séances de l'Académie des Sciences de Paris XLI1 :

953-959.

Brisson M.-J. 1760. — Ornithologie, ou Méthode

contenant ht division des oiseaux en ordres > sections,

genres, especes dr leurs variétés. A laquelle est joint une

description exacte de chaque espèce, avec les citations

des auteurs qui en ont traité, les noms qit 'ils leur ont

donnés, et que leur ont donné tes différentes nations,

& les noms vulgaires. Bauché, Parts, volume IV.

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Cuvier F. C. 1816. — Le Règne animal distribué

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425. 426

David-Beaulieu A. 1932. — Les Oiseaux de la région

de Honquan (Cochînchine). L'Oiseau et la Revue

française d'Ontithologte 2 : 133-154.

Davics Shetborn C. & Woodward B. B. 1906. —

Notes on the dates of publication of the natural

histoiy portions of sottie French voyages - Voyage

autour du Monde. , sur... la Coquille pendant...

1 822-25, The Armais and Magasine of Natural

History O) 17: 335-336.

Delaconr J. 193L - Description de neuf oiseaux

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çaise d'Ornithnlogie 1 : 473- 486.

Delacour J. tic Jabouille P. 1931. — Les oiseaux de

l'Indochine française. ?.. Exposition coloniale inter¬

nationale. Paris.

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7. Bertrand, Paris.

Deville F, 1851, — Notes sur quatre espèces nou¬

velles d'oiseaux de F expédition de M. Castelnau.

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Keijl G. O. 1997- — Fvpe spécimens of birds in the

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the Linnean Society. London X\ll: 133-200.

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Lesson R. P. 1842. —-Notes sur les oiseaux nouveaux

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Lesson R. P. & Garriot 1828 — Voyage autour du

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Levaillant F. 1808. — Histoire naturelle des Oiseaux

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Oustalet K. 1886. — Notice sur quelques oiseaux

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(Thrcskiornithinés) de la collection du Muséum

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Revue française WOrnhhuIogie 63 : 45-53.

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d’oiseaux des collections du Muséum national

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de Paris. 1. Péléçanif’otmes. L'Oiseau et la Revue

française d’Omithologtc 62 : 162-172.

Voisin J.-F. 1995. — Litre provisoire des types

d’oiseaux des collections du Muséum national

d’i listoire naturelle de Paris. 3, Podicipédiformes.

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Liste des types d’oiseaux des collections du

Muséum national d'Histoire naturelle (France). 5.

Proccdlariiformes (avec une note sur les catalogues

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Wells D. R. & Beckîng J. H. 1975- — Vocalisations

and status of little and himalayan cuckoos, Cuculus

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Ibis 117: 366-371.

Soumis le 28 février 1998 :

accepté le 15 janvier 1999.

402

ZOOSYSTEMA • 1999 • 21 (2)

Instructions aux auteurs

Ligne éditoriale

Zoosystema est une revue trimestrielle consacrée à

l’inventaire, l’analyse et l’interprétation de la bio¬

diversité animale. F.lle publie des résultats originaux

de recherches en zoologie, particulièrement en sys¬

tématique et domaines associes : morphologie com¬

parative, fonctionnelle et évolutive ; phylogénie ;

biogéographie ; taxinomie et nomenclature.

Un numéro de Zoosystema pourra être consacré h

un thème particulier sous la responsabilité d’un édi¬

teur inv'ité.

Informations générales

Soumettre un article pour publication dans

Zoosystema implique que celui-ci n’ait pas été sou¬

mis dans une autre revue, l.es droits de reproduc¬

tion de l'article, y compris les illustrations, sonr

réserves à la revue. La reproduction de tout ou par¬

tie de l’article doit faire l'objet d’une demande écri¬

te préalable, adressée à la rédaction.

Les auteurs devront suivie strictement le Code

International de Nomenclature Zoologujue. II est

recommandé que le matériel-type soit, au moins en

partie, déposé dans les collections du MNHN. La

revue porte une attention particulière aux dates de

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ZOOSYSTEMA • 1999 • 21 (2)

403

Instructions aux auteurs

- pour les descriptions systématiques, chaque des¬

cription doit suivre l’ordre suivant : nom du taxon

avec auteur et date, synonymie, matériel-type, éty¬

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Références bibliographiques

Dans le texte, les références aux auteurs d articles

seront en minuscules, sans virgule avant l'année,

ex. : Dupont (2001), Dupont (2001, 2002),

(Dupont 200 1 ; Durand 2002), (Dupont &

Durand 2003, 2005), (Dupont, Durand &

Dupond 200.5), Dupont (2001 : 1 ; 2003 : 4),

Dupont (2001 : fig. 2).

(.es auteurs de taxons sont indiqués avec une virgu¬

le, ex. : Mursia coseli Crosnier, 1997 ;

Neoheligmonella mastomysi Dioul, Bà & Durette-

Desset, 1998.

Dans la bibliographie, les références seront présen¬

tées comme ci-dessous, dans l’ordre alphabétique.

Les noms de revues ne doivent pas erre abrèges :

Hoeg J. T. & Lützen J. 1985. — Comparative

morphology and phylogeny of the Family

Thompsoniidae (Cirripedia: Rhizocephala:

Akentrogonida) with description of three nevv

généra and seven new species. Zaologica Scripta

22: 363-386.

Rock cl D.. Korn W. & Kohn A. L 1995. —

Matinal of the living (Jonidae . volume 1 : Indo-

Pacific région. Christa Hemmen, Wiesbaden.

517 p.

Schwaner T. D. 1985. — Population structure of

black figer snakes, Notechis ater niger , on off¬

shore islands of South Australia: 35-46, in

Grigg G., Shinc R. & Ehmann H. (eds),

Bio/ogy of Australasidn Frogs and Reptiles. Surrey

Beatty and Sons, Sydney.

Épreuves et tirés à part

Les épreuves seront adressées au premier auteur er

devront être retournées corrigées sous huitaine. Les

corrections, autres que celles imputables à la rédac¬

tion ou à l’imprimeur, seront it la charge des

auteurs. Les auteurs recevront gratuitement vingt-

cinq tirés à part ; les tirés a part supplémentaires

seront à commander sur un formulaire joint aux

épreuves.

404

ZOOSYSTEMA • 1999 • 21 (2)

Instructions to authors

Scope ofthe Journal

Zoosystema is a quarrerly journal devoted to the

inventory, analysis and interprétation of animal

biodiversity. Il publishes original rcsults of zoologi-

cal research, particularly in systematics and related

fields: comparative, functional and evolurionary

morphologv; phylogenv; biogeography; taxonomy

and nomenclature.

A complété issue of Zoosystema may be devoted to

several papers on a single topic as the responsibility

of an invited editor.

General information

The submission of a manuscript to Zoosystema

implies that it is not being offered for publication

elsewhere. Copyright ofpublished paper, including

illustrations, becomes the property of the journal.

Requests to reproduce material front Zoosystema

should be addressed to the editor.

Papers should strictly foliow the International Code

of Zoologieal Nomenclature. Wc recommend that

the authors should déposa in the MNHN collec¬

tions at leasr a part of the type ntatcrial. The jour¬

nal pays spécial attention to publication dates; a

table is published with the lirst issue ofcach vcar.

Manuscripts, without limitation of the number of

pages, rnust conform strictly with the instructions

to authors and will be sent to the Editor:

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Zoosystema

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fax: (33) (0)1 40 79 38 58

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Manuscripts which do not conform to instructions

will be returned. Each manuscript is reviewed bv at

least two referees.

Manuscripts must be submitted in triplicate in A4

format, double spaced, with margins of at least

3 cm and ail pages numbered. The original figures

should be sent with the révisée) manuscript, as well

as a 3.5” diskette Apple Macintosh or IBM-compa-

tible (Word) format, which will also contain tables

(Word, Excel) and possibly figures (Adobe

Illustrator, Photoshop; Deneba Canvas; EPS).

Format

Papers are to be written in simple and concise

French or English. Manuscripts in English should

be organîzed as follows:

— title in English; this should not include the name

of new taxa;

— title in French (exact translation);

— running head in English;

— namefs) of author(s), followcd by their full pro-

fcssiônal address(cs) and, if possible, fax number

and e-mail;

— abstracts in English, 200-250 words long; new

taxa name.s should be included in the abstract; the

abstract should bc précise and descriptive, in order

to bc rcproduccd as such iu daia bases; avoid vague

sentences sucli as three new species are described”

or “spccics are compared ro species already known”;

include précisé différenciai charaaers;

— résume in French (where feasible), which should

be au exact translation of the English abstract; the

journal may provide Help for translation;

— key words in English (5 to 10);

— key words in French;

— text of the article, in lhe following order:

Introduction, Materials and methods,

Abbrcviations used, Observations, Discussion,

Acknowledgements, Référencés, l.egends ro Figures

and Tables; the arrangement of rhe parts “Obser¬

vations” and “Discussion” may be modulated

according to the length and subject of the article;

very long papers may include a list o! contents;

— for systematic description, each description

should follow the order: name ol taxon with aurhor

and date, synonymy, type material, etymologv,

material exattiined, distribution, diagnosis and/or

description, commenta; this order may be adapred

according to the zoologieal groups: consult a recent

issue o f Zoosystema;

ZOOSYSTEMA • 1999 • 21 (2)

405

Instructions to authors

- use n. sp., n. yen., n. fam., etc. for new taxa;

- use italics for généra and species names;

- references to illustrations and tables should be

indicated as follows: (Fig. 1), (Fig. 2A, D),

(Fig. 2A-C), (Figs 3; 6), (Figs 3-5); (Table 1);

- captions to illustrations, on a separate sheet,

should includc abbreviations and scale values (e.g.:

scale bar; 1 cm);

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Line drawings must be in Indian ink or high quality

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Plates are not placcd at the end of the article: they

will be considered as figures and numbered as sueh.

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for each figure, are to be indicated on an accompa-

nying overlay, not on the original figure. They will

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each figure. No diagram or table is to exceed one

page; longer tables should be divided. Color plates

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References

In main text, references to authors, in lower case,

should be presented without comma before year, as

follows: Smith (2001), Smith (2001,2002), (Smith

2001), (Smith 2001; Jones 2002), (Smith & Joncs

2003, 2005), (Smith, Jones & Johnson 2003),

Smith (2001; I; 2003: 5), Smith (2001: fig. 2).

Authors of taxa arc indicated with a comma, e.g.;

Mursia coseli Crosnicr, 1997; Neoheligmonella mas-

tomysi Diouf, Bâ & Durette-Desset, 1998.

References should be presented as follows, in alpha-

betical order. Do not abbreviate journal names:

Hoeg J. T. & Lützen J. 1985. — Comparative

morphology and phylogeny of the family

Thompsoniidae (Cirripedia: Rhizoccphala:

Akentrogonida) with description ot threc new

généra and seven new species. Zooloeica Scripta

22: 363-386.

Rôckel D., Korn W. ôc Kohn A. J. 1995. —

Manudl of the living Conidae , volume 1 : Indo-

I’acific région. Christa Hemmen, Wiesbaden,

517 p.

Schwaner T. D. 1985. — Population structure of

black tiger snakes, Notechis ater niger, on off¬

shore islands of South Australia: 35-46, in

Grigg G., Shine R. & Ehntann H. (eds),

Biology ofAustralasian Frogs and Reptiles. Surrey

Beatty and Sons, Sydney.

Proofs and reprints

Proofs will be sent to the first author for correction

and must be returned within eight days. Authors

will be charged for excessive corrections. Authors

will reccive twenty-five reprints free of charge; fur-

ther reprints can bc ordered, at a charge indicated,

on a form supplied with the proofs.

406

ZOOSYSTEMA • 1999 • 21 (2)

Mise en page

Noémie de la Selle

Packaging Editorial

Achevé d’imprimer

sur les presses de l’Imprimerie Durand

28600 Luisant (France)

Juin 1999

Dépôt légal n° 11150

Printed on acid-free paper

Imprimé sur papier non acide

Date de distribution du fascicule n° 1, 1999:

17 mars 1999

Couverture : trichomes dorsaux de Polyxenus lagurus (Linné, 1758),

Myriapode, Diplopode, Penicillate

Photographie R. Cleva (MNHN, Laboratoire Zoologie-Arthropodes)

1999 • 21 ( 2 )

zoosystema

^MÜSECMÏ

'NAT ION Al)

IdHIST.NAT.

Cuif ).-P. & Perrin C.

Micromorphology and microstructure as expressions of scleractinian skeletogenesis in Favia fragum

(Esper, 1 795) (Faviidae, Scleractinia)

Neifar L., Euzet L. & Ben Hassine O. K.

Trois nouveaux Heterocotyle (Monogenea, Monocotylidae) parasites branchiaux de Taeniura grabata

(Euselachii, Dasyatidae) en Tunisie

Medinskaya A. I.

Foregut anatomy of the Cochlespirinae (Gastropoda, Conoidea, Turridae)

Simone L. R. L.

Comparative morphology and systematics of Brazilian Terebridae (Mollusca, Castropoda, Conoidea),

with descriptions of three new species ’WQK' > T

Ituarte C. F.

Pisidium chilense (d'Orbigny, 1846) and new species of Pisidium C. Pfeiffer, 1821 from Southern Chiie

(Bivalvia, Sphaeriidae)

Vidal |.

Taxonomie review of the eiongated cockles: généra Trachyca rdium, Vasticardium and Acroster igma

(Mollusca, Cardiidae)

Lourenço W. R. & Monod L.

Confirmation de la validité du genre Hormiops Fage, 1933 avec redesc ription d 'Hormiops davidovi

Fage, 1933 (Scorpiones, Ischnuridae) m

Crosnier A.

Un Heterocarpus nouveau (Crustacea, Decapoda, Pandalidae) du Pacifique Sud-Ouest

Clark P. F. & Ng P. K. L.

The identity of Chlorodius miliaris A. Milne Edwards, 1873, and the establishment of a new genus of

Chlorodiinae (Crustacea, Decapoda, Brachyura, Xanthoidea, Xant hida e ) from New Caledonia

Mauriès J.-P. & Kime R. D.

Description, écologie et chorologie de trois espèces nouvelles de diplopodes (Myriapoda, Diplopoda) |

des zones périphériques du Massif central et du centre de la France

Voisin C. & Voisin J.-F.

Liste des types d'oiseaux des collections du Muséum nationa l d'His toire naturelle de Paris. 6. Coucousl

(Cuculidae) ~

Conception Graphique : Isabel Gautray

Publication trimestrielle, juin 1999 / Quarterly, )une 1999

ISSN: 1280-9551

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