Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Comité DE REDACCIÓN (BOARD OF ÉDITORS)

EDITOR DE PUBLICACIONES (EDITOR-IN=CHIEF)

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EDITORA EJECUTIVA (MANAGING EDITOR)

Eugenia M* Martínez Cueto-Felgueroso Apdo. 156, Mieres del Camino, Asturias, España

EDITORES ADJUNTOS (ASSOCIATE EDITORS)

Benjamín Gómez Moliner Universidad del Poís Vasco, Vitoria, España

Ángel Antonio Luque del Villar Universidad Autónoma de Madrid, Madrid, España

Emilio Rolán Mosquera Universidad de Vigo, Vigo, España

José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Jesús S. Troncoso Universidad de Vigo, Vigo, España

Comité EDITORIAL (BOARD OF REVIEWERS)

Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España

Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España

Rafael Araujo Armero Museo Nacional de Ciencias Naturales, Madrid, España

Thierry Backeljau Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

Ridiger Bieler The Field Museum, Chicago, Estados Unidos

Sigurd v. Boletzky Loboratoire Arago, Banyuls-sur-Mer, Francia

Jose Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España

Karl Edlinger Noturhistorisches Museum Wien, Viena, Austria

Antonio M. de Frias Martins Universidade dos Acores, Acores, Portugal

José Carlos García Gómez Universidad de Sevilla, Sevilla, España

Gonzalo Giribet de Sebastián Harvard University, EE.UU.

Edmund Gittenberger National Natuurhistorisch Museum, Leiden, Holanda

Serge Gotas Universidad de Málaga, España

Angel Guerra Sierra Instituto de Investigaciones Marinas, CSIC, Vigo, España

Gerhard Haszprunar Zoologische Staatssammlung Múnchen, Múnchen, Alemania

Yuri 1. Kantor AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia

María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España

Jordi Martinell Calico Universidad de Barcelona, Barcelona, España

Ron K. 0'Dor | Dalhousie University, Halifax, Canada

Tokashi Okutani * : Nihon University, Fujisawa City, Japón

Marco Oliverio Universitá di Roma “La Sapienza”, Roma, Italia

Pablo E. Penchaszadeh * Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

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Carlos Enrique Prieto Sierra + Universidad del País Vasco, Bilbao, España

M* de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España

Paul G. Rodhouse * e - British Antarctic Survey, Cambridge, Reino Unido

Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España

María Carmen Salas Casanovas Universidad de Málaga, Málaga, España

Gerhard Steiner Institut fir Zoologie der Universitút Wien, Viena, Austria

Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España

Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia

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Four new species of Latirus (Gastropoda: Fasciolariidae)

from the Philippine Islands and the southern Caribbean

Cuatro nuevas especies de Latirus (Gastropoda: Fasciolariidae) de

Filipinas y el Caribe sur

Martin Avery SNYDER*

Recibido el 15-111-2002. Aceptado el 1-VI-2002

ABSTRACT

This paper describes four Latirus species in the neogastropod family Fasciolariidae, Latirus

cloveri, Latirus sarinae, and Latirus philippinensis, all from the Philippine Islands, and

Latirus abbotti from the southern Caribbean. Latirus cloveri is distinguished from Fascio-

laria walleri Ladd, 1976, also from the Philippine Islands, Latirus sarinae from Latirus

kandai Kuroda, 1950 from the western Pacific, Latirus philippinensis from Latirus elsiae

Kilburn, 1975 from South Africa, and Latirus abbotti from Latirus angulatus (Róding,

1798) from the Caribbean.

RESUMEN

Se describen cuatro especies de Latirus pertenecientes a la familia Fasciolariidae, Latirus

cloveri, Latirus sarinae, y Latirus philippinensis, procedentes de Filipinas, y Latirus abbotti

del Caribe sur. Latirus cloveri se diferencia de Fasciolaria walleri Ladd, 1976, también de

Filipinas, Latirus sarinae de Latirus kandai Kuroda, 1950 del Pacífico oeste, Latirus philip-

pinensis de Latirus elsiae Kilburn, 1975 de Sudáfrica, y Latirus abbotti de Latirus angulatus

(Róding, 1798) del Caribe.

KEY WORDS: Mollusca, Gastropoda, Fasciolariidae, Latirus, new species.

PALABRAS CLAVE: Mollusca, Gastropoda, Fasciolariidae, Latirus, nuevas especies.

INTRODUCTION

These four new species of Latirus

from the Philippine Islands and from

the Caribbean are new discoveries from

areas where several new species of

Latirus have been described in recent

years. Tangle net collecting in the Philip-

pine Islands has resulted in the discov-

ery of many new molluscan species

including Latirus aldeynzeri Garcia, 2001,

L. balicasagensis Bozzetti, 1997 and L.

martinorum Cernohorsky, 1987. Many

new species have also been discovered

recently in the Caribbean, especially

around Honduras, including “Latirus

anapetes” (Woodring) Petuch, 1981, L.

martini Snyder, 1988 and L. cuna Petuch,

1990. Almost all of the material

described herein was or is part of the

author's collection. These specimens

were collected in the last 10 years, either

as a byproduct of commercial fishing

operations or by scuba diving.

* Department of Malacology, Academy of Natural Sciences, 19% and Benjamin Franklin Parkway, Philadelphia,

PA 19103, USA.

Iberus, 21 (1), 2003

Abbreviations used

ANSP Academy of Natural Sciences of

Philadelphia, Philadelphia, Pennsy]l-

vania, USA

EC Everson collection, Louisville, Ken-

tucky, USA

MNHN Muséum national d'Histoire

naturelle, Paris, France

SC Snyder collection, Villanova, Penn-

sylvania, USA

USNM National Museum of Natural

History, Smithsonian Institution,

Washington D.C., USA

SYSTEMATICS

Family FASCIOLARIIDAE Gray, 1853

Subfamily PERISTERNIINAE Tryon, 1880

Genus Latirus Montfort, 1810

Type species: Latirus aurantiacus Montfort, 1810, a synonym of Latirus gibbulus (Gmelin, 1791),

Recent, Indo-Pacific, by monotypy.

Latirus cloveri spec.nov. (Figs. 1, 2)

Type material: Holotype ANSP 408331, length 52.7 mm, in tangle nets, depth 150 m. Paratype 1,

MNHN, length 40.4 mm, subadult, in tangle nets, Balicasag Island, Bohol, Philippine Islands.

Paratype 2, SC, length 49.6 mm, from type locality. Paratype 3, SC, length 48.1 mm, from type

locality.

Other material examined: 1 adult specimen 64.5 mm and 3 immature specimens, 34.2 mm, 38.7

mm, 43.3 mm, the first apparently live collected, all SC. 38.7 mm specimen from type locality; oth-

ers from tangle nets off Panglao, Bohol, Philippine Islands.

Etymology: The species is named for Phillip Clover of Glen Ellen, California, a friend and shell

dealer, who provided much of the type material.

Type locality: Aliguay Island, off Dipolog, northwest Mindanao, Philippine Islands.

Description: Shell medium size for

genus, fusiform, length of adult specimens

from 48.1 mm to 64.5 mm. Siphonal canal

short, narrowly open, curved, often twist-

ed away from aperture. Smooth pearly

white protoconch of 1 */4 to 1 */2 whorls,

with 8-9 additional whorls. Sculpture

begins on first teleoconch whorl, which is

also pearly white; balance of teleoconch

colored light tan to brown. Teleoconch

sculpture of 7-8 narrow axial ribs per

whorl with sharply angular nodes atshoul-

(Right page) Figure 1. Latirus cloveri spec. nov. Holotype. ANSP 408331, (52.7 mm), from tangle

nets off Aliguay Island, off Dipolog, northwest Mindanao, Philippine Islands, depth 150 m. Figure

2. Latirus cloveri spec. nov. Paratype 1. SC (40.4 mm), from tangle nets, Balicasag Island, Bohol,

Philippine Islands. Figures 3, 4. Fasciolaria walleri Ladd, 1976. SC (51.3 mm), in tangle nets at

night, Cebu, Philippine Islands, depth 130 m. Figure 5. Latirus sarinae spec. nov. Holotype.

ANSP 408332, (50.5 mm), from tangle nets off Mactan Island, Cebu, Philippine Islands, depth

160 m. Figure 6. Latirus sarinae spec. nov. Paratype 1. MNHN (48.9 mm), from type locality.

(Página derecha) Figura 1. Latirus cloveri spec. nov. Holotipo. ANSP 408331, (52,7 mm), de redes de

cerco frente a Aliguay Island, frente a Dipolog, noroeste de Mindanao, Filipinas, profundidad 150 m.

Figura 2. Latirus cloveri spec. nov. Paratipo 1. SC (40,4 mm), de redes de cerco, Balicasag Island,

Bohol, Filipinas. Figuras 3, 4. Fasciolaria walleri Ladd, 1976. SC (51,3 mm), en redes de cerdo de

noche, Cebu, Filipinas, profundidad 130 m. Figura 5. Latirus sarinae spec. nov. Holotipo. ANSP

408332, (50,5 mm), de redes de cerco frente a Mactan Island, Cebu, Filipinas, profundidad 160 m.

Figura 6. Latirus sarinae spec. nov. Paratipo 1. MNHN (48.9 mm), de la localidad tipo.

SNYDER: Four new species of Latirus from Philippine Islands and southern Caribbean

Iberus, 21 (1), 2003

ders crossed by numerous fine spiral cords,

axial ribs streaked white and dark brown,

shoulder nodes usually white. Suture

deeply impressed withimbricate scaly pro-

jections from abapically contiguous whorl.

Aperture ovate, lirate within, interior of

aperture pink to purple-brown with about

25-30 lirations stopping short of smooth

labral margin. Lip moderately thick with

minute dentations terminating in internal

lirations. Columella smooth with one rea-

sonably prominent plica at abapical end

of aperature by canal; small ridge on lip

approximately opposite plica.

Distribution: Latirus cloveri has been

collected in tangle nets at various loca-

tions in the central Philippine Islands.

Discussion: This species was figured

and discussed by SPRINGSTEEN AND LEO-

BRERA (1986: 332, pl. 94, fig. 12) as “Fasci-

olaria” walleri Ladd, 1976 (teratological)

although they mention that “it may rep-

resent a good taxon.” These authors state

that it is “almost morphologically indis-

tinguishable” from Fasciolaria walleri (Figs.

3, 4) which they redescribe from Recent

material (177, pl. 47, fig. 17). However,

apart from the shoulder nodules, which are

more rounded and less prominent in F.

walleri, and the suture, free of the scaly

projections in F. walleri, there are other dif-

ferences worth noting. The color pattern

of F. walleri is a random mix of segments

of brownish axial stripes and vertical

stripes, especially on the body whorl,

against a pale brown background color; the

coloration is much less pronounced than

in L. cloveri. The canal of F. walleri is pro-

portionally longer, straighter and often

more strongly recurved than in L. cloveri.

The status of Fasciolaria walleri deserves

comment. LADD (1976: 133, figs. 16-20)

described this species based upon Pleis-

tocene material from the New Hebrides

Islands, and placed it in the subgenus Pleu-

roploca. Subsequently (LADD, 1982: 47) he

transferred the species to Siphonofusus

[Buccinidae], while noting a superficial

resemblance to the fasciolariid genus Gran-

ulifusus. Later, BEETS (1987: 90) placed this

species in Buccinulum (Euthria) [Buc-

cinidae] and described a subspecies B.

walleri sedanense. These placements were

based on fossil material. This species was

subsequently discovered living in the

Philippine Islands (SPRINGSTEEN AND LEO-

BRERA, 1986: 177, pl. 47, fig. 17). Although

this species has been collected alive, no soft

parts have been preserved and no defin-

itive generic or familial placement can be

made at this time. One subadult speci-

men was obtained with an operculum

which presumably corresponds to the

shell. This operculum, ovate and corneus,

light brown, size and shape correspond-

ing to aperature, with terminal nucleus, is

consistent with assignment as a fascio-

lariid, as is the dentate lip (which buc-

cinids of this size lack) and the lack of

determinate growth.

The final generic placement of Latirus

cloveri is also problematic. At the begin-

ning of the siphonal canal there is a ridge

on the inside of the lip, reminiscent of

Siphonofusus lubrica (Dall, 1918), the type

of Siphonofusus [Buccinidae]. The shell of

L. cloveri is not entirely consistent with

placement in either Latirus or Fusinus al-

though it is apparently a fasciolariid. One

should also note the dentate lip and the

lack of determinate growth. GOFAs (2000:

15) has noted that columellar folds may

not be an absolute differentiator for the

genera Fusinus and Latirus.

Latirus sarinae spec. nov. (Figs. 5, 6)

Type material: Holotype ANSP 408332, length 50.5 mm, in tangle nets, depth 160 m. Paratype 1,

MNHN, length 48.9 mm, from type locality. Paratype 2, SC, length 55.0 mm, from type locality.

No live collected specimens seen, all type material adult.

Other material examined: Four subadult specimens, 34.7 mm and 37.3 mm, from type locality,

and two specimens, 31.3 mm and 44.3 mm, from tangle nets off Balicasag Island, Bohol, Philippine

Islands (all SC).

Etymology: The species is named for the author's daughter, Sarina.

Type locality: Mactan Island, off east central Cebu, Philippine Islands.

SNYDER: Four new species of Latirus from Philippine Islands and southern Caribbean

Description: Shell medium size for

genus, fusiform, length of adult speci-

mens from 48.9 mm to 55.0 mm.

Siphonal canal short, straight, angled

from apertural plane, umbilicus open in

larger specimens but as slit on smaller

specimens. Smooth whitish protoconch

of approximately 1 whorl, eroded on

most specimens; teleoconch of 8-9

whorls with 8 heavy rounded axial ribs

per whorl. Ribs on teleoconch solidly

colored yellow-orange to orange-brown,

interstices between with longitudinal

bands of predominate shell color inter-

spersed with white bands, 6-7 such

bands on body whorl with narrow

white band by suture, siphonal canal

mostly white. Weak spiral cords most

visible crossing axial ribs, with weaker

axial lines. Suture impressed with white

imbricate dentation from abapically

contiguous whorl. Aperture ovate, lip

weakly dentate, thin at edge and thick-

ened within, aperture lirate within after

4 mm smooth inner lip. Columella

smooth with one plica with traces of a

second plica in some specimens.

Distribution: Latirus sarinae has been

collected in tangle nets at various loca-

tions in the central Philippine Islands.

Discussion: Latirus sarinae is morpho-

logically closest to L. kandai Kuroda,

1950 (Fig. 7). This latter species was

described from material collected off

Cape Shirazaki, Kii Peninsula,

Wakayama Prefecture, Japan, but is also

collected in tangle nets in the Philippine

Islands. Latirus kandai is smaller than L.

sarinae and has more prominent spiral

cords that are white against a uniformly

orange-brown background color versus

cords that are the same color as the axial

ribs. The lip of L. kandai is weakly

dentate and the aperture is lirate right

up to the lip, whereas in L. sarinae these

lirae are interrupted by a smooth labral

margin. The protoconchs of both species

are similar in form, indicating that L.

sarinae may be a reasonably widespread

species.

Latirus philippinensis spec. nov. (Figs. 8, 9)

Type material: Holotype ANSP 408333, length 54.8 mm, in tangle nets, depth 160 m. Paratype 1,

MNHN, length 45.2 mm, in tangle nets, depth 160 m, Mactan Island, Cebu, Philippine Islands.

Paratype 2, SC, length 45.3 mm, same data as paratype 1. No live collected specimens seen.

Etymology: The species is named for the area of discovery, the Philippine Islands.

Type locality: Balut Island, off southernmost Mindanao, Philippine Islands.

Description: Shell medium size for

genus, fusiform, thin and elongate,

length of adult specimens from 45 mm

to 55 mm. Siphonal canal long,

recurved. Smooth whitish protoconch,

paucispiral of 1 */4 whorls, teleoconch 9

whorls. Axial sculpture of teleoconch

with prominent rounded ribs, 10 on

body whorl, decreasing in number

toward apex. Six spiral cords on first

five whorls of teleoconch. On sixth

whorl five additional weak cords

between six strong cords, sixteen on

penultimate whorl and numerous cords

on body whorl continuing up full length

of siphonal canal. Axial lines crossing

cords cut surface into small squares.

Suture slightly depressed. Three strong

folds on columella, one more prominent

than others. Aperture narrow and long,

parietal shield glossy, overlaying body

of shell, attached lip thin, weakly

dentate, shiny within. Shell white with

apical portion of early whorls brownish

with additional central brown color

band on body whorl, stripe visible

inside lip on labral margin.

Distribution: Latirus philippinensis is

known from just four specimens, all

from the type locality.

Discussion: Latirus philippinensis cannot

be confused with any other member of the

genus. The very strong columellar folds

resemble those of Latiromitra [Turbinelli-

dae], but the teleoconch sculpture, with

broad uncoronated ribs and numerous

closely-shaped spiral cords, is unlike any

species of Latiromitra. The color pattern of

Iberus, 21 (1), 2003

L. philippinensis is unlike species of

Latiromitra which are without color or

bands. The protoconch is inconclusive for

generic placement, and the other resem-

blances to Latiromitra are superficial.

Although the radula alone would permit

an unequivocal generic assignment, shell

morphology suggests placement in Latirus

rather than Latiromitra.

Latirus philippinensis is similar in size

and shape to Benimakia lanceolata (Reeve,

1847) from the tropical Indo-Pacific but B.

lanceolata has fewer axial ribs, a purple

aperture and a straight canal. It some-

what resembles L. elsiae Kilburn, 1975

from South Africa but that species is not

so elongate and is colored with axial

brown streaks along the ribs rather than .

brown spiral bands. L. philippinensis is

similar in shape to Pseudolatirus discrepans

Kuroda and Habe, 1971 from Japan, and

occasionally collected in tangle nets in

the Philippine Islands, but P. discrepans

grows much longer (to 87.5 mm), has

very faint brown color bands, less promi-

nent axial ribs on later whorls, and a

straight canal. Finally, L. philippinensis

may be compared to Pseudolatirus clausi-

caudatus (Hinds, 1844) from South Africa

but this latter species lacks the prominent

brown color bands, has a very small

aperture with a peculiar thickened lip,

and has a long, straight almost closed

siphonal canal.

Latirus abbotti spec. nov. (Figs. 10, 11)

Type material: Holotype ANSP 408334, length 49.1 mm, on sand, depth 6 m. Paratype 1, USNM

1001651, length 26.0 mm, from type locality. Paratype 2, EC, length 52.1 mm, by scuba at night on

turtle grass, depth 6 m, between Ragged Cay and Sandy Cay off Utila Island, Honduras. Paratype

3, SC, length 45.9 mm, under rubble, depth 3 m, Monte Cristi, Dominican Republic.

Other material examined: 4 immature specimens, 21.0 mm -— 36.0 mm, Utila Island, Honduras; 8

immature specimens, 22.5 mm -— 43.9 mm (largest specimen live collected), Roatan Island,

Honduras; 2 immature specimens, 26.2 mm, 27.1 mm, Cayo Gorda, (Mesquitia) Honduras; 1

immature live collected specimen, 23.6 mm, Punta Rusia, north coast Dominican Republic; 2

immature specimens, 23.4 mm, 25.6 mm, Quita Sueno Bank, Nicaragua (all SC).

Etymology: The species is named in memory of R. Tucker Abbott, one of the foremost researchers

on mollusks of America, and my early mentor.

Type locality: Utila Island, off north central Honduras.

(Right page) Figure 7. Latirus kandai Kuroda, 1950. SC (43.4 mm), in tangle nets, Panglao Island,

off southwest Bohol, Philippine Islands, depth 160 m. Figure 8. Latirus philippinensis spec. nov.

Holotype. ANSP 408333 (54.8 mm), in tangle nets, Balut Island, Cebu, Philippine Islands, depth

160 m. Figure 9. Latirus philippinensis spec. nov. Paratype 2. SC (45.3 mm), from tangle nets,

Mactan Island, Cebu, Philippine Islands, depth 160 m. Figure 10. Latirus abbotti spec. nov. Holo-

type. ANSP 408334, (49.1 mm), on sand off Utila Island, Honduras, depth 6 m. Figure 11.

Latirus abbotti spec. nov. Paratype 2. EC (52.1 mm), by scuba at night on turtle grass between

Ragged Cay and Sandy Cay off Utila Island. Honduras, depth 6 m. Figure 12. Latirus angulatus

(Róding, 1798). SC (54.7 mm), dived from sand, under rocks, Salvador, Bahia State, Brazil, depth

15-20 m.

(Página derecha) Figura 7. Latirus kandai Kuroda, 1950. SC (43,4 mm), en redes de cerco, Panglao

Island, al sudoeste de Bohol, Filipinas, profundidad 160 m. Figura 8. Latirus philippinensis spec. nov.

Holotipo. ANSP 408333 (54,8 mm), en redes de cerco, Balut Island, Cebu, Filipinas, profundidad 160

m. Figura 9. Latirus philippinensis spec. nov. Paratipo 2. SC (45,3 mm), en redes de cerco, Mactan

Istand, Cebu, Filipinas, profundidad 160 m. Figura 10. Latirus abbotti spec. nov. Holotipo. ANSP

408334, (49,1 mm), en arena frente a Utila Island, Honduras, profundidad 6 m. Figura 11. Latirus

abbotti spec. nov. Paratipo 2. EC (52,1 mm), de buceo nocturno sobre Thalassia testudinum entre

Ragged Cay y Sandy Cay frente a Utila Island. Honduras, profundidad 6 m. Figura 12. Latirus angula-

tus (Róding, 1798). SC (54,7 mm), arena, bajo rocas, Salvador, Bahia, Brasil, profundidad 15-20 m.

SNYDER: Four new species of Latirus from Philippine Islands and southern Caribbean

Iberus, 21 (1), 2003

Description: Shell medium size for

genus, fusiform, length of adult speci-

mens from 45.9 mm to 52.1 mm.

Siphonal canal short, straight, angled at

approximately 22” from axis of aperture,

colored brown to blackish-brown.

Smooth, cream-colored, bulbous proto-

conch of about 1 whorl and teleoconch of

8-9 whorls. Sculpture of teleoconch with

10-11 prominent rounded white to

cream-colored axial ribs with dark

brown interstices, converging above ribs

on body whorl to color canal dark

brown. Numerous reddish-brown spiral

cords crossing axial ribs; some speci-

mens with a single white band circling

from anal canal around body whorl to

approximate center of lip; band may

follow suture between penultimate and

body whorl and then becomes evanes-

cent. At suture, ribs terminate with a

brown band above. Columella with two

prominent plicae and an additional

smaller plica at beginning of canal. Aper-

ACKNOWLEDGEMENTS

Gary Rosenberg and Geerat Vermeij

offered helpful suggestions regarding

these new species. Philippe Bouchet

BIBLIOGRAPHY

BEETS, C., 1987. Notes on Buccinulum (Gas-

tropoda, Buccinidae), a reappraisal. Scripta

Geologica, 82: 83-100, pls. 6, 7, (1986). [Dated

in internal heading as January, 1987].

BOzZETTI, L., 1997. Study of the collection of Mr.

Emmanuel Guillot de Suduiraut with the

descriptions of three new gastropod species

(Fasciolariidae, Trochidae and Turridae). Bul-

letin of the Institute of Malacology, Tokyo 3 (4):

BULLOCK, R. C., 1974. A contribution to the sys-

tematics of some West Indian Latirus (Gas-

tropoda: Fasciolariidae). The Nautilus, 88 (3):

69-79.

CERNOHORSKY, W. O., 1987. The taxonomy of

some Indo-Pacific Mollusca. Part 14, with

descriptions of two new species. Records of the

Auckland Institute and Museum, 24: 107-122,

figs. 1-33.

ture narrowly elongate, lirate with thick

weakly dentate lip; sometimes bright

yellow within when freshly collected.

Umbilicus open only on large specimens.

Distribution: Latirus abbotti has been

widely collected in the southwestern

Caribbean.

Discussion: This distinctive species is

easily differentiated from known

Caribbean species of Latirus. The

unusual coloration and numerous axial

ribs are an immediate distinguishing

characteristic. The closest species mor-

phologically is Latirus angulatus

(Róding, 1798) which has an elongate

form (Fig. 12) from Cuba and South

America (see as well BULLOCK (1974: 74-

76, fig. 20)). Latirus angulatus has just 7-8

axial ribs on each whorl compared to 10-

11 in L. abbotti and has an orange to

orange-brown background with lighter

longitudinal cords; the color is uniform

throughout whereas in L. abbotti is

brown with lighter often whitish ribs.

offered a helpful discussion concerning

Latiromitra. Sarah Watson took the digital

photographs and composed the plates.

DaLtL, W. H., 1918. Notes on Chrysodomus and

other mollusks from the north Pacific Ocean.

Proceedings of the United States National

Museum, 54 (2234): 207-234.

GARCÍA, E. F., 2001. A new species of Latirus

(Gastropoda: Fasciolariidae) from the south-

ern Philippines. Novapex, 2 (4): 149-151, figs.

1-4, 1 text fig.

GOFEAS, S., 2000. Four species of the family Fas-

ciolariidae (Gastropoda) from the north

Atlantic seamounts. Journal of Conchology, 37

(1): 7-16, figs 123;

KILBURN, R. N., 1975. Taxonomic notes on South

African marine Mollusca (5), including

descriptions of new taxa of Rissoidae, Cerithi-

idae, Tonnidae, Cassidae, Buccinidae, Fasci-

olariidae, Turbinellidae, Turridae, Architec-

tonicidae, Epitoniidae, Limidae and Thraci-

idae. Annals of the Natal Museum, 22 (2): 577-

622, figs. 1-25.

SNYDER: Four new species of Latirus from Philippine Islands and southern Caribbean

KURODA, T., 1950. Descriptions of two new

species of marine gastropods, dedicated to

Mr. K. Kanda on his 77th birthday. Venus, 16

(1-4): 49-52.

KURODA, T. AND HABE, T., 1971. [Descriptions

of genera and species] In: T. Kuroda, T. Habe

and K. Oyama. The Sea Shells of Sagami Bay.

Tokyo: Maruzen. xix, 741 pp. [Japanese text),

121 pls., 489 pp. [English text], 51 pp., index,

map.

LapD, H. S., 1976. New Pleistocene Neogas-

tropoda from the New Hebrides. The Nautilus

, 90 (4): 127-138.

LADbD, H. S., 1982. Cenozoic fossil mollusks

from west Pacific islands; Gastropods (Eulim-

idae and Volutidae through Terebridae).

United States Geological Survey Professional

Paper, 1171: i-iv, 1-100, pls. 1-41.

PETUCH, E. J., 1981. A relict neogene caenogas-

tropod fauna from northern South America.

Malacologíia , 20 (2): 307-347.

PETUCH, E. J., 1990. A new molluscan faunule

from the Caribbean coast of Panama. The

Nautilus, 104 (2): 57-71.

SNYDER, M. A., 1988. Latirus martini (Gas-

tropoda: Fasciolariidae), a new species from

Honduras. The Nautilus, 102 (2): 54-55.

SPRINGSTEEN, F. J. AND LEOBRERA, F. M., 1986.

Shells of the Philippines. Manila: Carfel Seashell

Museum. 377 pp., 100 pls.

asin amados bre bnst Al SARA lote do ES

DE E

O Sociedad Española de Malacología ——_———— Iberus, 21 (1); 11-33, 2003

The littoral molluscs (Gastropoda, Bivalvia and Poly-

placophora) of Sáo Vicente, Capelas (Sao Miguel Island,

Azores): ecology and biological associations to algae

Los moluscos litorales (Gastropoda, Bivalvia y Polyplacophora) de

Sáo Vicente, Capelas (Sáio Miguel Island, Azores): ecología y asocia-

ciones biológicas con algas

Sérgio P. ÁVILA*

Recibido el 12-IV-2002. Aceptado el 26-VIIH-2002

ABSTRACT

A two-year systematic survey was conducted in a rocky exposed shore located at Porto

das Baleias (Sáo Vicente, Capelas) in the north coast of Sáo Miguel Island and suppos-

edly representative of the Azorean rocky shores. Zonation of the littoral molluscs [(Gas-

tropoda, Bivalvia and Polyplacophora) was established for the most abundant species and

possible molluscs/algae biological associations were studied.

Seventy-one taxa (56 Gastropoda, 13 Bivalvia and 2 Polyplacophora) were found in the

1996 and 1997 fieldwork, with a total of 35,960 specimens recorded. Bittium sp., an

Azorean endemism, was the commonest taxon, with 11,236 and 12,374 individuals

(61.1% and 75.4%, respectively in 1996 and 1997). Thirteen taxa accounted for about

96% of the total of specimens collected in both years. Some differences in the relative

abundance of the commonest taxa were found between the two years. Besides Bittium sp.,

the next most abundant taxa in 1996 were Tricolia pullus azorica (14,0%), Jujubinus

pseudogravinae and Manzonia unifasciata, both with 4%, and Parvicardium vroomi, Alva-

nia sleursi and Crassadoma pusio, all between 2-3%. In 1997, P. vroomi and T. pullus

azorica (6,4% and 6,1%) were the most abundant species after Bittium sp., followed by

Alvania sleursi and Jujubinus pseudogravinae with a little more than 1% of the total num-

ber of molluscs collected in that year.

The endemic rissoids Alvania angioyi, Manzonia unifasciata and Rissoa guernei, the also

endemic trochid Gibbula delgadensis, the Macaronesian Anachis avaroides and the small

bivalve Parvicardium vroomi were only abundant in shallow levels (down to 5-óm]),

whereas the endemic rissoid Alvania sleursi is especially abundant below 20m depth. Bit-

tium sp., Tricolia pullus azorica and Jujubinus pseudogravinae were found along all tran-

sect. No specific molluscs/algae associations were found.

RESUMEN

Se investigó durante dos años un tramo de costa rocosa expuesta en Porto das Baleias

(Sáo Vicente, Capelas) en la costa norte de Sáo Miguel que se suponía respresentativa de

las costas rocosas azoreanas. Se estableció la zonación de los moluscos litorales [(Gastro-

* Seccáo de Biologia Marinha and CIRN, Departamento de Biologia, Universidade dos Acores, 9501-801 Ponta

Delgada — Azores. e-mail: avilaGnotes.uac. pt

Iberus, 21 (1), 2003

poda, Bivalvia y Polyplacophora) más abundantes y se estudiaron posibles asociaciones

moluscos/algas.

Se encontraron 71 táxones (56 Gastropoda, 13 Bivalvia y 2 Polyplacophora) durante los

años 1996 y 1997. Bittium sp., un endemismo azoreano, fue la especie más abundante

(61,1 % y 75,4 % del total en el 96 y 97 respectivamente). Trece especies representaron

el 96% del total de especimenes recolectados en ambos años. Tras Bittium sp, los táxones

más abundantes en el 96 fueron Tricolia pullus azorica (14,0%), Jujubinus pseudogravi-

nae y Manzonia unifasciata, ambos con el 4%, y Parvicardium vroomi, Alvania sleursi y

Crassadoma pusio, entre el 2 y 3%. En el 97, tras Bittium sp. aparecen P. vroomi y T.

pullus azorica (6,4% y 6,1%), seguidos de Alvania sleursi y Jujubinus pseudogravinae con

algo más del 1%.

Los risoideos endémicos Alvania angioyi, Manzonia unifasciata y Rissoa guernei, el tam-

bién endémico trocoideoGibbula delgadensis, Anachis avaroides y el pequeño bivalvo

Parvicardium vroomi solo son abundantes en los niveles someros (hasta 5-6m), mientras

que Alvania sleursi es especialmente abundantes por debajo de 20 m. Bittium sp., Tricolia

pullus azorica y Jujubinus pseudogravinae se encontraron a lo largo de todo el transecto

No se encontró ninguna asociación molusco/alga

KEY WORDS: Mollusca, Azores, littoral, ecology, biological associations.

PALABRAS CLAVE: Mollusca, Azores, litoral, ecología, asociaciones biológicas.

INTRODUCTION

The first authors who studied the

marine molluscs of the Azores - Mac

ANDREW (1856), DRoUET (1858),

MORELET (1860), DAUTZENBERG AND

FISCHER (1896) and NOBRE (1924; 1930) —

were mainly concerned with the publi-

cation of commented checklists. The

most complete work with such a

methodology is still the classic first

volume of the Oceanographic Cam-

paigns of Prince of Monaco by

DAUTZENBERG (1889), where a large

number of new species were described.

Since the foundation of the Univer-

sity of the Azores, several scientific

expeditions were made by the Marine

Biology team of the Department of

Biology of this University to some of the

islands (“Graciosa/88”, “Flores/89”,

“Acores/89”, “Santa Maria and Formi-

gas/90”, “Pico/91” and “Faial/93”) pro-

viding new insights to the Azorean lit-

toral knowledge in general and an

improvement in the marine molluscs in

particular (AZEVEDO AND MARTINS,

1989; GOFaAs, 1989; AZEVEDO, 1990;

GOFAS, 1990; AZEVEDO AND GOFAS, 1990;

NETO AND AZEVEDO, 1990; AZEVEDO,

19918; ÁviLa, 1996; ÁVILA AND

AZEVEDO, 1996; ÁVILA, 1997; ÁVILA AND

AZEVEDO, 1997; ÁviLa, 1998; ÁVILA,

AZEVEDO, GONCALVES, FONTES AND

CARDIGOS, 1998; ÁVILA, AZEVEDO,

GONGALVES, FONTES AND CARDIGOS,

2000a; ÁVILA, GONCALVES, FONTES AND

CARDIGOS, 2000b; ÁVILA AND ALBER-

GARIA, 2002).

From 1996 on, a database with the

littoral molluscs of the Azores (<50m)

was constructed, based on samples col-

lected in all the islands of the archipel-

ago and an exhaustive and critical

examination of literature was done. This

data has provided the possibility to

further understand the biogeographical

relationships of the Azorean shallow-

water marine molluscs (ÁVILA, 2000a).

Molluscs/algae biological associa-

tions were first studied in the Azores by

CHAPMAN (1955), who was surprised by

the great abundance of molluscs in a

sample of Corallina sp. from Faial Island.

Later, BULLOCK, TURNER AND FRALICK

(1990) studied the micromolluscs associ-

ated to several monospecific algal

fronds in the rocky intertidal. Rissoa

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

guernei Dautzenberg, 1889, Tricolia pullus

azorica Dautzenberg, 1889, Rissoella sp.

and Omalogyra atomus (Philippi, 1841)

were found by these authors in high

numbers associated to Pterocladiella

capillacea, whereas Ulva rigida samples

had two very common small gastropods

in the intertidal, Skeneopsis planorbis

(Fabricius O., 1780) and Omalogyra

atomus, as well as Tricolia pullus azorica;

on Stypocaulon scoparia the best repre-

sented species were Skeneopsis planorbis,

Rissoa guernei and Omalogyra atomus.

AZEVEDO (1991a) studied the mala-

cological communities associated to the

algal fronds in the littoral of Sáo Miguel.

This author found Pisinna glabatra (Von

Munlfeldt, 1824) and Omalogyra atomus

to be the most abundant species in the

intertidal, whereas Omalogyra atomus

and, occasionally, Bittium sp. were the

most abundant taxa in the infralittoral.

At Lajes do Pico (Pico Island), in a

very shallow and sheltered coastal

lagoon, ÁVILA (1998) found Cingula tri-

fasciata (Adams J., 1798) to be the most

abundant species in the intertidal, with

maximum densities of 32,500 ind./m*.

According to BULLOCK (1995), in the

intertidal of the Ilhéu de Vila Franca, an

islet located in the south shores of Sáo

Miguel, Alvania mediolittoralis Gofas,

1989 and Crisilla postrema (Gofas, 1990),

two rissoids, can reach densities higher

than 10,000 ind. /m?.

More recently, COSTA AND ÁVILA

(2001) have studied the molluscs associ-

ated to monospecific samples of

Halopteris filicina and Stypocaulon sco-

paria at Sáo Miguel Island. The samples

were collected between 11 and 15m

depth and these authors found that 4

taxa (Bittium sp., present in all of the

samples and responsible for 85.6% of

the total number of molluscs collected,

Setiía subvaricosa Gofas, 1989, Tricolia

pullus azorica and Rissoa guernei) made

up 96.6% of the total number of mol-

luscs.

Algae provide shelter to the mol-

luscs, protecting them against heavy

waves and currents (DEAN AND

CONNEL, 1987), they work as a shelter

against predators (SEED, 1986; BULLOCK

ET AL., 1990) and give them food,

directly through the tissues of the algae

or of epiphytes diatoms or microalgae,

and indirectly, through sediments and

detritus that accumulate especially in

the base of the algal fronds (AZEVvEDO,

199%la; BuLLOCK, 1995). The structural

habitat complexity provided by

macroalgae, usually related to higher

specific richness (HICKs, 1986) is depen-

dent upon abiotic factors such as

hydrodynamic prevalent conditions,

intensity and type of light and the exis-

tence of pollutants; and of biotic factors,

such as predators occurrence, light

competition and algal morphology,

which in turn is related with the type of

ramification and growth of the alga.

BULLOCK ET AL. (1990) stated that algae

with ramified growth, foliose or tuft-

like non-coralline alga, possess a high

number of micromolluscs, in contrast

with species with blade-like branches

(Fucus spiralis), filamentous algae or

tuft-like coralline alga.

The main objectives of this work are

to establish the zonation of the most

abundant molluscs in Sáo Miguel Island

and to verify possible molluscs /algae

biological associations.

MATERIALS AND METHODS

Subtidal communities off Porto das

Baleias (Sáo Vicente, Capelas) located in

the north shore of Sáo Miguel Island

(Fig. 1) were surveyed under the project

PRAXIS/2/2.1/BIA/169/94, “Biodiver-

sity of the archipelago of the Azores”.

All samples were collected in July 1996

and July 1997 (see Table ID). A transect

was performed from low-tide level to

30m depth, with an extension of 450m

long (Fig. 2) across a rocky bottom

extensively covered by algae. Ptero-

cladiella capillacea, Ulva spp., Stypocaulon

scoparia, Asparagopsis armata, and Hypnea

musciformis dominated the shallow

levels, whereas Plocamium cartilagineum

and Zonaria tournefortil were more abun-

dant between 10 and 30m depth.

Asparagopsis armata, Halopteris filicina

and Dictyota dichotoma were present

Iberus, 21 (1), 2003

Table I. Samplings collected at Porto das Baleias, Sio Vicente, Capelas (Sáo Miguel Island).

DBUA: Reference collection of the Department of Biology of the University of the Azores.

Tabla I. Muestras recogidas en Porto das Baleias, Sáo Vicente, Capelas (Isla Sáo Miguel). DBUA: Refe-

rencia en el Departamento de Biología de la Universidad de Azores.

1996 1997

Depth (m) Date DBUA Depth (m) Date DBUA

4 19-7-96 107/6 5 16-7-97 166

4 19-7-96 107/1 5 16-7-97 785

4 19-7-96 107/0 5 16-7-97 188

5 19-7-96 708/H 6 10/7/97 184

5 19-7-96 108/1 6 10/7/97 181

5 19-7-96 708/F 6 10/7/97 182

8 18-7-96 708/€ 9 15-7-97 162

8 18-7-96 103/€ 9 15-7-97 eb

y 18-7-96 703/8 9 15-7-97 764

12 18-7-96 7103/H 10 15-7-97 177

12 18-7-96 704/A 10 15-7-97 169+791

12 18-7-96 704/D 10 15-7-97 'YA)

14 17-7-96 701/8 12 14-7-97 168

14 17-7-96 701/A 12 14-7-97 178

14 17-7-96 701/0 12 14-7-97 176

16 17-7-96 702/D 15 HN 183

16 17-7-96 702/€ 15 11/7/97 179

16 17-7-96 702/H 15 11/7/97 187

22 16-7-96 694/0 20 11/7/97 780+790

22 16-7-96 692/8 20 11/7/97 161

22 16-7-96 693/0 20 11/7/97 186

27 15-7-96 690/B 25 12/7/97 163+773

2 15-7-96 691/0 25 12/1/91 174

27 15-7-96 689/4 25 12/1/97 189

30 14-7-97 170

30 14-7-97 115

30 14-7-97 165

Porto das Baleias, Sáo Vicente,

Capelas

25” 30" W

37250” N

Figure 1. Sáo Miguel Island (Azores), with the location of Porto das Baleias, Sáo Vicente, Capelas.

Figura 1. Isla Sáo Miguel (Azores), con la localización de Porto das Baleias, Sáo Vicente, Capelas.

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

200 400

0 100

300

25 E

30 ES

Figure 2. Profile of the transects, Porto das Baleias,

200 400

300

Sáo Vicente, Capelas (Sáo Miguel, Azores) in

1996 and 1997. The arrows show the sampling sites of the 3 replicates.

Figura 2. Perfil de los transectos, Porto das Baleias, Sáo Vicente, Capelas (Sáo Miguel, Azores) en 1996

y 1997. Las flechas muestran los lugares de muestra de las 3 réplicas.

along all transect. In most of the tran-

sect, large boulders were dominant, but

especially bellow 20m depth, there were

large areas of bedrock interspersed with

sandy patches.

Minimum area for quantitative

studies of algae in the Azorean sublit-

toral is 50x50cm (NETO, 1997) larger

than the minimum area determined for

molluscs (25x25cm) (AZEVEDO, 1991a) so

the former area was used. Three

quadrates were scrapped in each

selected depth; the algae collected were

put in a labeled cotton drawstring bag.

In the laboratory, samples were washed

several times with seawater and animals

were removed by pouring the washing

water through 1 mm, 0.5 mm and 0.25

mm mesh sieves. Samples were labeled

and preserved in 70% ethanol. After

draining for about 30min, the wet

weight of the algae was determined

Table II. Most common taxa/species found at Porto das Baleias, Sáo Vicente, Capelas during the

1996 and 1997 campaigns. n: total number of specimens collected.

Tabla II. Especies más comunes en Porto das Baleias, Sáo Vicente, Capelas durante las campañas de

1996 y 1997. n: número total de especímenes recolectados.

1996 1997

n n

Bittium sp. 11,936 12,374

Tricolia pullus azorica IE 993

Parvicardium vroomi 557 1042

Jujubinus pseudogravinae 153 216

Manzonia unifasciata 7119 78

Alvania sleursi 526 284

Crassadoma pusio 447 36

Cardita calyculata 320 117

Alvania angioyi 205 159

Ocinebrina aciculata 145 162

Anachis avaroides 168 108

Gibbulo delgadensis 62 160

Rissoa guernei 195 35

Total 18,725 15,764

TOTAL (all taxa) 19,540 16,420

1996 1997

% %

61.1 15.4

14.0 6.1

1) 6.4

3.7 3

3./ 0.5

7, 1.7

LS 0.2

1.6 0.7

1.1 1.0

0.7 1.0

0.9 0.7

0.3 1.0

0.8 0.2

96.0 96.2

100.0 100.0

Iberus, 21 (1), 2003

Bittium sp.

1996

E Parvicardium vroomi

1996

200

100

3 5 8 12 14 16 DO 27

> Manzonia unifasciata

199

150

100

3 5 8 12 14 16 22 7

n ; :

> Alvania sleursi

1996

100

3 5 8 12 14 T6 22 27

3000 A

2500 7

o A A

Tricolia pullus azorica

9. 10.12 ,15* 201 234230

2 Parvicardium vroomi

1997

200

100

5 6 9 101215 20 "2530

Manzonia unifasciata

200

1997

150

100

5 6 9 100 12 15 201903750

a Alvanta sleursi

1997

100

Figure 3. Zonation of the most common molluscs species in 1996 and 1997 at Porto das Baleias,

Sáo Vicente, Capelas (Sáo Miguel, Azores) (maximum, mean and lower values by depth).

Figura 3. Zonación de las especies de moluscos más comunes en 1996 y 1997 en Porto das Baleias, Sáo

Vicente, Capelas (Sáo Miguel, Azores) (valores máximo, medio y mínimo por profundidaa).

(+0,01g). All molluscs found in the 1

mm mesh sieve were sorted, identified

and counted (see Appendices 1 and 2).

Only live molluscs were counted. Mol-

luscs retained in the 0.5 mm and 0.25

mm mesh sieves will be studied in

future works. About 300 days were

spent just sorting the material. All

AVILa: Littoral molluscs of Azores: ecology and biological associations to algae

Gibbula delgadensis

1996

Anachis avaroides

5) ,) 8 TAO 22 027

Jujubinus pseudogravinae

3 5 8 12 IGG: 2239127

Rissoa guernei

1996

Figure 3. Continuation.

Figura 3. Continuación.

samples were given a number and were

deposited in the DBUA (Department of

Biology of the University of the Azores)

marine molluscs reference collection.

Most problematic molluscan taxa

were photographed under the scanning

n e

80 Alvania angioyi

5 6 9 OPA TISAIO. +25 30

2 Gibbula delgadensis

1997

0 .

5 6 9 TOALLAS

a Anachis avaroides

5 6 9 VOD LOL ESO

p Jujubinus pseudogravinae

eS 1997

120

100

5 EAS TIA MIA SERLO 025 4130

Ñ 50 Rissoa guernel

1057

SAA AAA USA AA E

electronic microscope (protoconch,

teleoconch, and microsculpture of both);

these photos were quite useful in the

posterior identification. Algae were

identified by Dra. Ana Neto (University

of the Azores).

Iberus, 21 (1), 2003

5 6 9" 10 .12 15 20 271680

Figure 4. Taxa/species number (S), diversity index of Shannon-Wiener (H”) and equitatibility

index (J) by depth in 1996 and 1997 (maximum, mean and lower values by depth).

Figura 4. Número de especies (S), índice de diversidad de Shannon-Wiener (H) e índice de equitatibi-

lidad (J) por profundidades en 1996 y 1997 (valores máximo, medio y mínimo por profundidad).

STATISTICAL ANALYSIS

Data were analyzed with the statisti-

cal package PRIMER (version 5.2) (Ply-

mouth Routines in Multivariate Ecologi-

cal Research - Plymouth Marine Labora-

tory). For each depth sampled in both

campaigns (1996 and 1997), zonation

graphs were constructed for the most

abundant taxa with maximum, mean

and lower density values.

Species diversity was calculated in

each quadrate, using species richness (S)

and diversity indices of Shannon-Wiener

(H') and evenness (J) (KREBS, 1989;

BEGON, HARPER AND TOWNSEND, 1996).

Multivariate analysis was per-

formed, with raw data transformed by

VVGo (FIELD, CLARKE AND WARWICK,

1982; CLARKE AND AINSWORTH, 1993). In

O-analysis, dendrograms were con-

structed with Bray-Curtis similarity

index followed by UPGMA method

(FIELD ET AL., 1982); R-analysis was also

performed, dendrograms constructed

by using Euclidean distance and

UPGMA method.

ANOSIM/PRIMER program (“ran-

domization /permutation test”) (WAr-

WICK AND CLARKE, 1993) was used with

20,000 permutations, to test differences

between pre-selected groups of samples.

This program results in a value of R,

between —-1 and +1 (usually, R> 0). If R=

1, this means that all replicates of a given

depth are more similar than any repli-

cates of different depths; if R= 0, the sim-

ilarity between replicates is independent

of its depth and will be similar, in

average (CLARKE AND WARWICK, 1994).

Species with a ratio>1.40 (SIMPER/

PRIMER analysis) were considered as

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

Table III. Diversity indices by quadrate in 1996 and 1997. S: number of taxa/species; H': diversity

index of Shannon-Wiener; J: index of equitatibility.

Tabla 111. Índices de diversidad por cuadrado en 1996 y 1997. S: número de especies; H': índice de

diversidad de Shannon-Wiener; J: índice de equitabilidad.

1996

Quadrate— Depth S H J

1 3 28 1.42 0.426

2 3 21 1.35 0.443

3 3 30 1.70 0.500

4 5 23 1.58 0.504

5 5 23 1.45 0.462

6 5 18 1.42 0.491

/ 8 20 1.67 0.556

8 38 3 1.81 0.577

[0 ..)

— dl

—

(9 2]

[o]

(9 2]

[a]

mainly responsible for the differences

detected between pre-selected groups

(WARWICK, PLATT, CLARKE, AGARD AND

GOBIN, 1990). Three groups were selected,

based on the empirical knowledge of the

transect's bottom characteristics: shallow

levels (until 10m depth), medium depths

(between 10 and 20m depth) and deeper

levels (from 20 down to 30m depth). These

pre-selected groups were designed by the

letters s (“shallow”), m (“medium”) and

d (“deep”), respectively.

DAFOR scale, [D-dominant (>75%);

A-abundant (50%); F-frequent (25%); O-

occasional (10%); R-rare (5%)] was used

1997

Quadrate Depth S H' j

| 5 16 0.90 0.326

2 5 27 0.82 0.248

2 5 26 0.81 0.248

4 6 32 1.69 0.486

5 6 12 0.56 0.226

6 6 8 1.29 0.622

7 9 9 0.69 0.314

8 9 11 0.50 0.207

9 9 13 0.4] 0.159

10 10 15 0.75 0.276

11 10 5 0.39 0.245

12 10 9 1.04 0.475

13 12 14 2.21 0.837

14 12 6 0.67 0.373

15 12 13 177 0.691

16 15 / 0.67 0.345

17 15 i/ 0.93 0.476

25 30 / 0.83 0.429

26 30 5 1.16 0.721

21 30 6 0.41 0.231

to measure the relative abundance of the

algae species. As this is a semi-quantita-

tive scale, in contrast to the quantitative

molluscs data, the latter were transformed

according]ly, in order to test possible mol-

luscs /algae biological associations. A mul-

tivariate analysis was conducted over the

semi-quantitative molluscs-algae data

(most abundant species in both 1996 and

1997) and dendrograms were constructed

using Bray-Curtis similarity index fol-

lowed by UPGMA. Euclidean distance

was used to construct the dendrograms

of the analysis of pooled semi-quantita-

tive molluscs-algae data.

Iberus, 21 (1), 2003

O —KÁ — ==

40 50 60

70 80 90 100

Figure 5. All replicates of 1996 and 1997 grouped by depth-year. Data transformed by VWx, Bray-

Curtis similarity index, UPGMA.

Figura 5. Total de réplicas de 1996 y 1997 agrupadas por profundidad-año. Datos transformados por

WVx, índice de similaridad de Bray-Curtis, UPGMA.

RESULTS

A total of 35,960 specimens were

counted in both 1996 and 1997 cam-

paigns at Sáo Vicente, Capelas, belong-

ing to 71 taxa (56 Gastropoda, 13

Bivalvia and 2 Polyplacophora). Of

those, 19,540 specimens were collected in

1996 and 16,420 in 1997. Forty-one taxa

were found in both years (see Appen-

dices 1 and 2), 11 and 17 taxa exclusively

in 1996 and 1997, respectively.

Bittium sp. was the commonest taxa,

with 11,936 and 12,374 individuals

(61.1% and 75.4%, respectively in 1996

and 1997). Thirteen taxa accounted for

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

22-96

27-96

8-96

14-96

16-96

5-96

16-96

4-96

5-96

12-96

14-96

16-96

22-96

12-96

14-96

A 12-96

22-96

27-96

50 60 70 80 90 100

20-97

5-97

10-97

20-97

5-97

6-97

15-97

20-97

25-97

9-97

12-97

6-97

9-97

6-97

10-97

15-97

12-97

25-97

12-97

B 30-97

30-97

10-97

30-97

30 40 50 60 70 80 90 100

Figure 6. A: replicates of 1996; B: replicates of 1997 (grouped by depth-year). Data transfofmed

by VWx, Bray-Curtis similarity index, UPGMA.

Figura 6. A: réplicas de 1996; B: réplicas de 1997 (agrupadas por profundidad-año). Datos transfor-

mados por NNx, índice de similaridad de Bray-Curtis, UPGMA.

Iberus, 21 (1), 2003

1996 Zonaria tournefortii

Acrosorium spp.

Sphaerococcus coronopifolius

Plocamium cartilagineum

Dictyota dichotoma

Bittium sp.

Taonia tomaria

Stypocaulon scoparia

Ulva spp.

Jujubinus pseudogravinae

Alvania sleursii

Crassadoma pusio

Cardita calyculata

Halopteris filicina

Parvicardium vroomi

Manzonia unifasciata

Alvania angioyi

Asparagopsis armata

Tricolia pullus azorica

Hypnea musciformis

Pterocladiella capillacea

10 8 6 4 2 0

1997

Dictyota dichotoma

Asparagopsis armata

Halopteris filicina

Zonaria tournefortii

Acrosorium verulosum

Jujubinus pseudogravinae

Alvanta sleursii

Ocinebrina aciculata

Parvicardium vroomi

Tricolia pullus azorica

Hypnea musciformis

Pterocladiella capillacea

Stypocaulon scoparia

Ulva spp.

Bittium sp.

-——á> >A<<<«««<<<«IDIRO€E€FRIT-qC€O€P€>€-

15 10 5 0

Figure 7. Mollusc/algae collected in 1996 and 1997. Non-transformed semi-quantitative data,

DAFOR scale. Euclidean distance, UPGMA.

Figura 7. Moluscoslalgas recolectados en 1996 y 1997. Datos semicuantitativos no transformados, escala

DAFOR. Distancias euclidianas, UPGMA.

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

16 m+d

50 60 70 80 90 100

15 m+d

40 60 80 100

Figure 8. Molluscs/algae collected in 1996 and 1997. Non-transformed semi-quantitative data,

DAFOR scale. Bray-Curtis similarity index, UPGMA.

Figura 8. Moluscos/algas recolectados en 1996 y 1997. Datos semicuantitativos no tarnsformados, escala

DAFOR. Índice de similaridad de Bray-Curtis, UPGMA.

about 96% of the total of specimens col- found between the two years. Besides

lected in both campaigns (see Table ID. Bittium sp., most abundant taxa in 1996

Some differences in the relative abun- were Tricolia pullus azorica (14,0%), Juju-

dance of the commonest taxa were binus pseudogravinae Nordsieck, 1973

Iberus, 21 (1), 2003

Table IV. Pre-defined groups and symbols used in the multivariate analysis with ANOSIM and

SIMPER programs.

Tabla IV. Grupos predefinidos y símbolos usados en el análisis multivariantes con los programas

ANOSIM y SIMPER.

Depth (m) - year

4-96; 5-96; 6-96

591; 6-91; 9-97; 10-97

12-96; 14-96; 16-96

12-97; 15-97; 20-97

22-96; 21-96

2597; 30-97

Pre-defined groups

596 — shallow water, 1996

$97 — shallow water, 1997

m96 — medium water, 1996

m97 — medium water, 1997

d96 — deep water, 1996

d97 — deep water, 1997

Table V. ANOSIM results. Data transformed by VVx. Other symbols as in Table IV. |

Tabla V. Resultados de ANOSIM. Datos transformados por Vx. Resto de símbolos como en la Tabla IV

Comparisons between groups

596 / 597

m96 / m97

d96 / 497

597 / m97

m97 / 497

d96 / m97

(s96:597) / (m96-m97)

(596597) / (496-497)

(m96-m97) / (096-497)

(s96-m96-496) / (s97-m97-497)

and Manzonia unifasciata (Dautzenberg,

1889), both with 4%, and Parvicardium

vroomi van Aartsen, Menkhorst and Git-

tenberger, 1984, Alvania sleursi (Amati,

1987) and Crassadoma pusio (Linnaeus,

1758), all between 2-3%. In 1997, P.

vroomi and T. pullus azorica (6,4% and

6,1%) were the most abundant species

after Bittium sp., followed by Alvania

sleursi and Jujubinus pseudogravinae with

a little more than 1% of the total number

of molluscs collected in that year.

Bittium sp. was found in all quadrates

sampled in 1996 and 1997, whereas A.

sleursi, C. pusio and T. pullus azorica were

present in all quadrates in 1996 (Appen-

dices 1 and 2).

Some species were clearly only

abundant in shallow levels (down to 5-

6m) as the endemic rissoids Alvania

0.277

0.430

0.431

0.039

-0.031

0.137

0.056

0.198

0.046

0.337

angioyi van Aartsen, 1982, Manzonia uni-

fasciata and Rissoa guernei, the also

endemic trochid Gibbula delgadensis

Nordsieck, 1982, the Macaronesian

Anachis avaroides Nordsieck, 1975 and

the small bivalve Parvicardium vroomi.

On the other side, the endemic rissoid

Alvania sleursi, is especially abundant

below 20m depth, whereas Bittium sp.,

Tricolia pullus azorica and Jujubinus

pseudogravinae were found along all

transect. The highest variation detected

when both years are compared, is

related to the higher abundance of

Bittium sp. at 5m in 1997 (Fig. 3).

Species number and diversity in-

dices of Shannon-Wiener (H') and even-

ness (J), show lower variations between

replicates by depth in 1996 when com-

pared to 1997 (cf. Table Ill and Fig. 4).

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

When all 1996 and 1997 replicates

are pooled, the dendrogram (Q-analy-

sis) shows a clear split of replicates by

year (Fig. 5). Besides, and with only a

few exceptions, most of the clusters are

among replicates of similar depths. A

similar analysis by year shows that in

1996, replicates follow the depth gradi-

ent (Fig. 6A); in contrast, 1997 replicates

do not show such an evident pattern,

with higher mixture of replicates of dif-

ferent depths / levels (cf. Fig. 6B).

Cluster analysis of pooled algae-

molluscs data (Euclidean distance) is

not particularly elucidative. In the 1996

campaign (Fig. 7), Bittium sp. clusters

with the alga Dictyota dichotoma, Parvi-

cardium vroomi clusters with Halopteris

filicina and Tricolia pullus azorica clusters

with Asparagopsis armata. In the 1997

campaign, a strong dichotomy between

molluscs and algae was found, no mol-

luscs being associated to any alga

species. If the same procedure (cluster

analysis of pooled algae-molluscs data)

is repeated, now in R-mode, there is a

clear separation between shallow levels

(s) and medium-deep depths (Fig. 8).

Major differences among pre-

defined groups (see Table IV) do occur

between medium depth replicates of

1996 and 1997 (m96/m097), between the

deepestilevels of 1996 FPand 1997

(d96/d97) and between all samples of

1996 and 1997 (s96-m96-d96 / s97-m97-

d97) (see Table V).

Average similarity values of pre-

defined groups (s, m, d) is higher in

1996 than in 1997, Bittium sp. and Trico-

lia pullus azorica being responsible for

this. Average similarity decreases with

depth, when the same level is compared

in the two years (s96/s97, m96/m97 and

d96/d97). Similarities among shallow

and medium depth levels are best

explained by the two previous taxa plus

Jujubinus pseudogravinae. In the deepest

levels (d496/d97), we must add the

muricid Ocinebrina aciculata (Lamarck,

1822) to Bittium sp. and Tricolia pullus

azorica (cf. Table VD.

Crassadoma pusio and Alvania sleursi

are responsible for most of the differ-

ences detected between levels (s96/597,

m96/m97 e d96/d97), their importance

increasing with depth. Actually, total

number of Crassadoma pusio individuals

in 1997 is less than 10% the number in

1996; Alvania sleursi specimens in 1997

are about half total numbers in 1996. In

1996, Rissoa guernei and Alvaniía angioyi,

both common species in the shallow

levels, distinguish these levels from the

medium and deep ones, Parvicardium

vroomi, uncommon from 20m on,

explaining the medium-deep level dif-

ferences together with Bittium sp. In

1997, Bittium sp. and Tricolia pullus

azorica splits medium from deep levels.

When all levels are pooled by year (s96-

m96-d96 / s97-m97-d97), Alvania cancel-

lata, Alvania sleursi, Crassadoma pusio and

Manzonia unifasciata (more abundant in

1996), and Bittium sp. and Parvicardium

vroomi (more abundant in 1997) are

responsible for the differences detected

(Table VID.

DISCUSSION

No specific molluscs /algae associa-

tions were found in this work, however,

a certain algal preference was detected

because of the high densities of a few

molluscan taxa in some algae. The most

obvious examples were Bittium sp. in

Halopteris filicina and Pterocladiella capil-

lacea, Parvicardium vroomi in Halopteris

filicina, and Tricolia pullus azorica in

Asparagopsis armata, P. capillacea and Dic-

tyota dichotoma. It appears that the algae

provide a good microhabitat for the

molluscs, rather than being their main

source of food. It is interesting to note

the common presence of Tricolia pullus

azorica in Asparagopsis armata, a red

algae that is usually devoid of molluscs.

This algae was introduced into the

Atlantic and Mediterranean in the

beginning of the 20' century (FELD-

MANN AND FELDMANN, 1942) and is now

widely distributed in the north-eastern

Atlantic, from the Bristish isles south to

Senegal, the Azores, Madeira and

Canary Islands (DIXON, 1964).

Iberus, 21 (1), 2003

Table VI. SIMPER results. Data transformed by VVx. Aan: Alvania angioyi; Aca: Alvania

cancellata; Asl: Alvania sleursi, Ana: Anachis avaroides; Bit: Bittium sp.; Car: Cardita calyculata; Cra:

Crassadoma pusio; Gib: Gibbula delgadensis, Juj: Jujubinus pseudogravinae; Man: Manzonia unifas-

ciata; Mar: Marshallora adversa; Nas: Nassarius incrassatus, Oci: Ocinebrina aciculata; Par: Parvicar-

dium vroomi; Ris: Rissoa guernei, Tri: Trichomusculus semigranatus; pa: Tricolia pullus azorica.

Other symbols as in Table IV.

Groups Average similarity

596 16.29

m96 12.58

d96 45.71

597 47.33

m97 42.28

d97 45.71

s96 / 597 53.10

m96 / m97 50.10

d96 / 497 45.83

(s)6-m96-196) 67.13

(s97-m97-497) 43.88

BULLOCK ET AL. (1990) did not find

also any specific biological associations

of molluscs to algae in Sáo Miguel

Island. According to these authors, the

majority of the molluscs found in their

samples were common in several

species of algae (e.g.: Omalogyra atomus,

very common in Pterocladiella capillacea

and Stypocaulon scoparia, Gibbula magus

(Linnaeus, 1758) in Padina pavonica and

Stypocaulon scoparia, and the small

bivalve Lasaea adansoni (Gmelin, 1791) in

Sargassum vulgare and Gelidium spinulo-

sum). COSTA AND ÁVILA (2001) found

Bittium sp. with high-density values in

monospecific samples of Stypocaulon sco-

paria and of Halopteris filicina; with

lower densities, Setia subvaricosa, Tricolia

Taxa/Species

Bit — Tpa — Man — Juj — Par — Asl — Ris — Aan

Bit — Tpa — Juj — Par — Cra — As] — Car

Bit — Tpa — Asl — Cra - Oci

Bit - Tpa

Bit

Bit — Tpa - Mar

Bit — Tpa — Juj - Nas

Bit — Tpa -Juj - Nos

Bit — Tpa - Oci

Bit — Tpa — Juj — As] — Cra — Par — Car — Man — Oci— Aca - Nos

Bit - Tpa

pullus azorica and Rissoa guernei were

also common molluscs.

The differences detected in the rela-

tive abundances of the malacological

communities associated to algae, are usu-

ally related to different habitat condi-

tions, both biotic (species and type of

most common algae, algal biomass, pre-

dation intensity) as abiotic conditions (es-

pecially the hydrodynamics and the total

amount of sediment that algal fronds

may trap and hold) (AZEVvEDO, 1991a).

Feeding habits may influence

mollusc's distribution by depth. Bittium

sp. and Tricolia pullus azorica, the two

most abundant taxa, were found along

the entire transect. Such distribution

may be attributed to their detritivorous

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

Tabla VI. Resultados de SIMPER. Datos transformados con by WWx. Aan: Alvania angioyi; Aca:

Alvania cancellata; As/: Alvania sleursi; Ana: Anachis avaroides; Bit: Bittium sp.; Car: Cardita caly-

culata; Cra: Crassadoma pusio; Gíb: Gibbula delgadensis; Juj: Jujubinus pseudogravinae; Man:

Manzonia unifasciata; Mar: Marshallora adversa; Vas: Nassarius incrassatus; Oc¿: Ocinebrina aci-

culata; Par: Parvicardium vroomi; Rís: Rissoa guernei; 771: Trichomusculus semigranatus; 7pa: Tri-

colia pullus azorica. Resto de símbolos como en la Tabla IV

Toxa/Species (9) Cumulotive %

(Ratio of each taxa/species)

13.59 - 9.57 -7.21 — 6.91 — 6.52 — 6.49 — 5. 68 - 5.51 61.85

(10.00 — 6.49 - 3.77 —7.11 —8.01 — 8.86 — 10.24 — 7.97)

15.50 10.11 —7.33 — 6.95 — 6.88 — 6.45 — 6.42 59.64

(9.22 — 6.31 — 8.48 — 6.42 —7.31 —7.70- 5.25)

19.69 - 13.75 — 9.58 — 9.49 — 6.75 59.26

(4.94 — 4.13 — 4.51 — 3.30 — 5.08)

33.11 - 17.53 50.64

(2.79 - 2.58)

3172 31.72

(2.30)

32.87 — 15.84 — 15.46 64.17

(2.05 1.92 — 1.66)

24.32 — 13.82 — 7.50 -5.34 50:98

(3.18 - 3.32 -1.72-1.59)

22.11 - 11.35 — 7.82 — 6.33 48.11

(3.16-=1.72—1.73-1.42)

28.64 — 15.45 - 8.91 53.00

(2.70-2.90- 1.54)

16.44 - 11.12 7.69 —7.45— 6.96 — 5.20 — 4.88 — 4,54 — 4.04 - 3.51 - 3.22 75.04

(7.74 5.97 - 2.81 — 5.98 — 4.26-1.66 1.69 - 1.54 — 2.03 — 1.59 — 1.67)

33.88 — 15.39 49.27

(2.36 — 1.68)

habits, although Tricolia is thought to luscs included. Marshallora adversa

feed also on alga tissues and diatoms

(FRETTER AND GRAHAM, 1977; BORJA,

1986; GRAHAM, 1988). Shallow-water ris-

soids (Alvania angioy1, A. mediolittoralis,

Manzonia unifasciata and Rissoa guernel)

as well as deep-water (Alvania cancellata

(da Costa, 1778) and A. sleursi) also feed

preferably on diatoms and epiphyte

microalgae, as well as on the food

trapped by the sediment that accumu-

lates in the base and in the interstices of

the branches of algal fronds (FRETTER

AND GRAHAM, 1981; GRAHAM, 1988;

AVILA, 2000b). Anachis avaroides and

Nassartus incrassatus (Stróm, 1768) are

carnivorous or scavengers (N. incrassa-

tus) feeding on small invertebrates, mol-

(Montagu, 1803) probably feeds on

sponges (GRAHAM, 1988) and all

bivalves herein are suspension feeders.

The methodology followed in this

work was effective in the establishment

of the zonation of the most common

taxa of molluscs, but did not properly

answered to the question of the mol-

luscs /alga biological associations. This

is only possible with a broadly system-

atic study of monospecific algae

samples (see BULLOCK ET AL., 1990;

COSTA AND ÁVILA, 2001).

The time of sampling is also a non-

negligible factor. AZEVEDO (199la)

studied the infralittoral communities of

molluscs along a period of one year in

no)

5 |

Iberus, 21 (1), 2003

Table VII. SIMPER results. Data transformed by VWVx. Aan: Alvania angioyi, Aca: Alvania cance-

llata; Asl: Alvania sleursi; Ana: Anachis avaroides, Bit: Bittium sp.; Car: Cardita calyculata; Cra:

Crassadoma pusio; Gib: Gibbula delgadensis; Juj: Jujubinus pseudogravinae, Man: Manzonia unifas-

ciata, Mar: Marshallora adversa, Nas: Nassarius incrassatus; Oci: Ocinebrina aciculata; Par: Parvicar-

dium vroomi, Ris: Rissoa guernei; Tri: Trichomusculus semigranatus; Tpa: Tricolia pullus azorica.

Other symbols as in Table IV.

Groups Average dissimilarity

596 / 597 51.10

m96 / m97 56.4]

d96 / 497 60.00

596 / m96 29.18

m96 / d96 37.36

596 / 096 41.65

$97 — m97 52.82

m97 / 497 56.28

597 / 497 58.67

(596597) / (m96-m97) 49.92

(596597) / (096-497) 55415

(m96-m97) / (496-497) 52.66

(s76-m96-496) / (s97-m97-497) 56.22

two localities of Sáo Miguel Island

(Ribeirinha, located in the north and

Caloura, in the south shore). During this

time, he found that the molluscs'

density was higher in the beginning of

the summer, decreasing then and reach-

ing its lower values during the winter

times. At Ribeirinha, the most abundant

species (>1 mm) were Bittium sp. Tricolia

pullus azorica, Jujubinus pseudogravinae,

Nassarius incrassatus, Parvicardium

vroomi, Anachis avaroides and Ocinebrina

aciculata (AZEVEDO, 19910).

In this study, with the sole exception

Of N. incrassatus, all other taxa were also

the commonest at Sáo Vicente (Capelas)

in 1996 and in 1997. These results point

Taxa/species

Man — Par — Ris — Asl Cra

Bit — Par— Cra - Asl

Cra — Asl - Tpa

Ris — Ana — Aon - Gib

Par - Bit

Ris — Man — Par - Aan

Cra — Par — Bit — Mar — Man — Tri - Aca

Bit - Tpo

Cra — As — Man — Bit — Par - Aca

to a remarkable stability in species com-

position as well as in mean abundance

values of the malacological communi-

ties associated to macroalgae in the

rocky shores of the Azores archipelago,

however, Tricolia pullus azorica, Jujubinus

pseudogravinae, Manzonia unifasciata and

Crassadoma pusio were 2 to 4 times more

abundant in July 1996 than in 1997, and

Parvicardium vroomi was commonest in

1997. Some of the species found only in

one of the years are accidental (e.g.:

Melarhaphe neritoides, Pisinna glabatra,

Pedipes pedipes and Lasaea adansoni)

which belong to the supralittoral, and

others are just rare or encountered by

chance, as is the case of the not identi-

AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

Tabla VII. Resultados de SIMPER. Datos transformados con by WNx. Aan: Alvania angioyi; Aca:

Alvania cancellata; As/: Alvania sleursi; Ana: Anachis avaroides; Bit: Bittium sp.; Car: Cardita caly-

culata; Cra: Crassadoma pusio; Gíb: Gibbula delgadensis; Ju¿: Jujubinus pseudogravinae; Man:

Manzonia unifasciata; Mar: Marshallora adversa; Nas: Nassarius incrassatus; Ocí: Ocinebrina aci-

culata; Par: Parvicardium vroomi; Rís: Rissoa guernei; 771: Trichomusculus semigranatus; 7 pa: Tri-

colia pullus azorica. Resto de símbolos como en la Tabla IV.

Taxa/species (%)

Ratio of each taxa/species)

1.32—5.25—4.85 — 4.83 —4.51

(1.76=1.70—1.74—1.50— 2.06)

5.63 — 5.05 — 4.94 — 4.64

MS O SS)

6.47 —6.24—4.1|

(1.46 —1.72—1.66)

1.57 4.64 —4.40— 4.32

(3.06 —1.40—1.43—1.55)

5.68 4.16

(1.47 1.42)

6.68 — 6.43—6.01 - 4.77

NI IMOSASO)

Cumulotive %

26.16

25.51

17.41

20.93

9.84

26.96

3:09 3.02 24.30 —4.104/07 3:00 3.55 30.53

A SO 00 1011 2:20)

6.06 5.73

(1.57 1.49)

no taxa/species with ratio > 1.40 -

no taxa/species with ratio > 1.40

5.05 4.90 — 4.84 — 4.82 — 4.71 — 3.43

(1.60 — 1.49 — 1.40 —1.44—1.55— 1.58)

fied Eulimidae, associated with a host.

A more detailed study of the population

dynamics of their individual species

could help to clarify the situation,

mainly by stating the role of recruitment

on these disparate abundances.

ACKNOWLEDGEMENTS

This article was part of a MSc. thesis.

I am very grateful to José M. N.

Azevedo who critically read and orien-

tated the thesis. I also acknowledge Joáo

Carlos Marques (IMAR/ University of

Coimbra) and António Frias Martins

(University of the Azores) for advice

11:79

and corrections. Two anonymous

reviewers contributed many helpful

comments and criticisms. The Marine

Biology team of the University of the

Azores is acknowledged (José Azevedo,

Ana Neto and Ana Cristina Costa, as

well as the technicians Joáo Brum and

Sandra Monteiro). Several students

helped during the sorting phase (Joana

Duarte, Margarida Lourenco, Pedro

“Bué” Cerqueira, Hugo Renato Calado

and Rui Amen). This study was sup-

ported by PRAXIS/2/2.1/BIA/169/94.

The author was supported by grants

FCT” (PRAXIS —XXI/BIC/2788/96,

PRAXIS XXI BM/20521/99 and

SFRH/BD/5115/2001).

Iberus, 21 (1), 2003

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Iberus, 21 (1), 2003

Appendix 1. Molluscs collected at Porto das Baleias, Sáo Vicente, Capelas in July 1996 (non-trans-

formed data).

Apéndice 1. Moluscos recolectados en Porto das Baleias, Sáo Vicente, Capelas en Julio de 1996 (datos no

transformados).

Date (1996 July)

Depth (m)

DBUA

Toxo/Species M Quadrote

Aconthochitona fascicularis

Acmaeo virgineo

Alvania sp.

Alvanio angioyi

Alvanio concellata

Alvania mediolittoralis

Alvania poucheti

Alvanio sleursi

Anachis avoroides

Anomia ephippium

Aplysia cf. depilans

Aplysio sp.

Árco tetrogona

Bittium sp.

Calliostomo sp.

Cordita colyculata

Cerithiopsis tuberculoris

Cerithiopsis sp.

Cheirodonta pallescens

Columbella adansoni

Crassadomo pusio

Crisilla postrema

Envilia castanea

Gibbula magus

Gibbula delgadensis

Gouldia minima

Hoedropleura septangularis

Holiotis tuberculata

Jujubinus pseudogravinae

Losaeo odonsoni

Limaria hians

Lime loscombii

Manzonia unifasciata

Morshallora adversa

Mitromorpha azorensis

Monophorus erythrosoma

Nassarivs incrassatus

Ocinebrina aciculota

Odostomella doliolum

Odostomia sp.

Omologyra atomus

Porvicardium sp.

Parvicardium vroomi

Pedipes pedipes

Pisinna glabatra

Rhomboidella prideauxi

Rissoa guernei

Setia subvaricosa

Seto sp.

Similiphora similior

Sinezona cingulata

Trichomusculus semigranatus

Tricolia pullus azorica

Vitreolina sp.

TOTAL

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AVILA: Littoral molluscs of Azores: ecology and biological associations to algae

Appendix 2. Molluscs collected at Porto das Baleias, Sáo Vicente, Capelas in July 1997 (non-trans-

formed data).

Apéndice 2. Moluscos recolectados en Porto das Baleias, Sáo Vicente, Capelas en Julio de 1997 (datos no

transformados).

Date (1997 July)

Depth (m)

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O Sociedad Española de Malacología ——_—_—_—_——T— Iberus, 21 (1):35-42, 2003

The genus Granulina (Gastropoda, Marginellidae) from the

Atlantic Iberian Pliocene with description of a new species

from Portugal

El género Granulina (Gastropoda, Marginellidae) en el Plioceno

Atlántico Ibérico y descripción de una especie nueva para Portugal

Rafael LA PERNA*, Bernard LANDAU** and Carlos Marques da SILVA***

Recibido el 27-X1-2001. Aceptado el 16-1X-2002

ABSTRACT

The marginellid Granulina choffati n.sp. is described from Pliocene, uppermost Zanclean

to lower Piacenzian, shallow-water sandy beds cropping out in central-west Portugal. This

is the first record of Granulina from the Atlantic European Pliocene. G. choffati n.sp. is

markedly similar to G. elliptica La Perna, 2000, known from the Pliocene of Sicily, and

both species share close similarities to the Recent West African species G. nofronii

Smriglio, Gubbioli and Mariottini, 2001. The two Pliocene species are regarded as ther-

mophilic elements of southern affinity. The Pliocene distribution of Granulina in the East

Atlantic ranged at least up to 40%N along the Iberian coasts and it is hypothesized that

this limit shifted southward because of the Plio-Pleistocene cooling pulses.

RESUMEN

Se describe una nueva especie de Marginellidae, Granulina choffati n.sp., procedente de

afloramientos arenosos someros del Plioceno (Zancliense superior-Piacenziense) en el cen-

tro-oeste de Portugal. Esta es la primera cita del género Granulina para el Plioceno Atlán-

tico Europeo. G. choffatti n.sp. es muy similar a G. elliptica La Perna, 2000, conocida en

el Plioceno de Sicilia, y ambas especies muestran marcadas semejanzas a la especie

actual del oeste de Africa, G.nofronii Smriglio, Gubbioli y Mariottini, 2001. Las dos espe-

cies pliocénicas se consideran elementos termófilos con afinidad meridional. La distribu-

ción de Granulina en el Plioceno del Este Atlántico llegaba hasta almenos 40? N en la

costa ibérica y se presume que este límite se haya trasladado hacia el sur a causa de cri-

sis de enfriamiento plio-pleistocénico.

KEY WORDS: Gastropoda, Granulina, new species, paleobiogeography, palaeoclimatology, Pliocene,

Portugal.

PALABRAS CLAVE: Gastropoda, Granulina, especie nueva, paleobiogeografía, paleoclimatología, Plioceno,

Portugal.

* Dipartimento di Geologia e Geofisica, Universitá di Bari, Via E. Orabona 4, 1-70125 Bari, Italia (r.laper-

naO2geo.uniba.it)

** International Health Centres, Avenida Infante de Henrique 7, Areias Sáo Joáo, P-8200 Albufeira, Portugal

(bernie.landauCbtinternet.com)

*** Departamento e Centro de Geologia, Universidade de Lisboa, Rua da Escola Polítecnica 58, P-1250-102

Lisboa, Portugal (Paleo.CarlosCfc.ul.pt)

Iberus, 21 (1), 2003

W 8? 43

PORTUGAL A

Pombal A]

Vale de Freixo O / ¡M0 ]

/) |

SS ><

847 LEIRIA

Coimbra

E 7 3 LISBOA

to Lisboa / 78 km

Figure 1. Location of the Vale de Freixo Pliocene outcrop in central-west Portugal.

Figura 1. Localización del yacimiento de Vale de Freixo en en centro-oeste de Portugal.

INTRODUCTION

Several recent works devoted to the

marginellid genus Granulina Jousseame,

1888 have provided a fairly good

knowledge of the composition of this

genus in the Mediterranean and the

adjacent Atlantic (Ibero-Moroccan Gulf

south to Gulf of Guinea), and also in the

Mediterranean Pliocene and Pleistocene

(La PERNA, LANDAU AND MARQUET,

2002, with references).

Granulina is a “southern” genus, not

present in the North European waters.

The Iberian coasts are the northernmost

area of distribution of this genus in the

East Atlantic, but little is known about

its Occurrence in this area. NOBRE (1936)

reported G. clandestina (Brocchi, 1814)

from Portugal with no precise indication

of locality, but this record was based on a

misidentification, as G. clandestina is an

extinct Pliocene species whose name was

widely and uncritically used until the

revision made by GOFAsS (1992). A list of

modern FEuropean and Northwest

African marginellids (MUÑIZ SOLIS,

1987) and a recent popular book on Por-

tuguese molluscs (MACEDO, MACEDO

AND BORGES, 1999) reported several

species of Granulina, including “G. clan-

destina”. None of the other species

reported, G. guancha (d'Orbigny, 1839),

G. minusculina (Locard, 1897) and G.

parvulina (Locard, 1897) (= G. occulta

Monterosato, 1869) is proved to occur

north of the Ibero-Moroccan Gulf. GOFAS

(1992) stated that no species of this

genus has ever been found in the Biscay

Bay. Although the occurrence of Gran-

ulina in the Ibero-Moroccan Gulf is doc-

umented (GOFAs, 1992), no species was

found in the Sagres and Olháo areas

(southern Portugal, Ibero-Moroccan-

Gulf), in spite of extensive samplings

carried on by the Muséum National

d'Histoire Naturelle (Paris) in 1988

(Gofas, pers. comm.). Therefore, the

northern limit of Granulina in the Eastern

Atlantic must be within the Ibero-Moroc-

can Gulf excluding the southern Portu-

gal, at least for the shelf species (Fig. 10).

La PERNA ET AL.: Atlantic Iberian Pliocene Granulina and a new species from Portugal

No representative of Granulina had

been so far reported from the Atlantic

European Pliocene, but there are a few

records from the Upper Oligocene and

Lower Miocene of the Aquitaine basin

and from the Upper Oligocene of the

North Sea basin (LOZOUET, 1997 and

pers. comm., 2002). Recent studies on a

rich molluscan fauna from Pliocene

beds cropping out at Vale de Freixo, in

central-west Portugal (Fig. 1), allowed

the discovery of several specimens of

the genus, most of them belonging to an

undescribed species. Nevertheless, this

new species is not the only one of the

SYSTEMATICS

genus occurring at Vale de Freixo, a

single shell of another one was found,

but it is too badly preserved to be

unequivocally determined. It is however

similar to G. detruncata, recently

described from the Pliocene of Malaga

(southern Spain) by LA PERNA ET AL.

(2002).

Like the previous study on the

Pliocene species from the southern

Iberian Peninsula (LA PERNA ET AL.,

2002), the present work focuses on the

history of Granulina in the Mediter-

ranean and adjacent Atlantic through

the Plio-Quaternary.

Class GASTROPODA Cuvier, 1797

Order NEOGASTROPODA Thiele, 1929

Family MARGINELLIDAE Fleming, 1828

Genus Granulina Jousseame, 1888

Granulina choffati n.sp. (Figs. 2-6)

Type material: Holotype and 3 paratypes, National Natural History Museum of the Lisbon Uni-

versity (MNHN /UL.I1.413-416), 9 paratypes in B. Landau coll., 1 paratype in R. La Perna coll.

Examined material: Only the type material.

Type locality: Vale de Freixo, Pombal region in central-west Portugal (Fig. 1), Pliocene, upper-

most Zanclean to lower Piacenzian fine sandy beds.

Etymology: After Léon-Paul Choffat (born in 1849, in Porrentruy; died in 1919, in Lisbon), classi-

cal Portuguese geologist and palaeontologist of Swiss origin who discovered and studied the

first Pliocene outcrops and molluscan faunas from Portugal.

Description: Shell minute, with

immersed spire, elliptic-elongate in

shape, maximum diameter at mid-

length of shell, lenght/diameter ca 1.5.

Posterior end slightly truncated to

rounded, covered by a thin more or less

developed callus. Siphonal notch faintly

distinct. Lip moderately thickened,

forming a regular arch, broader at mid

length. Lip denticulations moderately

fine, well defined, somewhat roundish.

_ Four columellar plications, the upper-

most two slightly smaller, obscurely

excavated inside aperture by a shallow

sulcus. Thin ill-defined inner parietal

callus ridge running from posterior end

to columellar plications. Outer parietal

callus not distinct. Aperture narrow.

Surface smooth but crossed by fairly

well distinct growth striae. Holotype:

lenght 2.1 mm, diameter 1.4 mm.

Paratypes: lenght 1.8 to 2.2 mm.

Distribution: So far, only known from

the type locality. The material comes

from a Pliocene mostly sandy sequence,

containing a particularly diverse and

well preserved fauna. Recent studies on

this fauna (GILI, SILVA AND MARTINELL,

1995; SILVA, 1996; NOLF AND SILVA, 1997;

SILVA, LANDAU AND MARTINELL, 2000;

SILVA, 2001) indicate a marine shallow-

water (infralittoral) environment with

relatively high water temperatures. The

age of the Vale de Freixo fauna was

referred to uppermost Zanclean to lower

Piacenzian (Discoaster tamalis, CN12a,

biozone of OKADA AND BUKRY, 1980) by

CACHAO (1990; emended by SILVA, 2001,

Iberus, 21 (1), 2003

by means of Strontium *%Sr/*Sr dating)

and to the Mediterranean Pliocene Mol-

luscan Unit 1 of RAFFI AND MONEGATTI

(1993) and MONEGATTI AND RAFFI (2001)

by SILVA (2001). For general stratigrafical

setting, Vale de Freixo graphic columnar

section, and additional references see

SILVA ET AL. (2000).

Remarks: The most similar fossil

species is Granulina elliptica La Perna,

2000 (Figs. 7-9) described from the

Lower-Middle Pliocene of Sicily (LA

PERNA, 2000) (Fig. 10). In both species

the shell has a markedly elliptic shape,

with maximum diameter at mid length.

Such a shape is rather unusual for the

genus, since the maximum diameter

often is within the posterior (adapical)

third, giving an ovate (rather than ellip-

tic) shell outline. G. choffati n.sp. is even

more regularly elliptic than G. elliptica,

in which the maximum diameter tends

to be slightly above the mid point. A

parietal callus ridge almost inside the

aperture is present in the shell of both

species, as well as an inner parietal

sulcus producing an “excavation” of

columellar plications (LA PERNA ET AL.,

2002). However, the callus ridge, sulcus

and excavated plications are much more

developed in G. elliptica than in G. chof-

DISCUSSION

Although most species of Granulina

differ from one other only by subtle con-

chological differences, a number of shell

features have been recently considered

(GOFAs, 1992; LA PERNA, 1999; BOYER

AND ROLÁN, 1999; LA PERNA ET AL.,

2002). As discussed by LA PERNA ET AL.

(2002), morphological affinities may be

of help to infer phyletic closeness within

this genus, in spite of such a monoto-

nous conchological pattern. An attempt

of clustering some Pliocene, Pleistocene

and Recent species was made by LA

PERNA ET AL. (2002).

When G. elliptica was described, the

impossibility to recognise any marked

similarity with other fossil or living

species was stressed, but the finding of

G. choffati n.sp. now deprives G. elliptica

fati n.sp., and these are the main distinc-

tive features between these species. Lip

denticulations are somewhat roundish

in both species, particularly in G. ellip-

tica, in which they are also notably

coarser. The outer parietal callus (LA

PERNA, 1999; LA PERNA ET AL., 2002) is

not distinct in both species. In G. elliptica

the shell is more swollen, and the outer

lip lacks the central broadening which is

more or less developed in G. choffati

n.sp. It is also worth noting the shallow-

water (infralittoral) distribution of both

species.

A Recent shelf species, G. nofronii,

recently described by SMRIGLIO, GUBBI-

OLI AND MARIOTTINI (2001) from North-

west Africa (Western Sahara and Mauri-

tania) shares similarities with the two

Pliocene species. Its elliptic shape is par-

ticularly similar to that of G. choffati

n.sp., including a certain broadening of

the lip in its median part. Also plications

shape and callus ridge strength are par-

ticularly similar to those of G. choffati

n.sp., while the coarse lip denticulation .

resembles that of G. elliptica. G. nofronii

differs from both species by being a

little larger, slightly more slender and

rostrated, and with a better defined

siphonal slope-break.

of such a “uniqueness”. These two

species are so similar that a close

phyletic relation between them is highly

likely.

LA PERNA ET AL. (2002) recognised

amongst the Mediterranean Pliocene

shelf species of Granulina a mixture of

warm and warm-temperate taxa, due to

the occurrence of some extinct species

with “African affinities”, such as G. clan-

destina (a similar Recent species, G.

parilis Gofas and Fernandes, 1988, is

known from the Gulf of Guinea),

together with a more temperate stock,

well represented by G. marginata

(Bivona, 1832) and G. boucheti Gofas,

1992, which are now the most common

shallow-water species of the genus in

the Mediterannean.

La PERNA ET AL.: Atlantic Iberian Pliocene Granulina and a new species from Portugal

Figures 2-6. Granulina choffati n.sp. 2, 3: holotype, 2.1 mm; 4: paratype 1, 2.0 mm; 5: paratype 2,

2.0 mm, 6: paratype 3, 1.8 mm. Figures 7-9. Granulina elliptica. 7: holotype, 2.1 mm; 8: paratype,

2.1 mm; 9: paratype, 2.3 mm.

Figuras 2-6. Granulina choffati n.sp. 2, 3: holotipo, 2.1 mm; 4: paratipo 1, 2.0 mm; 5: paratipo 2,

2.0 mm; 6: paratipo 3, 1.8 mm. Figuras 7-9. Granulina elliptica. 7: holotipo, 2.1 mm; 8: paratipo,

2.1 mm; 9: paratipo, 2.3 mm.

Iberus, 21 (1), 2003

G. choffati (Plio)

G. elliptica (Plio)

occurrences

400

370 0

G. nofronii

(Recent) >=

200

400 200

| Present day

Pliocene

occurrences

Plio-Pleistocene

“ shouthward shift

Figure 10. Present day Eastern Atlantic and Mediterranean distribution of Granulina and occur-

rences of Granulina choffati n.sp., G. elliptica La Perna, 2000, and G. nofronii Smriglio et al., 2000.

Figura 10. Distribución actual en el Atlántico este y Mediterráneo de Granulina y citas de Granulina

choffati n.sp., G. elliptica La Perna, 2000, and G. nofronii Smriglio et al., 2000.

Also G. elliptica and G. choffati n.sp.

may thus represent a group of species

with a southern affinity, if the West

African G. nofronii is regarded as a

living representative of the same

lineage. This does not imply a direct

phyletic relation between these two

species (1.e., G. elliptica may be not the

ancestor of G. choffati n.sp., or vice-

versa). For the time being, the key point

is to recognize groups of closely related

species, sharing similar ecologic and

biogeographic distribution. Changes in

the distribution of these groups should

be referred to major climatic changes,

whose strong effects on distribution

and diversity of Plio-Pleistocene

benthic faunas are well known (e.g., DI

GERONIMO, DI GERONIMO, LA PERNA,

Rosso AND SANFILIPPO, 2001; MON-

EGATTI AND RAFFI, 2001). Also the distri-

bution and composition of Granulina

and of marginellids in general, which

typically have a warm-water distribu-

tion, must have been controlled by the

Plio-Pleistocene cooling events. Actu-

ally, the diversity of Granulina in the

shelf waters was higher in the Pliocene

than in the present days (LA PERNA ET

AL., 2002 and unpubl. data; CHIRLI,

2002). Extinction of warm-water taxa,

and/or local disappearance due to

southward shifting, is the most impor-

tant aspect of the Plio-Pleistocene

history of shelf molluscs (e.g., STANLEY

AND RUDDIMAN, 1995; MONEGATTI AND

RAFFI, 2001).

CONCLUSIONS

During the Pliocene, the genus Gran-

ulina ranged at least up to ca. 40N (the

latitude of Vale de Freixo) along the

West Iberian coasts (Fig. 10). It should

be admitted that such a Pliocene distrib-

ution represented but a step in the

overall southward shift of Granulina in

the Eastern Atlantic since its appearance

(or its first documented occurrence) at

higher latitudes in the Upper Oligocene

(see introduction). Other Portuguese

La PERNA £7 4£.: Atlantic Iberian Pliocene Granulina and a new species from Portugal

Pliocene gastropod taxa show evidences

of a similar southward migration (e.g.,

among Nassarius, GILI, SILVA AND MAR-

TINELL, 1995; Solariella, SILVA, LANDAU

AND MARTINELL, 2000). From the Middle

Pliocene (ca. 3.0 Ma) on, the southward

shift of the Atlantic species of Granulina

was, probably, more intense than previ-

ously, reflecting the sucessive Plio-Pleis-

tocene sharp cooling pulses, the species

with stronger termophilic affinities

becoming rapidly extinct. The same

pattern of extinction and /or local disap-

pearance also occurred in the Mediter-

ranean, Where, simultaneously, some

endemic taxa were appearing (LA

PERNA ET AL., 2002).

BIBLIOGRAPHY

BOYER, F. AND ROLÁN, E. 1999. Granulina fer-

nandesi (Gastropoda: Volutacea), a new

species from Cape Verde Islands, and some

considerations on the genus Granulina. Iberus,

17 (2): 1-10.

CACHAO, M.,, 1990. Posicionamento biostrati-

gráfico da Jazida Pliocénica de Carnide

(Pombal). Gaia, 2: 11-16.

CHIRLI, C., 2002. Malacofauna Pliocenica Toscana.

Vol. 3. Muricoidea e Cancellaricidea. Stamperia

e Legatoria Pisana, Agnano Pisano, 92 pp. +

37 pls.

DI GERONIMO, I., DI GERONIMO, R., LA PERNA,

R., ROSSO, A. AND SANFILIPPO, R., 2001.

Cooling evidence from Pleistocene shelf

assemblages in SE Sicily. In Hart, M.B (Ed.):

Climates: Past and Present. Geological

Society, London, Special Publication, 181:

113-120.

GILI, C., SILVA, C.M. DA AND MARTINELL, J.,

1995. Pliocene nassariids (Mollusca: Neogas-

tropoda) of central-west Portugal. Tertiary

Research, 15 (3): 95-110.

GOFAS, S., 1992. Le genre Granulina (Margin-

ellidae) en Méditerranée et dans 1'Atlan-

tique Oriental. Bollettino Malacologico, 28 (1-

4): 1-26.

LA PERNA, R., 1999. Pleistocene and Recent

Mediterranean species of Granulina (Gas-

tropoda, Marginellidae), with description of

four new species. Bollettino Malacologico, 34

(1-4): 43-52.

La PERNA, R., 2000. Granulina elliptica n. sp.

and comments on the Mediterranean

Pliocene species of Granulina (Gastropoda,

Marginellidae). Bollettino Malacologico, 35 '

(1-4): 53-55.

ACKNOWLEDGEMENTS

The authors wish to acknowledge

Pierre Lozouet (Muséum National

d'Histoire Naturelle, Paris) and Serge

Gofas (Universidad de Málaga) for rele-

vant information, Isabel Puigdevall

(Universita di Bari) for her kind assis-

tance with Spanish language, two

anonymous referees and Serge Gofas for

critical reading and comments.

Contribution of the Portuguese FCT

Project 32724 /99-Comparative (palaeo)-

environmental analysis of oceanic and

coastal domains, over the last 20 Ma,

based on calcareous nannoplankton

(CANAL), co-financed by FEDER.

LA PERNA, R., LANDAU, B. AND MARQUET, R.,

2002. Species of Granulina (Gastropoda, Mar-

ginellidae) from the Pliocene of Málaga

(southern Spain) with description of four

new species. Cainozoic Research, 1 (1-2): 111-

120.

LOozoOUErT, P., 1997. Le domaine atlantique

européen au Cenozoique moyen: diversité et

évolution des gastéropodes. Unpublished

Thesis (Paris MNHN), 309 pp. + annexes 52

pp., 112 figs.

MACEDO, M.C., MACEDO, M.I. AND BORGES, J.P.,

1999. Conchas Marinhas de Portugal. Editorial

Verbo, Lisboa, 516 pp.

MONEGATTI, P. AND RAFEL S., 2001. Taxonomic

diversity and stratigraphic distribution of

Mediterranean Pliocene bivalves. Palaeo-

geography, Palaeoclimatology, Palaeoecology,

16SA7ISIOS:

MUÑNIZ SOLIS, R., 1987. Catalogo preliminar de

la familia Marginellidae Fleming, 1828 (Gas-

tropoda, Prosobranchia) en las provincias

Lusitana, Mediterránea y Mauritana. Reseñas

Malacologicas, V, Sociedad Española de Mala-

cologia, 60 pp.

NOBRE, A., 1936. Moluscos Marinhos de Portugal.

II vol. Companhia Editora do Minho, Barce-

los, 378 pp.

NOLF, D. AND SILVA, C.M. DA, 1997. Otolithes

de poissons pliocenes (Plaisancien) de Vale

de Freixo, Portugal. Revue de Micropaléon-

tologie, 40 (3): 273-282.

OKADA, H. AND BUKRY, D., 1980. Supplemen-

tary modification and introduction of code

numbers to the low-latitude coccolith bios-

tratigraphic zonation. Marine Micropaleon-

tology, 5: 321-325.

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RAFFL S. AND MONEGATTI, P., 1993. Bivalve tax-

onomic diversity throughout the Italian

Pliocene as a tool for climatic-oceanographic

and stratigraphic inferences. Proc. 1%* RCANS

Congr., Lisbon 1992. Ciéncias da Terra (UNL),

12: 45-50.

SILVA, C.M. DA, 1996. Moluscos pliocénicos da

regiáo de Caldas da Rainha - Marinha Grande

- Pombal (Portugal). IM. Neogastropoda.

Conidae. Gaza, 12: 37-43.

SILVA, C.M. DA, 2001. Gastrópodes Pliocénicos

Marinhos de Portugal. Sistemática, Paleoecolo-

gia, Paleobiologia, Paleobiogeografia. Ph. D.

Thesis, University of Lisbon, 747 pp.

SILVA, C.M. DA, LANDAU B. AND MARTINELL J.,

2000. The genus Solariella (Mollusca,

Archaeogastropoda) from the Pliocene of

Vale de Freixo, Portugal: palaeobiogeo-

graphic and palaeoclimatic implications. Con-

tributions to Tertiary and Quaternary Geology,

37 (3-4): 57-65.

SMRIGLIO, C., GUBBIOLI, F. AND MARIOTTINL P.,

2001. New data concerning the presence of

Granulina Jousseaume, 1888 (Neogastropoda,

Cystiscidae) along the West African coast

and description of four new species. La

Conchiglia, 287: 54-59.

STANLEY, S.M. AND RUDDIMAN, W.F., 1995.

Neogene Ice Age in the North Atlantic

Region: Climatic Changes, Biotic Effects, and

Forcing Factors. In AAVV: Effects of Past

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Council, National Academic Press, Wash-

ington D.C., pp. 118-133.

O Sociedad Española de Malacología —_—_—_—_—_——T— Iberus, 21 (1): 43-56, 2003

Solenogastres molluscs from the BENTART Collection

(South Shetland Islands, Antarctica), with a description of

a new species

Moluscos Solenogastres de la Colección BENTART (Islas Shetland

del Sur, Antártida), con la descripción de una nueva especie

Oscar GARCÍA-ÁLVAREZ and Victoriano URGORRI”

Recibido el 27-VI-2002. Aceptado el 25-[X-2002

ABSTRACT

Four specimens from the BENTART Collection are studied (South Shetland Islands and Brans-

field Strait, Antarctic). Two specimens were identified as: Rhopalomenia carinata Salvini-Plawen,

1978 collected on a muddy, gravel bottom at a depth of 710 m, off Livingston Island, and

Rhopalomenia rhynchopharyngeata Salvini-Plawen, 1978 from a muddy bottom at a depth

of 235 m off Livingston Island. One specimen, collected from a fine muddy and sandy bot-

tom at a depth of 80 m off Livingston Island, belongs to the genus Neomenia, presenting fea-

tures which well distinguish the same from known species. lt is not, however, described as a

new species since the anatomic organisation of the posterior part of the animal is unknown.

And one specimen, collected on a muddy bottom at a depth of 80 m off Livingston Island,

is described as a the new species Dorymenia parvidentata. This article report on the

Solenogastres previously studied in the BENTART Collection.

RESUMEN

Se estudian cuatro ejemplares de la Colección BENTART (Islas Shetland del Sur y estrecho

de Bransfield, Antártida). Dos ejemplares fueron identificados como: Rhopalomenia carinata

Salvini-Plawen, 1978 recogida en un fondo fangoso y de gravas a 710 m de profundidad

en la Isla Livingston y Rhopalomenia rhynchopharyngeata Salvini-Plawen, 1978 procedente

de un fondo fangoso a 235 m de profundidad en la Isla Livingston. Un ejemplar, procedente

de un fondo de fango fino y arena a 80 m de profundidad en la Isla Livingston, pertenece

al género Neomenia y presenta rasgos que lo separan claramente de las especie conoci-

das, pero no es descrito como una nueva especie ya que no se conoce la organización ana-

tómica de la parte posterior del animal. Y un ejemplar, recogido en una fondo fangoso a

80 m de profundidad en la isla Livingston, es descrito como la nueva especie Dorymenia

parvidentata. Se informa sobree los Solenogastres ya estudiados de la Colección BENTART.

KEY WORDS: Neomenia sp., Rhopalomenia carinata, Rhopalomenia rhynchopharyngeata, Dorymenia parviden-

tata, Mollusca, Solenogastres, Antarctica.

PALABREAS CLAVE: Neomenia sp., Rhopalomenia carinata, Rhopalomenia rhynchopharyngeata, Dorymenia

parvidentata, Moluscos, Solenogastros, Antártida.

* Marine Biology Station A Graña, University of Santiago de Compostela. Casa do Hórreo, Rúa da Ribeira, 1.

A Graña, E-15590 Ferrol. Spain. E-mail: baoscarOusc.es

** Department of Animal Biology, Faculty of Biology, University of Santiago de Compostela, E-15782 Santiago

de Compostela. Spain. E-mail: bavitucoOusc.es

Á3

Iberus, 21 (1), 2003

INTRODUCTION

During the Spanish campaigns

BENTART'94 and BENTART'95, for the

study of the Antarctic benthos off the

Livingston and Deception Islands (South

Shetland Islands) and in the Bransfield

Strait, north of the Antarctic Peninsula,

17 specimens of Solenogastres Molluscs

were collected. Research had previously

been conducted in this area to gain

knowledge of the benthonic fauna (U.S.

Antarctic Research Program, 1961-1972),

resulting in the description of 23 species

of Solenogastres (SALVINI-PLAWEN, 1978).

Study of the BENTART Collection of

Solenogastres revealed that the majority

of specimens collected belonged to new

species. Dorymenia troncosoi García-

Alvarez, Urgorri and Salvini-Plawen,

1998, 5 specimens: 4 collected south of

Livingston Island on muddy bottoms at

a depth of 65-66 m, and 1 specimen col-

lected north of the same Island on a

muddy bottom at a depth of 240 m

(GARCÍA-ÁLVAREZ, URGORRI AND

SALVINI-PLAWEN, 1998). Dorymenia hes-

peridesi García-Álvarez, Urgorri and

Salvini-Plawen, 2000, 1 specimen col-

lected on a muddy bottom at a depth of

235 m south of Livingston Island; Dory-

menia menchuescribanae García-Álvarez,

Urgorri and Salvini-Plawen, 2000, 6

specimens: 5 collected south of Liv-

ingston Island on Ophidiogorgia paradoxa

Bayar, 1980, at a depth of 50 m, and 1

specimen collected south of the same

Island on a muddy bottom at a depth of

66 m (GARCÍA-ÁLVAREZ, URGORRI AND

SALVINI-PLAWEN, 2000). Ocheyoherpia

bursata García-Álvarez and Urgorri,

2003, 1 specimen collected off Deception

Island on a gravel bottom at a depth of

248 m (GARCÍA-ALVAREZ AND URGORRI,

2003, in press). In this paper, 4 specimens

from the BENTART Collection are

RESULTS

studied. Two specimens were identified

as belonging to two species previously

known in the study area: Rhopalomenia

carinata Salvini-Plawen, 1978 collected

on a muddy and gravely bottom at a

depth of 710 m off Livingston Island and

Rhopalomenia rhynchopharyngeata Salvini-

Plawen, 1978 from a muddy bottom at a

depth of 235 m off Livingston Island.

One specimen belonging to the genus

Neomenia presented features, which well

distinguish it from known species,

although it is not described as a new

species since the anatomic organisation

of the posterior part of the animal is

unknown, this specimen was collected

from a fine muddy and sandy bottom, at

a depth of 80 m off Livingston Island.

And one specimen, collected on a

muddy bottom at a depth of 80 m off

Livingston Island, is described as a the

new species Dorymenia parvidentata.

MATERIAL AND METHODS

The specimens studied were fixed

and preserved in 70% alcohol. The scle-

rites were studied by separation of small

pieces of cuticle from the central dorsal

area of the body and from the ventral

groove. These pieces were treated with

5% sodium hypochlorite for 12 hours in

order to isolate the sclerites; they were

them rinsed with distilled water, dried

under a heater at 40"C and mounted

using synthetic resin. For the anatomical

study, the specimen were decalcified in

an ethylenediaminetetracetic acid

(EDTA) solution 12 hours, embedded in

paraffin and a series of 10 um cross sec-

tions cut which were stained with Azan

of Heidenhain. The anatomy was recon-

structed from the serial sections.

Order NEOMENIAMORPHA Pelseneer, 1906

Family NEOMENIIDAE Ihering, 1876

Genus Neomenia Tullberg, 1875

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

Neomenta sp.

Material examined: One specimen, 3.7 mm in length and 1.4 mm in width at the anterior part, and

0.7 mm in width at the posterior part (sectioned into 10 ym transversal series). Collected at station

7-BOX-3 (62? 44' 17” S; 60* 28' 11” W) with a box-corer trawl on a fine muddy and sandy bottom,

at a depth of 80 m off Livingston Island (South Shetland Islands, Antarctic) during the Spanish

campaign, BENTART'95, for the study of the Antarctic benthos,.

Description: The body of the animal

is wider at the anterior end than at the

posterior end, and gradually narrows in

diameter between the two (Fig. 14). The

sclerites do not appear to protrude from

the cuticle, but they have a very shiny

appearance. The ventral groove is well

visible. In alcohol the colour of the

animal is yellowish white. The cuticle is

30-50 um thick with papillae at the base.

Underneath the epidermis there is a

thick subepithelial layer matrix up to 70

um (Fig. 24). The mantle produces three

types of sclerites: elongated laminate

scales not forming grooves (120 ym x 14

um) (Fig. 1B); solid slightly curved acic-

ular spicules (100um x 4.5 mm) (Fig.

1C); elongated scales in the shape of

grooves (100 um x 16 um) (Fig. 1D). The

pedal pit is located below the oral

opening and the anterior part of the

pharynx (Fig. 1E). In a preserved state, it

has a narrow opening. Its epithelium is

ciliated and at its end the three folds are

visible that continue on to the pedal

groove (Fig. 2B). These folds are like-

wise ciliated and the middle one is

larger than the two lateral ones. At the

end of the body, the two lateral folds

become smaller, and only the middle

fold is still present. In the pallial cavity

10 to 12 respiratory folds are visible, but

due to the poor condition of the animal,

it is impossible to affirm that this is the

correct number. Moreover, the number

of folds may depend on the size and the

maturity of the animal. The buccal

opening is found at the posterior area of

the atrium (Fig. 1E). It represents the

anterior end of a short tube with three

internal thickenings or large lips, two

dorsolateral and one ventral that are in

rostral prolongation of the pharynx

(Figs. 1E, 2A). The two dorsolateral lips

are separated by a mid dorsal slit, but

they are never separated and set off

from the walls where the buccal tube is.

A lateral slit from each dorsolateral lip

separates the ventral lip. These slits also

continue in a ventral space of this lip,

separating the latter as a tongue-like for-

mation from the wall of the buccal tube.

The foregut continues into a long

pharynx having a longitudinally pleated

folded wall, its epithelium is covered by

a fine cuticular layer. The middle

portion of the pharynx is surrounded by

glands and by a strong circular muscu-

lature which cause it to narrow (Figs.

1E, 2C). It opens into the midgut

through a sphincter formed by a very

strong circular musculature (Figs. 1E,

2D). There is no radula or radular sac

and no ventral foregut glandular organs

are elaborated. No rostral caecum of the

midgut is present (Fig. 1E), the latter

shows lateral constrictions due to the

dorsoventral musculature. The cerebral

ganglion is large, located dorsally to the

pharynx in the middle portion above

the ring of circular musculature (Fig.

1E). At either side of the cerebral gan-

glion, there is one lateral ganglion (Fig.

1E). A short distance behind these

lateral ganglion, there are two connec-

tives which emerge from each side of

the cerebral ganglion: the strongest pair

(270 um x 20 um each) runs vertically to

join the buccal ganglion (Figs. 1E, 2C).

The two buccal ganglia are located ven-

trolaterally of the pharynx throughout

the area of strong circular musculature.

They are interconnected by a ventral

commissure of the pharynx. The second

connectives runs to the ventral ganglia

that are large and rounded (100 um in

diameter), joined with each other by a

single commissure (Fig. 2B) and located

above the beginning of the pedal groove

and ventrally in the mid-area of the

pharynx with its strong circular muscu-

lature. The atrium opens at the anterior

Iberus, 21 (1), 2003

end of the body through a narrow slit in

a dorsoventral direction. The atrium is

it presents several papillae forming

groups of 4-7, joined together in one

large and in its dorsal and lateral walls, base (Fig. 2A).

Order CAVIBELONIA Salvini-Plawen, 1978

Family RHOPALOMENIIDAE Salvini-Plawen, 1978

Genus Rhopalomenia Simroth, 1893

Rhopalomenia carinata Salvini-Plawen, 1978

Material examined: One specimen, 7 mm in length by 0.6 mm in width, (sectioned in 10 yum seri-

ated cuts), was collected at station A-30 (62* 01 24” S; 60* 26' 16” W) with a Assasiz trawl drag, on

a muddy bottom at a depth of 710 m, off Livingston Island (South Shetland Islands, Antarctic)

during the Spanish campaign, BENTART'95, to study the Antarctic benthos.

Description: The specimen studied

was slightly flattened, laterally, in the

anterior part, presenting a barely pro-

truding dorsal cuticular keel, although

this is visible in the transversal cuts

(Figs. 3A-C). In alcohol the colour of the

animal is light brown. Cuticle is not

thick, up to 60 ym, attaining 90 ym in

the keel. Sclerites are arranged in

several layers within the cuticle, are

hollow, straight or slightly arched acicu-

lars in varying sizes, up to 100 ym in

length, and are similar to those

observed in the other species of the

genus (see Fig. 151 in SALVINI-PLAWEN,

1978). Blade shaped solid scales, of up to

60 um length, are found in the pedal

groove. A single fold is presented in the

pedal groove (Figs. 3D, E), which does

not access the pallial cavity. The pallial

cavity is small, presenting no respira-

tory folds, lacking abdominal spicules

and copulatory spicules, the anus

leading out dorsally in it, whereas the

spawning duct does so ventrally. In the

specimen studied, no presence of a sen-

sitive dorsoterminal organ was noted.

The mouth (Fig. 3B) opens separated

from the atrium (Fig. 3C) and is located

at the end of a short duct located within

the buccal cavity. The radular sac in the

mid part has a characteristic ventral

epithelial fold (Fig. 3D). It presents a

pair of ventral foregut glandular organs

formed by two long ducts, which lead

into supepithelial glandular follicles

(type A according SALVINI-PLAWEN,

1978). Anteriorly, these ducts are located

parallel to the end part of the radular

sac until leading laterally into the same.

These three ducts (the two glandular

organs and the radular sac) are jointly

surrounded by circular musculature

(Figs. 3D,E). The oesophagus is very

(Right page) Figure 1. Neomenía sp. A: habitus; B: elongate laminate scales; C: acicular spicules; D:

groove-shaped scales; E: Schematic organization of anterior body. At: atrium; Bg: buccal ganglion; -

Bt: buccal tube; Cg: cerebral ganglion; Cm: nervous commissure; Co: connective; Gl: glands; Lg:

lateral ganglion; Li: lip; Ma: mantle; Mg: midgut; Mt: matrix; Mu: musculature; Ph: pharynx; Pp:

pedal pit; Ps: pedal groove folds; Sp: sphincter; Sv: ventral blood sinus; Vg: ventral ganglion. 1-4

lines corresponding to cross-sections A-D in Figure 2. Figure 2. A-D. Microphotographs of the

cross-sections of the anterior region of the body corresponding to lines 1-4 in Figure 1.

(Página derecha) Figura 1. Neomenia sp. A: habitus; B: escamas laminares alargadas; C: espículas aci-

culares; D: escamas excavadas; E: organización esquemática de la parte anterior del cuerpo. At: atrio; Bg:

ganglio bucal; Bt: tubo bucal: Cg: ganglio cerebral; Cm: comisura nerviosa; Co: conectivo; Gl: glándulas;

Lg: ganglio lateral; Li: labio; Ma: manto; Mg: intestino; Mit: matríz; Mu: musculatura; Ph: faringe; Pp:

fosa pedia; Ps: pliegues del surco pedio; Sp: esfínter; Su: seno sanguíneo ventral; Vg: ganglio ventral. 1-4

líneas que corresponden a los cortes en sección A-D en la Figura. Figura 2. A-D. Microfotografías de los

cortes en sección de la región anterior del cuerpo correspondientes a las líneas 1-4 de la Figura 1.

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

E ' Ea pere “54

Iberus, 21 (1), 2003

long and is located dorsally to the

radular sac and to the ventral foregut

glandular organs until leading into the

midgut. The midgut has a large dorso-

rostral caecum, which reaches the level

of the cerebral ganglion (Figs. 3B-D).

The gonad is full of spermatozoids and

ovules, and has a pair of dorsal seminal

receptacles at the spawning ducts.

Remarks: The specimen studied is

from a geographical area (Livingston ls-

land, South Shetland Islands) close to

part of the material studied in the origi-

nal description (Elephant Island /

Joinville Island, South Shetland Islands),

although at a considerably greater depth

710 m, as opposed to 119-220 m (SALVINI-

PLAWEN, 1978). It presents a thinner cuti-

cle than the model material, 60 um in the

present specimen 7 mm in length by 150-

225 ym for specimen up to 35 mm; no

pedal fold was observed within the pal-

lial cavity, nor was there the presence of a

dorsoterminal sensitive organ, as noted

in the original description. But other im-

portant features particular to this species

are well defined: the mouth is located at

the end of a horn and separate from the

atrium, the ventral glandular organs of

the pharynx are subepithelial, and the an-

terior tubular part with the radular sac

are jointly surrounded by circular muscu-

lature; it presents a ventral epithelial fold

in the mid part of the radular sac; and the

general structure of the gonopericardic

system is similar.

Rhopalomenia rhynchopharyngeata Salvini-Plawen, 1978

Material examined: One specimen 13 mm in length and 2.1 mm in width (sectioned into 10 ym

seriated cuts), collected at station A-19 (62%43'43”5; 60%31'27”"W) with an Agassiz drag trawl on a

muddy bottom at a depth of 235 m, off Livingston Island (South Shetland Islands, Antarctic) during

the Spanish campaign, BENTART'95, for the study of the Antarctic benthos.

Description: Rolled up animal with

no keel or protuberances (Fig. 4A),

cylindrical in section, with a bristling

appearance due to the sclerites standing

out from the mantle. In alcohol the

colour of the animal is light brown.

Thick cuticle of up to 150 jm. The scle-

rites are arranged in several layers, both

obliquely and radially on the cuticle,

with different sized hollow, straight or

slightly arched acicules, similar to those

appearing in other species of the genus

(see Figure 151 in SALVINI-PLAWEN,

1978), attaining maximum lengths of

200 um. Blade shaped solid scales are

found in the pedal groove of up to 80

pm in length. In the first third of the

body, the pedal groove presents three

folds, a longer central one of up to 120

pm, and two shorter lateral folds of up

to 70 yum. In the posterior part only one

fold is found, which does not access the

pallial cavity (Figs. 4C-E). The pallial

cavity is small, without respiratory

folds, lacking abdominal spicules and

copulatory spicules (Fig. 4B), and the

anus open dorsally into the cavity. The

. Spawning duct opens independently of

the pallial cavity, since it ends unpaired,

free and directly outside the cavity on

the ventral part (Fig. 4B). There is a dor-

soterminal sensitive organ located at the

posterior end of the body. The mouth

opens separated from the atrium. lt

lacks a radula but presents a short

radular sac. The ventral foregut glandu-

lar organs are formed by two long ducts

into which subepithelial glandular folli-

cles open (Type A according to Salvini-

Plawen, 1978). Anteriorly, these ducts

are located freely to both sides of the

radular sac, until they open laterally

into the radular sac, and are not jointly

surrounded to the radular sac by a

common circular musculature. The

oesophagus is very long, and open into

the ventral part of the midgut. The

midgut has a dorso-rostral caecum,

which extends to the level of the cere-

bral ganglion. The rectum (Figs. 4C,D,E)

is circular in section (diameter up to 150

um). The gonad is full of spermatozoids

and ovules. The pericardium is not very

voluminous and is circular in section

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

Figure 3. Rhopalomenia carinata Salvini-Plawen, 1978. A: habitus; B-E: microphotographs of the

cross-sections of the anterior region of the body. At: atrium; Bo: buccal opening; Cg: cerebral gan-

glion; Dc: dorsal caecum; Fd: ventral epithelial fold of the radular sac; Ke: keel; Lg: lateral ganglion;

Mu: circular musculature; Ph: pharynx; Rs: radular sac; Vfg: ventral foregut glandular organ.

Figure 3. Rhopalomenia carinata Salvini-Plawen, 1978. A: habitus; B-E: microfotografías de los cortes

en sección de la región anterior del cuerpo. At: atrio; Bo: abertura bucal; Cg: ganglio cerebral; Dc: ciego

dorsal; Ed: pliegue epitelial ventral del saco radular; Ke: cresta; Lg: ganglio lateral; Mu: musculatura

circular; Ph: faringe; Rs: saco radular; Vfg: órgano glandular ventral de la faringe.

(diameter of 200-250 pm), the heart is

located dorsally in the pericardium, is

relatively large and perfectly bilobu-

lated (Fig. 4E). There is a pair of seminal

receptacles lying dorsal to the spawning

ducts (Fig. 4P). The spawning ducts

(Figs. 4C-F) are circular in section (up to

400 jm in diameter), and with their

glandular walls, in the posterior part

fuse into a single duct (Fig. 4E), also cir-

cular in section (up to 450 yum in diame-

ter) and with glandular walls.

Remarks: The specimen studied here is

from a geographical area (Livingston

Island, South Shetland Islands) close to

part of the material studied in the origi-

nal description (Elephant Island / Joinville

Island, South Shetland Islands) and at a

similar depth (SALVINI-PLAWEN, 1978).

This specimen is of a larger size (13 mm x

2.1 mm) than those studied in the origi-

nal description (10 mm x 1 mm); the

cuticle is thicker, 150 qm as opposed to

120 um, the same occurring with the

Iberus, 21 (1), 2003

length of the folds in the pedal groove,

where the central groove attains 120 yum

and the lateral grooves 70 um, as opposed

to 85 hm and 60 ym in the model speci-

men. The gonad is not observed as being

divided into two by a septum, as indicated

in the original description. The character-

istics of this species are well defined (see

Table 4, p. 159, in SALVINI-PLAWEN, 1978):

the specimen presents a bristly appear-

ance due to the radially arranged sclerites;

the spawning duct open directly into the

pallial cavity; no radula; the radular sac

is short; and the anterior parts of the

ventral foregut glandular organs are

located laterally to the radular sac, without

circular musculature surrounding the

three ducts.

Family PRONEOMENIIDAE Simroth, 1893

Genus Dorymenia Heath, 1911

Dorymenia parvidentata sp. nov.

Type material: Holotype measuring 7.0 mm in length, 0.6 mm in width (spicule slide, specimen in

seriated sections).

Type locality: Livingston Island (station A-7, BENTART'95) (South Shetland Islands, Antarctic)

62%44'07”S, 60%27'42”W from a silt bottom at 80 m depth.

Deposit and derivatio nominis: The holotype is deposited in the “Museo Nacional de dencia

Naturales” of Madrid, number: MNCN 15.02/12. The specific name refers to the few radular teeth

it has (from the Latin: parvum: small amount, dens: tooth).

Diagnosis: Body 7.0 x 0.6 mm, in

rounded section, with no keel or protu-

berances. Not thick cuticle (50 qm). With

hollow acicular sclerites. Radula with 10-

12 short based teeth with a pointed,

slightly curved apex. Pallial cavity with

walls with no diverticles, extending ante-

riorly into a ventral sac. Unpaired spawn-

ing duct leading into the dorsal wall of the

pallial cavity. A pair of four pointed cop-

ulatory spicules star shaped in section.

Without abdominal spicules. Elongated

erythrocytes with no granulations. With

a dorsoterminal sensitive organ.

Description: Animal with an elon-

gated, cylindrically shaped body, with

no protuberances or keel (Fig. 5A).

Smooth mantle surface, with no project-

ing sclerites and a visible ventral groove.

In alcohol the colour of the animal is

light brown. Cuticle is not thick, measur-

ing about 55 ym, with hollow, slightly

arched acicular sclerites of up to 180 qm

in length (Fig. 5C) arranged in layers.

Along the pedal groove there are two

further types of sclerites: solid, slightly

curved acicular spicules of up to 160 ym,

with one of its two ends pointed and

other rounded (Fig. 5D) and blade

shaped scales of up to 110 yum in length

(Fig. 5E). The pedal groove starts in a

small pedal pit (Fig. 5F) and presents a

single fold (Fig. 6D), which does not ac-

cess the pallial cavity. The pallial cavity

opens onto the exterior by a narrow ven-

tro-posterior opening, and presents no

respiratory folds or diverticles in its

walls (Fig. 6F). It has a pair of copulatory

spicules, in section star-shaped with four

pointes, arranged ventrolaterally on

some small protuberances in the walls of

the pallial cavity (Figs. 5G, 6E). It lacks

abdominal spicules. The anus opens out

into the dorsal wall of the pallial cavity

(Fig. 5G). The pallial cavity presents an

ample sac, which extends ventro-anteri-

orly below the pericardioducts and the

rectum (Figs. 5G, 6E). The atrial sensitive

organ presents several simple papilla on

its walls, and dorsally to this organ lies

the cerebral ganglion (Figs. 5F, 6A), the

only part of the nervous system to be ob-

servable in this specimen. It has a single

dorsoterminal sensitive organ (Fig. 5G).

The mouth opens into the atrial cavity,

occupying a dorsoposterior position,

and is located at the end of duct, which

in the case of the specimen studied, was

found to be evaginated (Figs. 5F 6B). The

pharynx is short and in the midgut pre-

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

Ele 4. Rhopalomenia ROI aaedó Salvini- Pa 1978. E EDI. B- E TITAN

graphs of the cross-sections of the posterior region of the body. Ht: heart; Pc: pallial cavity; Pd:

pericardioduct; Pr: pericardium; Re: rectum; Sd: spawning duct; Sr: seminal receptacle.

Figure 4. Rhopalomenia rhynchopharyngeata Salvini-Plawen, 1978. A: habitus; B-E: microfotogra-

fías de los cortes en sección de la región posterior del cuerpo. Ht: corazón; Pc: cavidad paleal; Pd: peri-

cardioducto; Pr: pericardio; Re: recto; Sd: conducto de desove; Sr: receptáculo seminal.

sents a very short dorso-rostral caecum

(Figs. 5FE, 6B). It has a short radular sac,

and a polystichous /polyseriate radula,

consisting of 10-12 teeth measuring 20-25

um length, with a short base and elon-

gated, slightly curved apex (Fig. 5B).

Teeth are located isolated upon a basal

membrane. The pair of ventral foregut

glandular organs (Figs. 5F, 6C) are ep-

ithelial and tubular shaped (Type C, ac-

cording to SALVINI-PLAWEN, 1978), lead-

ing laterally into the anterior part of the

Iberus, 21 (1), 2003

40 um

200 pm

Figure 5. Dorymenia parvidentata sp. nov. A: habitus; B: radular teeth; C: hollow spicules; D:

groove solid spicules; E: groove scales; F: schematic organization of anterior body; G: Schematic

organization of posterior body. At: atrium; Cg: cerebral ganglion; Cs: copulatory spicule; De:

dorsal caecum; Dso: dorsoterminal sense organ; Ht: heart; Ma: mantle; Mg: midgut; Pc: pallial

cavity; Pd: pericardioduct; Ph: pharynx; Pp: pedal pit; Pr: pericardium; Re: rectum; Rs: radular

sac; Vfg: ventral foregut glandular organ. 1-6 lines corresponding to cross-sections A-F in Figure 6.

Figura 5. Dorymenia parvidentata sp. nov. A: habitus; B: dientes radulares; C: espículas huecas; D:

espícula maciza del surco pedio; E: escamas del surco pedio; F: organización esquemática de la parte

anterior del cuerpo; G: organización esquemática de la parte posterior del cuerpo. At: atrio; Cg: ganglio

cerebral; Cs: espídula copuladora; Dc: ciego dorsal: Dso: órgano sensitivo dorsoterminal; Ht: corazón;

Ma: manto; Mg: intestino; Pc: cavidad paleal; Pd: pericardioducto; Ph: faringe; Pp: fosa pedia; Pr:

pericardio; Re: recto; Rs: saco radular; Vfg: órgano glandular ventral de la faringe. 1-6 lineas que corres-

pondes a los cortes en sección A-F en la Figura 6.

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

AAA

Figure 6. Dorymenia parvidentata sp. nov. A,B,C: microphotographs of the cross-sections of the

anterior region of the body corresponding to lines 1,2,3 in Figure 5. D,E,F: microphotographs of

the cross-sections of the posterior region of the body corresponding to lines 4,5,6 in Figure 3. At:

atrium; Cg: cerebral ganglion; Cs: copulatory spicule; Dc: dorsal caecum; Ht: heart; Mg: midgut;

Pc: pallial cavity; Pd: pericardioduct; Ph: pharynx; Pr: pericardium; Ra: radula; Rs: radular sac;

Vífg: ventral foregut glandular organ.

Figure 6. Dorymenia parvidentata sp. nov. A,B,C: microfotografías de los cortes en sección de la región

anterior del cuerpo correspondientes a las líneas 1,2,3 de la Figura 5. D,E,F: microfotografías de los

cortes en sección de la región posterior del cuerpo correspondientes a las líneas 4,5,6 de la Figura 5. At:

atrio; Cg: ganglio cerebral; Cs: espícula copuladora; Dc: ciego dorsal; Ht: corazón; Mg: intestino; Pc:

cavidad paleal; Pd: pericardioducto; Ph: faringe; Pr: pericardio; Ra: rádula; Rs: saco radular; Vfz:

órgano glandular ventral de la faringe.

radular sac, and are located ventrally un-

der the first third of the midgut. The ery-

throcytes are elongated (15 um length)

and present no granulations. The heart is

clearly bilobulated ventrally (Fig. 6E),

and is located inside a large peri-

cardium. The pair of pericardioducts

lead out from the pericardium on the

posterior-lateral part, and are laocated

ventrolaterally to the digestive duct. The

exemplar was immature, no ovules or

spermatozoids were observed in the go-

nads, the periocardioducts can be ob-

served as narrow ducts until they van-

ish, no spawning duct is present (Figs.

5G, 6E) and no seminal receptacies or the

anterior part of the reproductive appara-

tus Were noted.

(9)

Iberus, 21 (1), 2003

DISCUSSION

Neomenia sp. belongs to the order

Neomeniamorpha, as it presents solid

acicular sclerites together with groove-

shaped scales; there are no ventral glan-

dular organs in the pharynx; with respi-

ratory folds, and it is assigned to the

family Neomeniidae, as it has a rela-

tively thick cuticle with epithelial papil-

lae and subepithelial matrix, solid acicu-

lar sclerites and elongated groove-

shaped scales, it lacks a radula and has a

pedal groove with several folds

(SALVINI-PLAWEN, 1978).

At present two genera of the family

Neomeniidae are known: Neomenia Tull-

berg, 1875 and Heathimenia Salvini-

Plawen, 1967. The specimen is assigned

to the genus Neomenia as it fulfils the

main characteristics of the genus: a

somewhat thick cuticle with papillae;

solid, grooved sclerites; buccal opening

in the atrium; absence of a radula and

ventral glandular organs in the pharynx;

midgut with lateral constrictions and

the presence of respirators folds (WIREN,

1892; SALVINI-PLAWEN, 1978). The status

of the genus Heathimenia with H. verrilli,

collected in the Gulf of Saint Lawrence

on the east coast of North America

(HEATH, 1918) is not settled and the

species needs re-examination (SALVINI-

PLAWEN, 1967, 1978).

Currently six Antarctic or Sub-

Antarctic species are known to belong to

the genus Neomenia. N. permagna Salvini-

Plawen, 1978 is a South Pacific species

collected at a depth of 549 m. The follow-

ing Characteristics distinguish it from

Neomenia sp.: the body size (12 cm x 3

cm); the presence of only groove-shaped

scales and solid acicular sclerites; a pedal

groove having 25 folds; a rostral tube-

like pharynx, without a terminal sphinc-

ter. N. labrosa Salvini-Plawen, 1978 was

collected on the South Shetland Islands

(Elephant Island) at a depth of 220-240

m. It is differentiated from Neomenia sp.

by the following features: the presence of

only groove-shaped scales and solid aci-

cular sclerites; a pedal groove having 13-

1 folds; a pharynx with a lateral slit at

each side in the rostral portion which

separates the dorsal and ventral lips,

without a terminal sphincter. N. trapezi-

formis Salvini-Plawen, 1978 is a South

Pacific species that was collected on the

Antipode Islands at a depth of 2010-2110

m (SALVINI-PLAWEN, 1978).

The main characteristics distinguish-

ing it from Neomenia sp. are: a body with

strong lateral edges and an even

number or dorsal lumps that give it its

shape in the typical trapezoidal section;

the presence of only groove-shaped

scales; arrowhead-shaped sclerites and

solid acicular sclerites; a pedal groove

having 23-3 folds; a pharynx with a

short lateral slit at each side in the

rostral portion, separating the dorsal

and ventral lips; the ventral lip is not

separated from the wall of the buccal

cavity. N. crenagulata Salvini-Plawen,

1978 is a South Indian species, collected

from the Kerguelen Islands at 585 m.

The characteristics that separate it from

Neomenia sp. are: the presence of only

groove-shaped scales and solid acicular

sclerites; a pedal groove having 13-1

folds, a pharynx with a ventral slit in the

rostral portion; the dorsal wall of the

pharynx has pronounced folds that

hang over the buccal space. N. laminata

Salvini-Plawen, 1978 was collected from

the South Orkney Islands at a depth of

298-302 m. The main traits serving to

differentiate it from Neomenia sp. are: a

pedal groove with 7-3 folds; a pharynx

without slits to delimit the lips. N. propi-

etecta Salvini-Plawen, 1978 was collected

from the Ross Sea near Victoria Land at

a depth of 344-351 m. The characteristics

separating it from Neomenia sp. are: the

presence of only groove-shaped scales

and acicular sclerites with keels; a pedal

groove with one fold; a pharynx

without slits to delimit the lips.

Due to the considerable damage of

the posterior body, it was only possible

to study and reconstruct the anterior

part of the single specimen available.

The differences between Neomenia sp.

and other species of this genus are quite

clear, especially owing to the presence of

the three large lips in the rostral portion

of the pharynx, the elongated laminate

mantle scales, as well as its geographical

GARCÍA-ÁLVAREZ AND URGORRI: Solenogastres from the Antarctica, and a new species

location. But as its important posterior

organs are unknown, it is not described

as a new species until it is confirmed

with new data.

Dorymenia parvidentata sp. nov.

belongs to the order Cavibelonia, as it

presents hollow acicular sclerites

ordered into several layers within a rela-

tively thick cuticle, and is classified in

the family Proneomeniidae based on the

fact that it has a polystic/polyseriated

radula and tubular epithelial ventral

foregut glandular organs of type C

(SALVINI-PLAWEN, 1978). The characteris-

tics which locate this new species within

the genus Dorymenia are well defined:

the mouth opens into the atrium,; the

genital orifice is impair, it presents a dor-

soterminal sensitive organ, has copula-

tory spicules and the pallial cavity pre-

sents no respiratory folds.

Taking into account the radular

structure and teeth form (SALVINI-

PLAWEN, 1978: GARCÍA-ÁLVAREZ et al.,

2000), the species of the genus Dory-

menia may be classified into three

general groups: one group presenting

numerous short radular teeth with a

curved apex and long base; another

group with short based teeth and one or

two medium sized teeth; and a third

group, which includes Dorymenia parvi-

dentata sp. nov., characterised by pre-

senting a radula with few, very elon-

gated and short based apex teeth, con-

stituting: Dorymenia acutidentata Salvini-

Plawen, 1978; Dorymenia paucidentata

Salvini-Plawen, 1978 and Dorymenia sin-

gulatidentata Salvini-Plawen, 1978.

BIBLIOGRAPHY

GARCÍA-ÁLVAREZ, O. AND URGORRI, V. 2003.

Ocheyoherpia bursata a new species of Phyl-

lomeniidae (Mollusca, Solenogastres: Ster-

rofustia) from the South Shetland Islands

(Antarctica). Journal of Molluscna Studies (en

prensa)

GARCÍA-ALVAREZ, O., URGORRI, V. AND SALVINI-

PLAWEN, L.v., 1998. Dorymenia troncoso! sp.

nov. (Mollusca Solenogastres: Proneomeni-

idae), a new species from the South Shet-

land Islands (Antarctica). Polar Biol, 20: 382-

387.

With each of the three species cited

above, Dorymenta parvidentata sp. nov.

presents significant differences (see

Table 1 in GARcÍíA-ÁLVAREZ et al., 2000).

Dorymenia parvidentata sp. nov. presents

10-12 radular teeth, whereas D. acutiden-

tata has 22-26, D. paucidentata 12-14 and

D. singulatidentata 14. The pallial cavities

are very different, Dorymenia parviden-

tata sp. nov. presents no diverticles and

extends anteriorly in a ventral sac,

whereas in D. actuidentata it has numer-

ous diverticles and a dorsoanterior sac,

in D. paucidentata lacks diverticles and

presents a pair of lateral sacs and a pair

of ventroanterior sacs and in D. singu- :

latidentata lacks diverticles and presents

a pair of ventroanterior sacs. Also, sec-

tioning of the copulatory spicules in D.

acutidentata and D. singulatidentata

reveals a circular formation and not a

four-pointed star as in Dorymenia parvi-

dentata n. sp. Finally, in D. acutidentata

the pedal fold enters the pallial cavity,

whereas this is not the case in Dorymenia

parvidentata sp. nov.

ACKNOWLEDGEMENTS

We thank Dr. Luitfrid v. Salvini-

Plawen of the University of Vienna for:

his assistance. This work was supported

by the BENTART projets (ANT94-1161-

E, ANT95-1011; ANT97-2097-E) and the

Integrated Spanish-Austrian Co-opera-

tion Actions (HU1997-0002; HU2000-

0010). Thanks also to lan Emmett for the

English translation.

GARCÍA-ÁLVAREZ, O., URGORRI, V. AND SALVINF-

PLAWEÉN, L.v., 2000. Two new species of Dory-

menia (Mollusca: Solenogastres: Pro-

neomeniidae) from the South Shetland

Islands (Antarctica). Journal of the Marine Bio-

logical Association of the United Kingdom, 80:

835-842.

HEATH H 1918. Solenogastres from the Eastern

Coast of North America. Memoirs of the

Museum of Comparative Zoology at Harvard

College, 45 (2): 183-263.

¡0

Iberus, 21 (1), 2003

SALVINI-PLAWEN, L.v., 1967. Kritische

Bermerkungen zum System der Solenogas-

tres. Zeitschrift fúr zoologische Systematik und

Evolutionsforschung, 5 (4): 419-437.

SALVINI-PLAWEN, L.v., 1978. Antarktische und

subantarktische Solenogastres. Eine Mono-

graphie : 1898-1974. Zoologica (Stuttgart), 128 :

1-315.

WIREN. A., 1892. Studien úber Solenogastres

II. Kunglige Svenska Vetenskaps- Akademiens

Handlingar, 25 (6): 1-100.

O Sociedad Española de Malacología —_—_——T— lIberus, 21 (1):57-66, 2003

Cephalopod species collected in the upper continental slope

off Alicante (Western Mediterranean)

Especies de cefalópodos capturadas en el talud superior de Alicante

(Mediterráneo occidental)

Sergi SORIANO”, José L. SÁNCHEZ LIZASO* and Angel GUERRA**

Recibido el 4-VII-2002. Aceptado el 29-X-2002

ABSTRACT

The cephalopod fauna collected in 24 hauls carried out in the upper continental slope off

Alicante (Western Mediterranean) by commercial trawlers is analysed. Samples were

taken at depths between 237 and 611 m from April 1998 to December 1999. Eighteen

species in 8 families were captured. Todarodes sagittatus, Todaropsis eblanae, Rossia

macrosoma, Sepietta oweniana, Eledone cirrhosa and Octopus salutii¡ were the most abun-

dant species. Cephalopods represented 3.9 % of the total catch (94 kg). Average

cephalopod biomass estimated was 89.2 g/hectare. The known depth range for Octopus

defilippi in the Mediterranean is expanded up to 370 m. Observed seasonal changes in

biomass of main species are discussed.

RESUMEN

En el presente trabajo se analizan los cefalópodos capturados por arrastreros comerciales

en 24 lances realizados en el talud superior de la provincia de Alicante (Mediterráneo

occidental). Las muestras fueron tomadas entre los 237 y 611 m de profundidad, entre

abril de 1998 y diciembre de 1999. Se capturaron 18 especies pertenecientes a 8 fami-

lias. Todarodes sagittatus, Todaropsis eblanae, Rossia macrosoma, Sepietta oweniana,

Eledone cirrhosa y Octopus salutii fueron las más abundantes. Los cefalópodos representa-

ron un 3,9 % de la captura total (94 kg.). La biomasa de cefalópodos media estimada fue

de 89,2 gr./hectárea. El rango .de profundidad conocido para Octopus defilippi en el

Mediterráneo fue ampliado hasta los 370 metros. Se discuten los cambios estacionales de

biomasa de las especies más importantes.

KEY WORDS: Cephalopods; Trawl fishery; Seasonality; By-catch; Western Mediterranean.

PALABRAS CLAVE: Cefalópodos, Pesquería de Arrastre, Estacionalidad, Pesca Acompañante, Mediterráneo

occidental.

INTRODUCTION

An important trawling fishery is con- Alicante (Western Mediterranean). The

ducted in the upper continental slope off main target species of this fishery are

* Unidad de Biología Marina, Departamento de Ciencias Ambientales y Recursos Naturales, Universidad de

Alicante, P.O. Box 99, 03080 Alicante, Spain.

** ECOBIOMAR, Instituto de Investigaciones Marinas (CSIC), C/ Eduardo Cabello 6, E-36208 Vigo, Spain.

' Corresponding author. Present adress: Institut de Ciéncies del Mar (CSIC), P. Marítim de la Barceloneta 37-

49, E-08003 Barcelona, Spain.

Iberus, 21 (1), 2003

38.8

38.6

3 siósa

38.4

Ca de l'Horta

38.27 Cap Sta. Pola

100 200 400

-0.6 -0.4 -0.2

ap de la Náo

0 0% 0,4 0.6

Figure 1. Location of samples in the study area. Lines link initial and final sampling positions.

Figura 1. Localización de las muestras en el área de estudio. Las líneas unen las posiciones iniciales y

finales del muestreo.

Norway lobster (Nephrops norvegicus), rose

shrimp (Aristeus antennatus), Plesionika

spp., European hake (Merluccius merluc-

cius), blue whiting (Micromesistius poutas-

sou) and great fork-beard (Phycis

blennoides). Cephalopods are not target

species but contribute to by-catch

(SORIANO, 2000).

Faunistic composition and spatial

distribution of the cephalopods in the

Spanish Mediterranean waters is rela-

tively well known (see GUERRA, 1992 for

a review). However, this information

mainly corresponds to the Catalan Sea

(41 N to 43% N). Thus, the results of

cluster analysis in this area, as well as in

the northern Tyrrhenian Sea, showed

the presence of clear associations related

with depth (SÁNCHEZ, BELCARI AND

SARTOR, 1998). Cephalopod assemblages

and some biological aspects of deep-sea

cephalopods species were studied in the

continental shelf and upper slope off

Balearic Islands (QUETGLAS, ALEMANY

AND SÁNCHEZ, 2000). On the other hand,

different seasonal abundances were

found in the bathymetric distribution of

some species in the Catalan Sea that

could be related to differences in the

spawning and/or recruitment periods

for each species (SÁNCHEZ, 1986;

SÁNCHEZ ET AL., 1998). Depth ramees

recorded for 10 species from the bathyal

basin (1000-2000 m depth) of the North-

Western Mediterranean were analysed

and discussed by VILLANUEVA (1992),

who suggested an up-slope ontogenic

migration of Bathypolypus sponsalis and

Neorossia caroli. Furthermore, the distrib-

ution and abundance of bathyal sepi-

olids in this area has notably increased

as stated by VILLANUEVA (1995).

The results obtained by SARTOR,

BELCARI, CARBONELL, GONZÁLEZ, QUET-

SORIANO ET AL.: Cephalopod of the upper continental slope off Alicante

Table I. Data summary on trawling operations

Tabla I. Resumen de los datos de las operaciones de pesca.

Total Ceph

Min Max Surface catch cotch

Haul Date Initial position Final Position Depth Depth (km2) (kg) (kg)

1 14/04/1998 38-28-645N /0-32-266E 38-11-052N / 0-13-704€ 333 522 0.89% 210.5 3.8

2 10/07/1998 38-18-942N / 0-20-236E 38-10-771N /0-19-096E 433 444 0.285 695 63

3 10/07/1998 38-09-195N / 0-21-305E 38-07-122N / 0-06-325E - 602 602 0.530 70.6 12

4 17/07/1998 38-18-942N /0-20-236E 38-10-771N /0-19-096E 433 444 0.285 417 19

5 17/07/1998 38-09-500N /0-12-384E 38-10-761N /0-17-889€ 426 500 0.183 410 77

6 21/08/1998 38-19-349N / 0-20-432E 38-10-621N /0-19-119E 444 444 0.285 963 12

7 21/08/1998 38-10-693N / 0-17-879E 38-07-177N /0-10-024E 426 426 0.611 268.9 1.0

8 04/09/1998 38-19-830N / 0-20-375E 38-11-671N /0-18:-255E 407 463 0.346 1772 42

9 04/09/1998 38-10-701N /0-17-889E 38-07-524N / 0-07-347E 389 426 0.530 141.7 37

10 16/10/1998 38-02-764N /0-04-881W 38-07-692N / 0-07-625E 611 611 0.407 384 1.1

11 16/10/1998 38-07-328N /0-03-068E 38-11-094N /0-15-578E 351 444 0.469 685 24

12 23/10/1998 38-19-262N / 0-20-377€ 38-10-649N / 0-19-090E 407 433 0.318 576 13

13 23/10/1998 38-14-639N /'0-16-871€ 38-18-838N / 0-20-721E 388 426 0.570 1693 3,5

14 13/11/1998 38-05-303N / 0-00-374€ 38-08-106N /0-11-955E 574 611 0.367 167 0.7

15 13/11/1998 38-09-829N / 0-14-004€ 38-07-143N /0-01-413E 420 426 0.489 921 26

16 07/12/1998 38-10-701N /0-17-889E 38-02-090N /0-11-960W 444 454 0937 892 42

17 12/03/1999 38-21-907N /0-21-092E 38-15-411N /0-12:-516E 237 244 0.367 97.8 6.2

18 12/03/1999 38-16-044N /0-15-958E 38-27-538N / 0-29-082E 366 377 0.652 170.8 7.3

19 18/03/1999 38-19-544N /0-20-219E 38-10-594N / 0-14-243E 366 407 0.407 937 45

20 18/03/1999 38-15-978N /0-16-526E 38-25-409N /0-27-173E 352 370 0.566 1329 4.4

21 23/04/1999 38-06-891N /0-02-817E 38-09-785N /0-18-263E 504 509 0.469 1152 47

22 23/04/1999 38-10-535N / 0-18-540E 38-08-542N / 0-06-104€ 426 444 0.530 859 39

23 04/06/1999 38-25-010N /0-28-575E 38-08-627N /0-04-798E 444 474 0.947 193.1 94

24 27/12/1999 38-19-693N /0-20-497E 38-07-000N / 0-04-545W 426 370 0.926 473.8 6.1

GLAS AND SÁNCHEZ (1998) showed that

few cephalopods are discarded from

trawlers operating in the Western

Mediterranean. Although discarding, of

cephalopod was practically negligible in

terms of biomass in all-bathymetric

strata analysed by the authors, in terms

of number of species the discard compo-

nent was, however, notable.

In Alicante and VALENCIA REGION,

SÁNCHEZ AND OBARTI (1993) studied the

biology and fishery of Octopus vulgaris

and BLANCO, AZNAR AND RAGA (1995)

analysed the cephalopod composition in

the diet of the striped dolphin Stenella

coeruleoalba. Despite this, little is known

about the upper slope continental

cephalopods from the Central Spanish

Mediterranean Sea. Although GUERRA

(1992) reported some species from this

area, the cephalopod fauna of this

region have not received especial atten-

tion. It is therefore important to deter-

mine what cephalopod species capture

the trawling fishery operating off Ali-

cante, which is one of the most impor-

tant fishing areas of the Western

Mediterranean (OLIVER, 1983). This

paper provides a list of the cephalopod

species caught during a project carried

out with the objective of analysing the

discards of the upper continental slope

off Alicante, as well as some information

on the bathymetric distribution and sea-

sonal changes in the main cephalopod

species abundance.

10)

Iberus, 21 (1), 2003

Table IT. List of species collected in this paper after SWEENEY AND ROPER (1998).

Tabla II. Lista de especies obtenidas en el presente estudio según SWEENEY Y ROPER (1998).

Class Cephalopoda Schneider, 1784

Order Sepiida Zittel, 1895

Family Sepiídae Keferstein, 1866

Rhombosepion orbignyana (Férussac, 1826)

Order Sepiolida Grimpe, 1921

Family Sepiolidae Leach, 1817

Subfamily Rossinae Appellúf, 1898

Rossia macrosoma (Delle Chiaje, 1830)

Neorossia caroli (Joubin, 1902)

Subfamily Sepiolinae Appellof, 1898

Sepietta oweniana (Orbigny, 1840)

Order Teuthida Naef, 1916

Family Loliginidae Lesueur, 1821

Loligo media (Linnaeus, 1758)

Family Enoploteuthidae Pfeffer, 1900

Abralia veranyi (Rippell, 1844)

Family Onychoteuthidae Gray, 1849

Onychoteuthis sp.

Family Histioteuthidae Verill, 1881

Histioteuthis bonnellii (Férussac, 1834)

Histioteuthis reversa (Verrill, 1880)

Family Ommastrephidoe Steenstrup, 1857

Subfamily /Mlicinae Posselt, 1891

Illex coindetii (Vérany, 1839)

Subfamily Todarodinae Adam, 1960

Todaropsis eblanae (Ball, 1841)

lodarodes sagittatus (Lamarck, 1798)

Order Octopoda Leach, 1818

Family Octopodidae Orbigny, 1840

Subfamily Octopodinae Grimpe, 1921

Octopus salutii Vérany, 1836

Octopus defilippi Vérany, 1851

Scaeurgus unicirrhus (Delle Chiaje, 1840)

Pteroctopus tetracirrhus (Delle Chiaje, 1830)

Subfamily Eledonidae Grimpe, 1921

Eledone cirrhosa (Lamarck, 1798)

Subfamily Bathypolypodinae Robson, 1928

Bathypolypus sponsalis (Fischer and Fischer, 1892)

MATERIAL AND METHODS

During the April 1998-December 1999

period, twenty four individual samples

were collected on board two commercial

trawlers at a number of sampling locations

at the same fishing ground on the upper

continental slope off Alicante. The studied

area is situated between 38” 13” N//0* 32

E and 37” 54 N//0* 6' W (Fig. 1). Initial

and final tow position, tow duration and

initial and final depths were registered on

board. Depth range varied between 237

and 611 m. Hauls were carried out during

daytime. Mean tow duration was 4.2 h

and towing speed was 2.5 knots. Effective

SORIANO ET AL.: Cephalopod of the upper continental slope off Alicante

Table III. Mean abundances (+ SE) expressed in g/hectare and ocurrences (in %) of the species of

cephalopods caught.

Tabla IT. Abundacias medias (+ ES) expresado en g/hectárea y ocurrencias (en %) de las especies de

cefalópodos capturadas.

SPECIES Abundance (* SE) Ocurrence

Rhombosepion orbignyana 0.40 * 0.34 8.70

Rossia macrosoma 9.18* 2.55 69.57

Neorossia caroli 0.14*%0.14 4.35

Sepietta oweniana 4.00 * 1.24 13.91

Loligo media 0.09 * 0.07 13.04

Abralia veranyi 0.03 * 0.02 17.39

Onychoteuthidae 0.01 * 0.01 4.35

Histioteuthis bonnelli 1.39* 1.39 4.35

Histioteuthis reversa 0.74*0.51 13.04

Illex coindetii 287151 21.74

Todaropsis eblanae 6:63 "231 47.83

Todarodes sagittatus 30.06 * 4.90 86.96

Octopus salutii LED 52.17

Octopus defilippi 0,45% 0.45 4.35

Scaergus unicirrhus 0.80 * 0.80 4.35

Pteroctopus tetracirrhus 1.02 * 0.49 21.74

Eledone cirrhosa 25.19 * 16.37 26.09

Bathypolypus sponsalis 0.26 * 0.20 8.70

gear opening width was 22 m. The surface

sampled during each tow was estimated

using tow duration, towing speed and

gear opening.

Commercial individuals were identi-

fied and weighed on board using a

dynamometer. A random subsample of

the discarded fraction of the total catch in

each haul was taken. These subsamples

were transported fresh to the laboratory.

Cephalopod individuals in these sub-

samples were sorted, identified and

weighed to the nearest 0.1 g and the results

were extrapolated to the total amount of

discards. Then, the individuals were fixed

in 10% formaline and preserved in 70%

alcohol. Finally, the abundance of each

cephalopod species was estimated and

expressed as grams per hectare.

RESULTS

Table I shows a summary of infor-

mation on trawling operations. A total

of 93.7 kg of cephalopods were cap-

tured. Cephalopods represented 3.86%

in weight of the total catch. The mean

biomass of cephalopod estimated was

-89.2 g/hectare.

The cephalopods collected com-

prised 18 species in 8 families (Table ID.

One individual of the family Onycho-

teuthidae was caught but its stage of con-

servation did not allowed precise identi-

fication. Female Argonauta argo empty

shells were caught. However, they have

not been considered due to the lack of

animal and the fact that currents can

drift the empty shells.

Table III shows the mean abundance

and percentage of occurrence of the

species of cephalopod collected in the

present study. Todarodes sagittatus was

the most abundant species followed by

Eledone cirrhosa and Rossia macrosoma.

These three species together constituted

72.75% of the total cephalopods abun-

dance. Octopus salutii, Todaropsis eblanae -

and Sepietta oweniana were very frequent

Iberus, 21 (1), 2003

Table IV. Depth ranges of the collected species off Alicante and in other areas.

Tabla IV. Rango de profundidades de las especies capturadas en Alicante y en otras zonas.

Species Depth range (m) Source Location

Rhombosepion orbignyana 40-460 Sánchez, 1986 Mediterranean

Rossia macrosoma 125-450 Sánchez, 1986 .

Neorossia caroli 150-1744 Sánchez, 1986; Villanueva, 1992 ¿

Sepietta oweniand 22-974 Lumare, 1970; Villanueva, 1992 y

Loligo media 0-200 Sánchez, 1986 '

Abrolio veranyi

Onychoteuthidae

Histioteuthis bonelli 220-430 Sánchez, 1986 p

Histioteuthis reversa 0-1766 Joubin, 1900; Villanueva, 1992 E

Illex coindetii 50-280 Sánchez, 1986 ,

lodaropsis eblanae

lodarodes sagittatus 20-1000 Mangold-Wirz, 1963 ;

Octopus salvtii 30-800 Sánchez, 1986 - ]

Octopus defilippi 6-60 Wirz, 1958 h

Scaeurgus unicirrhus 235-700 Sánchez, 1986 e

Pteroctopus tetracirrhus 110-570 Sánchez, 1986 ;

Eledone cirrhosa 10-570 Sánchez, 1986 :

Bathypolypus sponsalis 120-1835 Mannini, 1989; Villanueva, 1992 h

in the hauls but they were not abundant

in terms of biomass. Finally, 10 of the 18

species were caught in less than 20% of

the hauls and none of them achieved

abundances greater than 2 g/hectare.

The highest cephalopod catches

occurred in summer and winter. S. owe-

niana, T. eblanae and O. salutii showed

their highest abundances in winter,

while E. cirrhosa showed a peak in

summer and T. sagittatus a decreasing

pattern from summer to spring. R.

macrosoma achieved its highest abun-

dance both in summer and winter (Fig.

Table IV shows the depth ranges for

the eighteen species collected in the

present study, together with the known

maximum and minimum depths

recorded for the same species in the

Mediterranean and in other areas.

DISCUSSION

Faunistical composition of the

present study is similar to other studies

carried out in the Western Mediter-

ranean (SÁNCHEZ, 1986; SÁNCHEZ ET AL.,

1998; QUETGLAS ET AL., 2000) and Gulf of

Cádiz (GUERRA, 1982) for the same

bathymetric stratum.

Depth ranges of the species caught

in the sampling were similar to those

shown in SÁNCHEZ (1986), GUERRA

(1992) and QUETGLAS ET AL. (2000).

Despite these similiarities, an individual

of Octopus defilippi was caught in a trawl

between 366 and 377 m depth. This

species is rare in the Iberian peninsula

coasts and its bathymetric distribution is

between 6 and 60 m depth in the

Mediterranean (Guerra, 1992). Never-

theless, it has been caught until 350 m

depth in the Southern Coast of Portugal

(Reis, CABIDO AND LEAL, 1984). Consis-

tently, the bathymetric distribution

range of this species in the Mediter-

ranean should be extended. However, it

was caught only once, so more data will

be needed in order to confirm this fact.

On the other hand, individuals of the

squid Loligo media have been caught

between 237 and 426 m depth, when the

SORIANO ET AL.: Cephalopod of the upper continental slope off Alicante

Table IV. Continuation.

Tabla IV. Continuación

Depth range (m) Source

32-600 Mangold-Wirz, 1963

367-1332 Chun, 1913; Joubin, 1924

29-475 Bas et al., 1976: Guerra, 1982

70-2000 Mangold-Wirz, 1963

0-1332 Lu and Roper, 1979; Joubin, 1924

48-500 Mangold-Wirz, 1963

85- 660 Mangold-Wirz, 1963

0-1000 Clarke, 1966

<350 Reis et al, 1984

70-430 Mangold-Wirz, 1963

70-680 Mangold-Wirz, 1963

10-770 Mangold-Wirz, 1963

358-930 Pérez-Gándaros and Guerra, 1978; Fischer and Joubin, 1906

known bathymetric distribution range

of this species extends between the

surface until 350 m depth (GUERRA,

1992):

Cephalopods represent a commer-

cially important resource in Western

Mediterranean but, deeper than 350 m,

cephalopod catch rates are low and the

majority of species have little or no

commercial value (SARTOR ET AL.,

1998). In the Alicante upper slope

fishery they contributed more to the

discards (4.5%) than to the commercial

catch (3.5%), being fishes and crus-

taceans the most important groups

(SORIANO, 2000). The most abundant

cephalopod species in this study were

the squids T. sagittatus and T. eblanae,

the octopuses E. cirrhosa and O. salutii

and the sepiolids R. macrosoma and S.

oweniana. The relatively high biomass

of octopuses found in the trawl hauls

may be explained by the benthic way

of life of these animals, which

increases the catchability of these

species by the trawl gear. Moreover,

the agreggational behaviour of squids

Location Depth range (m) Source

237-377 Present study

NE Atlantic 237-509 z

NW Atlantic 426-500

NE Atlantic 237-611 3

237-426 ,

352-426 »

366-377 s

? 333-522 ;

NW Atlantic 351-611 :

SW Atlantic 352-611 ;

SW Atlantic 237-509 :

NE Atlantic 351-611 j

231-237 z

NE Atlantic 366-377 :

Caribean Sea 237-244 z

Caribean Sea 352-509 j

- NE Atlantic 366-500

NE Atlantic 400-444 E

(GUERRA, 1992) and sepiolids (VILLA-

NUEVA, 1995) may explain the rela-

tively high abundances of these groups

in the catch.

Many authors studied changes in

size distribution (¡.e. VILLANUEVA, 1992;

QUETGLAS, ALEMANY, CARBONELL,

MERELLA AND SÁNCHEZ, 1998a), abun-

dance (i.e. SÁNCHEZ, 1986; VILLANUEVA,

1995), biomass (i.e. SARTOR ET AL., 1998;

QUETGLAS ET AL., 2000), reproductive

parameters (i.e. QUETGLAS ET AL., 1998a)

and ecological parameters (i.e. QUET-

GLAS ET AL., 2000) along the bathymetric

gradient of cephalopods in the Western

Mediterranean. To our knowledge,

SÁNCHEZ (1986), QUETGLAS, ALEMANY,

CARBONELL, MERELLA AND SÁNCHEZ

(1998b), SÁNCHEZ ET AL. (1998) and

SÁNCHEZ AND MARTÍN (1993) studied

changes in cephalopods abundance

according with season in this area. Nev-

ertheless, they were not focused on

deep-sea species. In this sense, informa-

tion about biomass seasonal patterns of

target and non-target species and the

explanation of these patterns could be

Iberus, 21 (1), 2003

Mean Biomass (g/hectare)

SUMMER

Figure 2. Seasonal abundances (in g/hectare).

Figura 2. Abundancias estacionales (en g/hectárea).

useful to understand the complex eco-

logical relationships in the Western

Mediterranean deep-sea and, in conse-

quence, it may provide data for a better

management of the fishery.

Seasonal changes of biomass

observed in this study may be related

with species life history (spawning

and /or recruitment). R. macrosoma was

caught in summer, autumn and winter,

showing two peaks of abundance

(summer and winter), being spring and

autumn the spawning periods for this

species in Western Mediterranean

(GUERRA, 1992). In the present study no

individuals of R. macrosoma were caught

in spring, which agrees with observa-

tions by VILLANUEVA (1995). In the same

way, S. oweniana spawns from March to

November between 10 and 45 m depth

(GUERRA, 1992), and its highest abun-

dance level was observed in winter. It .

seems reasonable to think that both

AUTUMN

Rossia macrosoma

Sepietta owentana

Todarodes sagittatus

Todaropsis eblanae

Octopus salutii

—E

—O—

—k—

——

—B-

——

Eledone cirrhosa

WINTER SPRING

species may carry out reproductive

migrations to depths out of present

study sampling area.

The relatively high catch rates of T.

sagittatus found in summer and autumn

could be explained arguing that the

spawning period of this species extends

through these seasons, as observed in

the Balearic Sea (QUETGLAS ET AL.,

1998a). On the other hand, the high

catch rate could be also due to the fact

that individuals are more concentrated

during this period and, therefore, its

catchability by the trawl gear increases.

The decreasing trend in biomass from

summer to spring could be related with

changes in the intensity of spawning

during the year, linked with a process of

desaggregation after the spawning and

due to a gradual ontogenic migration to

deeper waters than local trawl fleet

operation depths. In this sense, could be

made a hypothesis about the existence

SORIANO ET AL.: Cephalopod of the upper continental slope off Alicante

of a variable spawning depth according

to season, in which, individuals would

spawn in shallow waters in summer

while it would perform an increasing in

the spawning depth until winter, when

spawning period finishes for this

species.

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ACKNOWLEDGEMENTS

Authors wish to thank the collabora-

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helpful comments on the manuscript.

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O Sociedad Española de Malacología —__—_——— lberus, 21 (0% 57-97, 2003

On the phylogenetic significance of the aplacophoran

Mollusca

Sobre la significación filogenética de los moluscos aplacoforos

Luitfried von SALVINTEPLAWEN*

Recibido el 9-1X-2002. Aceptado el 13-XII-2002

ABSTRACT

The increase in our knowledge of the organisation of the aplacophoran Mollusca over the

last decades is outlined and the arguments with respect to their systematic position and

phylogenetic significance are discussed.

First, the analysis of the mantle cavity in Neomeniomorpha and Chaetodermomorpha

reveals an independent"evolutionary transformation of both: in neomeniomorphs by a lat-

eral narrowing of the mucociliary-gliding body with internalisation of the (posterior) mantle

grooves, in chaetodermomorphs by a terminal shift and inversion of the (posterior) mantle

grooves and the reduction of the foot to burrow in sediment, with midventral fusion of the

mantle rims from posterior to anterior (HOFFMAN, 1949). This confirms their diphyletic status

derived from a common ancestral group; together with the absence of true synapomorphies,

this allows the chaetodermomorphs to be separated as a proper clade Caudofoveata

(Boettger, 1956) in contrast to the neomeniomorph aplacophorans or Solenogastres.

Second, the comparison of the alimentary tract and the excretory system in both classes

with those in the monophyletic Testaria (Placophora + Conchifera) contradicts an origin of

the aplacophoran Mollusca by regressive derivation from Placophora (as assumed by

Pelseneer, 1890); rather, it confirms the conservative level of the aplacophoran organisa-

tion, which is also strongly supported by cladistic analysis.

Third, the mantle cover, the musculature, and the elaboration of the alimentary tract reflect

a gradual anagenesis within Mollusca from the aplacophoran to the polyplacophoran and

mono-placophoran (conchiferan) levels.

Fourth, within the Mollusca the Solenogastres have retained a primitive stage of the radula

which, in its most conservative level, appears to be characterized by the monoserial type

(median bars or two teeth with symphysis). In correlation with the cnidaria-vory in

Solenogastres, a molluscan origin from ciliary-gliding, mesenchymate Spiralia is accepted,

and the plesiomorphic conditions of molluscan characters are outlined.

RESUMEN

Se pone de manifiesto el mayor conocimiento sobre la organización de los moluscos

aplacóforos en las últimas décadas y se discuten su posición sistemática y relevancia filo-

genética. |

El análisis de la cavidad del manto en Neomeniomorpha y Chaetodermomorpha revela

una transformación evolutiva independiente en ambos grupos: en el primero por un adel-

gazamiento lateral del cuerpo mucociliar con internalización de los pliegues posteriores

* Institut fúr Zoologie, Universitát Wien, A-1090 WIEN, Althanstrafie 14.

Iberus, 21 (1), 2003

- del manto, en los Chaetodermomorpha por un cambio terminal e inversión de los pliegues

posteriores del manto y la reducción del pie para enterrarse en el sedimento, con una

fusión ventral de los bordes del manto de atrás adelante (HorrMAN, 1949). Esto confirma

su estatus difilético a partir de un ancestro común. Junto con la ausencia de verdaderas

sinatopomorfías, esto permite separar a los Chaetodermomorpha como verdaderos

Caudofoveata (Boettger, 1956) en contraste con los Solenogastres.

La comparación del tracto alimentario y el sistema excretor en ambas clases con las de

los Testaria (Placophora y Conchifera) contradice un posible origen de los aplacóforos por

derivación regresiva de los Plocophora [tal y como sugirio Pelseneer, 1890); esto con-

firma el nivel conservativo de la organización aplacófora, lo cual se sostiene también por

análisis cladístico.

La cubierta del manto, la musculatura y la elaboración del tracto alimentario refleja una

anagénesis gradual dentro de los moluscos, desde los aplacóforos a los poliplacóforos y

los conchífera.

Dentro de los moluscos, los solenogastros han conservado un estadío primitivo de rádula

que, en sus niveles más conservativos, es del tipo monoserial (piezas medias o dos

dientes con sínfisis). En correlación con los solenogastros comedores de cnidarios, se

acepta un origen a partir de Spiralia mesenquimáticos con desplazamiento ciliar, y se

indican las condiciones plesiomórficas de los caracteres de moluscos.

KEY WORDS: *Aplacophora*, Solenogastres, Caudofoveata, molluscan phylogeny.

PALABRAS CLAVE: *Aplacophora*, Solenogastres, Caudofoveata, filogenia de moluscos.

INTRODUCTION

The purely marine aplacophoran

molluscs are classified in two taxa, the

chaetodermomorphs or Caudofoveata

and the neomeniomorphs or Solenogas-

tres (Fig. 1), known since 1845 (Chaeto-

derma: Lovén) and 1875 (Neomenia: Tull-

berg), respectively. A first period of

enlargement in knowledge began in

1877 and stretched to 1921, with the

description of many new species of both

chaetodermatids and neomeniids

(mainly Koren and Danielssen,

Hubrecht, Kowalevsky and Marion,

Pruvot, Wirén, Thiele, Nierstrasz,

Heath, Odhner; see SIMROTH, 1893b,

HOFFMANN, 1929). A second period may

be outlined from1938-1950, with investi-

gations by Baba, Stork, Leloup,

Hoffman and Schwabl (see FISCHER-

PIETTE AND FRANC, 1960, HYMAN, 1967);

a third period began in 1967 involving

L. v. Salvini-Plawen and since 1972 like-

wise A. Scheltema. Overviews and com-

pilations of organisation are given by

SALVINI-PLAWEN (1971, 1972a, 1985) and

SCHELTEMA, TSCHERKASSKY AND

KUZIRIAN (1994). Most of these investi-

gations concern organisational diversity

based on descriptions of species, but

comprehensive collections still await

elaboration. Due to the habitat of

members of both the chaetodermo-

morphs/Caudofoveata and the neome-

niomorphs /Solenogastres (mostly

below 40 meters depth) living observa-

tions, biological data and physiological

investigations are still rare. However,

the work over the last decades has con-

tributed considerably to elucidating the

systematic position of the apla-

cophorans and to underlining their phy-

logenetic significance.

Scientific results in the phylogenetic

context suffer from two properties: they

are often evaluated one-sidedly in rela-

tion to a predominant or central group

(for molluscs: gastropods or conchifer-

ans in general), and there is an inflexible

trend to press them into some beloved

theory (e.g. molluscs as altered coelo-

mates or even articulates) rather than to

adapt or reject hypotheses based on new

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

TASAS GOTO

ros

Caudofoveata

Figure 1. Aplacophoran Mollusca. External aspect of an animal (Solenogastres: Nematomenia

banyulensis, Caudofoveata: Falcidens sagittiferus), and organisational sketch of alimentary tract,

gono-pericardial system and mantle cavity. cg: cerebral ganglion; go: gonad; md: glandular duct;

mg: midgut; mgs: unpaired midgut sack; mt: spawning duct (internalised mucous tract of mantle

groove); pc: pericardium; pg: pedal gland; ps: pedal shield; so: dorsoterminal or osphradial sense

organ.

Figura 1. Moluscos Aplacóforos. Aspecto exterior de un animal (Solenogastres: Nematomenia banyulen-

sis; Caudofoveata: Falcidens sagittiferus), y esquema de la organización del tracto digestivo, del sistema

gono-pericárdico y de la cavidad paleal. cg: ganglio cerebral; go: gónada; md: conducto glandular; mg:

intestino medio; mgs: saco impar del intestino medio; mt: conducto del desove (tracto mucoso internal-

izado del surco paleal); pc: pericardio; pg: glándula pedal; ps: escudo pedal; so: órgano sensorial dorsoter-

minal u osfradial.

facts and results. The aplacophoran

molluscs, as two small, conservative

taxa, additionally suffer from the great

effort and expense to collect them. Our

standard of knowledge is therefore

weak compared with other molluscan

groups. They are thus often not seri-

ously considered in comparative evalua-

tions, be it due to their low number, be it

that the “a-placophoran” condition sug-

gests a regressive state, or be it that they

are regarded as aberrant inconveniences

that are not equivalent to “true” mol-

luscs.

1. THE DIPHYLY/PARAPHYLY OF

*APLACOPHORA *

The two aplacophoran taxa exhibit a

dorsal integument (mantle) with calcare-

ous (aragonitic) products, a mantle cavity

with body outlets, a radula, a tetraneu-

rous nervous system, and a gono-peri-

cardium including a heart that performs

open circulation and ultrafiltration. In spite

of their unfamiliar appearance they are

thus true Mollusca. Additional characters

in each of the two groups unequivocally

confirm them as members of the phylum.

Iberus, 21 (1), 2003

When IHERING (1876) created (within

Amphineura) the Aplacophora in con-

trast to Placophora, he emphasized the

different organisation of Chaetodermati-

dae and Neomeniidae from chitons.

GEGENBAUR (1878: 139) proposed for the

group which has “a more distinct sepa-

ration of the ventral side through the

differentiation of a furrow”, the name

Solenogastres; unfortunately and mis-

leadingly he also included Chaetoderma

based on the erroneous description by

GRAFF (1877) of the dorsoterminal sense

organ as the vestige of a ventral furrow.

As early as 1883, Lankester classified

Neomeniae and Chaetoderma as

equivalent orders to Polyplacophora

within his Isopleura. In Lankester's

Treatise, PELSENEER (1906) again

grouped his orders Neomeniomorpha

and Chaetodermomorpha below Apla-

cophora, a classification that has been

retained by many authors up to present.

Decisive insights, however, were pre-

sented by HOFFMAN (1949). His detailed

histological, comparative investigation

on the mantle cavities of Chaetoderma

nitidulum Loven, 1845, Proneomenia

“antarctica” (= Dorymenia hoffmani S.-

Plawen, 1978) and Leptochiton asellus

(Gmelin, 1791) revealed the diphyletic

status of recent aplacophorans (Fig. 2).

The posterior mantle grooves in lower

representatives of the (secondarily much

flattened) Placophora are provided with

mucous tracts as well as with the outlets

of gonoducts and excretory organs (Fig.

2A); these tracts parallel in extension at

least the merobranchiate elaboration of

ctenidia. This paired section of the

mantle cavity in the ctenidia-less

neomeniomorphs /Solenogastres is rep-

resented by the glandular spawning

ducts (also known as shell glands or

“lower gonoducts”). Consequently, as in

other molluscs, the mesodermal pericar-

dioducts open directly into the ectoder-

mal mantle cavity, i.e. here into the

internalised mucous tracts or spawning

ducts (Fig. 2B). This condition is clearly

evidenced also in juvenile specimens, in

which the anlagen of the pericardio-

ducts come from the mesodermal peri-

cardium and the anlagen of the spawn-

ing ducts emerge from the ectodermal

mantle cavity rudiment (cf. e.g.

BABA,1938 for Epimenia babai S.-Plawen,

1997, SALVINI-PLAWEN, 1978 for Neome-

nia laminata S.-Plawen, 1978, or TopT

AND SALVINI-PLAWEN, 2003 for Spiomenia

prematura).

In contrast, in chaetodermo-

morphs/Caudofoveata the mucous

tracts of the posterior mantle grooves as

well as the ctenidia (one pair) are

retained as free structures; here,

however, they are arranged upside

down ventrally in the mantle cavity

(HOFFMAN, 1949). In addition, the peri-

cardioducts are not directly connected

to the (mucous tracts of the) mantle

cavity; rather, they each lead out into a

pair of voluminous glandular organs of

unknown homology (see below); only

these glandular ducts or sacks open lat-

eroventrally into the mucous tracts (Fig.

2C). Both the arrangement of the

mucous tracts (present in all females,

rarely in males) and the position of

those openings demonstrate an inver-

sion of the organs (HOFFMAN, 1949).

Thus, the mantle cavity configurations

of neomeniomorphs and chaetodermo-

morphs are clearly different from each

other. In neomeniomorphs they reflect

an evolutive slendering of their forerun-

ners: the ciliary-gliding body narrows,

with the foot becoming a mere pedal

groove, and the (posterior) mantle

grooves internalise to become the

present spawning ducts. In contrast, the

precursors of the chaetodermomorphs

— as sediment burrowers — underwent

a terminal shift of the (posterior) mantle

grooves and a ventral fusion of the

mantle rims into a tube (cf.

Scaphopoda); this included an inversion

of the mucous tracts and ctenidia. This

development was paralleled by the

reduction of the foot with midventral

fusion of the mantle rims from posterior

to anterior (see the midventral anterior

mantle suture in some Scutopus species).

The result was a cylindrical, worm-like

habit.

These scenarios clearly reflect two

totally different evolutionary processes:

neither can the chaetodermophorph

SALVINI-PLAWEN: Significance of-aplacophoran Mollusca

GOA OOOO AAN

eo reste MU. o...

Figure 2. Relation of pericardium, pericardioducts and mantle cavity in A Placophora (with cteni-

día), B Solenogastres and C female Caudofoveata (with ctenidia). At left schematic views from

lateral, at right schematic views projected from behind according to oblique cross sections indi-

cated at left. go: gonad; md: glandular duct; ml: musculus longitudinalis; mt: mucous tract of

mantle groove; pc: pericardium; so: osphradial sense organ.

Figura 2. Relación del pericardio, de los pericardioductos y de la cavidad paleal en A Placophora (con

ctenidios), B Solenogastres y C Caudofoveata femeninos (con ctenidios). A la izquierda vistas esquemati-

cas de lateral, a la derecha vistas esquematicas por atrás según las secciones transversales indicadas. go:

gónada; md: conducto glandular; ml: musculus longitudinalis; mt: tracto mucoso del surco paleal; pc:

pericardio; so: órgano sensorio osfradial.

Iberus, 21 (1), 2003

Figure 3. Structures of ventral body adjacent to mantle rim in A Caudofoveata (region of pedal

shield), B Solenogastres (region of spawning ducts / mucous tracts), C Placophora (region of

mucous tracts) (after SALVINI-PLAWEN, 1991). ce: microvillious epithelium of mantle groove; cu:

irregularly-arranged microvilli within matrix of glycocalix; ml: musculus longitudinalis; mr: mantle

rim (inner mantle fold); mt: mucous tract of mantle groove; ps1: pedal shield (cerebrally innerva-

ted); ps2: pedal sole (ventrally innervated); sg: glands of pedal shield or sole.

Figura 3. Estructuras del cuerpo ventral contiguo al margen del manto A en Caudofoveata (i región del

escudo pedal), B en Solenogastres (región de los tractos mucosos / conductos de desove), C en Placophora

(región de los tractos mucosos) (según SALVINI-PLAWEN, 1991). ce: epitelio microvilloso del surco paleal;

cu: estrato de glycocalix con microvilli irregulares ; ml: musculus longitudinalis; mr: margen del manto

(pliegue interor del manto); mt: tracto mucoso del surco paleal; psl: escudo pedal (de inervación cere-

bral); ps2: suela pedal (de inervación ventral); sg: glándulas del escudo pedal o de la suela pedal.

configuration be derived from the

neomeniomorph conditions, nor vice

versa (see Fig. 2). Both arrangements are

comprehensible only when each derived

from ancestors whose posterior body

bore a peripedal mantle groove with

mucous tracts and the outlets of the

pericardioducts. Such a relation of the

two aplacophoran groups clearly

reflects a diphyly (or, including the

more flattened Placophora, a triphyly).

Thus, BOETTGER (1956) separated the

chaetodermomorphs as Caudofoveata

from other, neomeniomorph apla-

cophorans with a pedal groove: Ventro-

plicida (Boettger, 1956) = Solenogastres

(Gegenbaur, 1878, emend. SIMROTH,

1893a). SALVINI-PLAWEN (1967) finally

accepted both as independent classes

equivalent to Placophora. Are there

synapomorphies of one of these classes

with the Placophora (or other molluscan

group)? Up to the present no conclusion

can be drawn about whether — within

paraphyletic aplacophorans -— the

Caudofoveata or the Solenogastres are,

from the cladistic point of view, the

sister-group of Placophora. Referring to

the Solenogastres, we may assume that

their precursors were likewise provided

with anterio-lateral mantle grooves that

underwent regression during evolutive

transformation (HOFFMAN, 1949). This is

supported by the presence of a pedal

gland, which suggests a former separa-

tion of the cerebrally innervated peri-

oral region (snout, “head”-portion) from

the ventrally innervated foot with pedal

gland. This would equalise the atrial

sense organ or vestibulum of Solenogas-

tres with the pre-oral mantle groove

(HOFFMAN, 1949). On the other hand, it

is not clear whether such precursors

were already provided with ctenidia

(see below). In contrast, at present we

cannot advance similar assumptions

about the regression of mantle grooves

in Caudofoveata. They show no vestige

of a pedal gland, and consequently we

do not know whether the mantle

grooves of their flattened ancestors also

extended into the anterior body. The

post- to peri-oral, secondarily cuticu-

larised buccal plate or pedal shield

reflects a histology and configuration

similar to that of the pedal groove in

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

A o

dl, O SL nsl

cs do A

cg nsl

SS víg

y 0 UR

ES Y

P8 Si

Figure 4. Innervation areas in the ventral anterior body (with identical marking) of A Caudo-

foveata, B Solenogastres, C Placophora. as: atrial sense organ; cg: cerebral ganglion; eg: esophageal

gland; fs: foot (pedal sole); hs: head plate; mc: anterior mantle groove; mg: midgut; nsl: lateral

nerve cord; nsv: ventral nerve cord; oe: esophagus; pf: peri-oral fold (snout); pg: pedal gland; ps:

pedal shield; vfg: ventral foregut gland.

Figura 4. Áreas de inervación del cuerpo anterior ventral (marcadas identicamente) en A Caudofoveata,

B Solenogastres, C Placophora. as: órgano sensitivo atrial; cg: ganglio cerebral; eg: glándula esofágica; fs:

pie (suela pedal); hs: cabeza; mc: surco paleal anterior; mg: intestino medio; nsl: cordón nervioso lateral

; nsv: cordón nervioso ventral; ve: esófago; pf: hocico; pg: glándula pedal; ps: escudo pedal; vfz: glándula

ventral de la faringe.

Iberus, 21 (1), 2003

Solenogastres and the foot in Pla-

cophora (HOFFMAN, 1949, SALVINI-

PLAWEN, 1972a; Fig. 3); the fine struc-

tural organisation (SALVINI-PLAWEN,

1991, SCHELTEMA ET AL., 1994) neither

supports nor contradicts the suggestion

that the shield might be derived from an

anterior portion of the ancestral foot.

Yet, the pedal shield is cerebrally inner-

vated (SALVINI-PLAWEN, 1972a; Fig. 4),

and it could therefore be interpreted to

represent the remnant of a formerly

more extended (ventrally and cerebrally

innervated) ventral locomotory organ

(SALVINI-PLAWEN, 1972a, 1981a); the

mantle grooves in such case were

restricted to the posterio-lateral body.

Despite these as yet unsettled condi-

tions regarding the configuration of the

common ancestral anterior body (no

organogenesis known in Caudofoveata),

the different organisation of the posterior

body in the two groups (mantle cavity;

Fig. 2) unequivocally demonstrates the

diphyly of Solenogastres and Caudo-

foveata. That diphyly is also strongly evi-

denced by cladistic analysis, which either

proposes a paraphyly of aplacophorans

with (dependent on character choice and

coding) the Solenogastres or the Caudo-

foveata as first offshoot (SALVINI-PLAWEN

AND STEINER, 1996, HASZPRUNAR, 2000),

or results in a compromise-polytomy (tri-

phyly) of Solenogastres, Caudofoveata

and Placophora (which in strict Henni-

gian cladistics always includes a not yet

solved paraphyly). To date there are no

comparative data from molecular se-

quences to give support to one or the

other side: The available 185 rDNA se-

quence of Scutopus (Caudofoveata) clus-

ters inmidst of the Mollusca (WINNEPEN-

NINCKS, BACKELJAU AND DE WACHTER,

1996); attempts at sequencing the 185

rDNA gene of threee different species of

Solenogastres merely identified gut con-

tents as diverse Cnidaria (Dr. Hermann

Dreyer, Wien, unpublished), a result also

obtained by other investigators (pers.

comm.; see also WINNEPENNINCKS ET AL.,

1996); probably only DNA sequences of

non-feeding larvae might provide valu-

able information. Polytomy is thus

presently preferred, being supported by

the manifold differences in the organisa-

tion between the three groups (e.g.

SALVINI-PLAWEN, 1972a, 1985; see Table D.

Diphyly of recent aplacophoran molluscs

is even strikingly evident in “mono-

phyletic” representations (SCHELTEMA,

1978, SCHELTEMA ET AL., 1994), with their

separate treatment of organ systems of

each group. Finally, besides the sexual

condition (hermaphroditism in Soleno-

gastres), the single preoral ciliary trochus

in larvae of Solenogastres also contrasts

to the three preoral trochi in Scutopus and

Chaetoderma (Caudofoveata; SALVINI-

PLAWEN, 1991, NIELSEN, 1995) as well as

in other groups (Okusu, 2002).

Paraphyly of *Aplacophorans* with

diphyly of recent groups is in accor-

dance with the (prerequisite) lack of true

synapomorphies (in contrast to SCHEL-

TEMA, 1996, Ivanov, 1996). The mantle

cover of cuticle and unicellularly

formed aragonitic sclerites, likewise

elaborated at the girdle/perinotum of

Placophora (HOFFMAN, 1949, HAAS,

1981), represents a plesiomorphic char-

acter, as does the lack of excretory

organs (emunctoria) for the secondary

urine in aplacophoran organisation (see

below). The so-called ganglionated

nervous system is only present in the

Solenogastres: They mostly possess

medullary ventral and lateral body

cords combined with more or less dis-

tinctly outlined, fairly serial ganglia; all

stages between a predominantly

medullary condition and a true separa-

tion of carya into ganglia interconnected

by pure fibrous portions have been

observed. In Caudofoveata, however,

the medullary system shows only a few

ganglia (buccal ganglia, large ventral

ganglion at the beginning ventral cords,

precerebral ganglia; SALVINI-PLAWEN,

1972a); this reflects a similar condition

in Placophora (buccal and subradular

ganglia). Thus, in contrast to Solenogas-

tres, no ganglionisation of body cords

can be stated in Caudofoveata. The orig-

inally paired osphradial sense organ is

also clearly a molluscan plesiomorphy

(SALVINI-PLAWEN, 1981la, HASZPRUNAR,

1987). The position of the latter in

Caudofoveata is at the dorsal border of

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

Table 1. Comparison of characters in Solenogastres, Caudofoveata and Placophora.

Tabla I. Comparación entre los caracteres de Solenogastres, Caudofoveata y Placophora.

with turgescent cells

Ventral foregut glandular organs paired follicles

paired follicles

Character Solenogastres Coudofoveata Placophora

Chitinous cuticle present present present

Montle sclerites scales, spicules scales scales at girdle

Foot groove with folds reduced large foot

Pedal gland present no gland present in lorvae

Cerebral portion mouth /snout+ atrium mouth + buccal pedal shield head disc+anterior groove

Osphradial sense organ at dorsoterminal mantle area atdorsoterminal mantle border within mantle cavity

Ctenidia no ctenidia one pair 5-88 pairs

Mucous tracts of mantle cavity internalised free, upside down free, at roof

“Lower gonoducts” of literature part of mantle cavity “glandular ducts” distal portion of

(spawning ducts/ between pericario- proper gonoducts

mucous tracts) ducts and mantle cavity

Excretory organ (emunctoria) no emunctoria function assumed by “glandular ducts”? central portion

of pericardioducts

- Outlet of gonads gono-pericardioducts gono-pericardioducts gonoducts

Body wall musculature present, only below mantle — present, ventrally fused below mantle vestigial, only below mantle

M. longitudinalis present vestigially present present (?)

Bundles of dorso-ventral musculature — serially arranged serial, but vestigial 16 poirs

Nervous sytem ganglionated medullary medullary

Latero-terminal commissure supro-ectal supro-ectal supro-ectal

-Subradular organ not present not present present

Radula membrane no separate pre-ibbon separate pre-ibbon separate ribbon

Radulo monoserialdistichous, distichous polystichous in paired anlage

polystichous-polyserial

Radula support very simple to complex two compact bolsters two vesicles with cartilages

vestigial pouches

to elaborate organs of-subradular sack

Anterior midgut body-filling body filling narrow esophagus

+ stomach + glands

Posterior midgut body-filling midgut duct+ midgut sack narrowed to looped intestine

Heart (ventricle) invagination of pericardium in part invagination of pericardium invagination of pericardium

Aorta no aorta present present

Sex hermaphroditism separate sexes mostly separate sexes

Sperm derived introsperms modified aquasperms modified aquasperms

Larval type pericalymma, stenocalymma stenocalymmo pseudo-trochophora

Preoral trochi of larvae one three one or wo

Locomotion muco-ciliary gliding burrowing muco-ciliary gliding

Nutrition carnivory micro-omnivory predominantly herbivory

the mantle cavity and in Solenogastres

dorsally outside the mantle cavity (in

Neomenia labrosa S.-Plawen, 1978, the

position is likewise outside the retracted

mantle cavity: misinterpretation by

IVANOV, 1996). This dorsoterminal loca-

tion is different in detail and is thus too

ambivalent to serve as a good synapo-

morphy. This also holds true for the

gonopericardioducts (paired intercon-

nection between gonads and peri-

cardium) in both aplacophoran groups.

Iberus, 21 (1), 2003

mo sro ve

Figure 5. Regionated alimentary tract of Testaria (Placophora + Conchifera) (after MIZZARO-

WIMMER AND SALVINI-PLAWEN, 2001). (cr): crop (Conchifera only); dfg: salivary gland(s); eg:

esophageal gland; ep: esophageal pouch; in: intestine; (j): jaw (Conchifera only); mg: midgut

gland; mo: mouth opening; oe: esophagus; ph: pharynx; pp: pharyngeal pouch; ras: radula sheath;

re: rectum; sro: subradular organ; (ss): style sack (Conchifera only); st: stomach; vfg: ventral

foregut gland(s).

Figura 5. Tracto digestivo subdividido de los Testaria, (según MIZZARO-WIMMER Y SALVINI-PLAWEN,

2001). (cr): papo (solo Conchifera); dfg: glándula(s) salivar(es); eg: glándula esofágica; ep: bolsa esofá-

gica; in: intestino delgado; (¡): mandíbula (solo Conchifera); mg: glándula del intestino medio; mo:

boca; oe: esófago; ph: faringe; pp: bolsa faríngea; ras: bolsa de odontoblastos; re: recto; sro: órgano subra-

dular; (55): saco del estilo (sólo Conchifera); st: estomago; vfe: glándula(s) ventral(es) de la faringe.

This condition either is a retained ple-

siomorphy of pre-testarian configura-

tion, or it represents a paedomorphy (as

convergent apomorphy or synapomor-

phy?). Based on the different evolutive

transformation of the posterior body

(see above), it may also reflect true

autapomorphies in both Caudofoveata

and Solenogastres (see below, chapt. 6).

2. MIDGUT AND EMUNCTORIA

A comprehensive analysis of the

digestive systems in Mollusca (SALVINI-

PLAWEN, 1981b, 1988a) demonstrates

two principal configurations of the

postradular alimentary tract: either a

simple, basically body-filling midgut

(*Aplacophora*; Fig 1), or a midgut

axially regionated into oesophagus with

paired glandular pouch, stomach with

paired digestive gland, and a narrowed,

wound to enrolled intestine (Testarla;

Fig. 5). The detailed coincidence of the

organ system between Placophora and

conservative Conchifera also includes

the subradular organ, the radular

bolster, the anterior oesophagus with

dorsal food channel and the simple pos-

terior esophagus (SALVINI-PLAWEN

1988a). All these clearly synapomorphic

elaborations in Placophora and

Conchifera define the monophyletic Tes-

taria (SALVINI-PLAWEN, 1972a, 1985,

1991; WINGSTRAND, 1985).

In contrast, the

possess a through-going, uniformly

body-filling midgut (middorsally con-

fined by the gonads) without regiona-

tión.The Caudofoveata show a basically

similar midgut, yet with a posterior

splitting into midgut sack and midgut

duct (“intestine”). This latter exit canal

is an asymmetrically dorsal-right, longi-

tudinal subdivision of the posterior

midgut, leaving the main portion as a

single, voluminous blind sack (Fig. 1).

As demonstrated elsewhere (SALVINI-

PLAWEN, 1981b, 1988a), this longitudinal

splitting is evidenced by the most con-

servatively represented configuration in

some members of the Limifossoridae

le.g. Psilodens elongatus (S.-Plawen,

1972)1: in these the undivided (anterior)

midgut and the midgut sack still exhibit

an identical histological elaboration,

thus still forming a histological entity.

Terminally, the midgut duct continues

in the ventrally inclined hindgut to open

at the bottom of the mantle cavity. Thus,

in contrast to the Testaria (Fig. 5), in

which the midgut glands principally are

a paired lateral pouching of the stomach

Solenogastres --.

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

Figure 6. Gono-pericardial system of Testaria (Placophora + Conchifera) (after MIZZARO-

WIMMER AND SALVINI-PLAWEN, 2001). ao: aorta; aob: bulb of aorta; at: heart auricle; em: emu-

nctorium (excretory organ); exd: lower pericardioduct or excretory duct (“ureter”); exs: excretory

sack; gd: gonoduct; gdg: gonoduct gland; go: gonad; gpd: gono-pericardial interconnection; pc:

pericardium; pd: upper pericardioduct (“reno”-pericardial canal); ve: ventricle.

Figura 6. Sistema gono-pericárdico de los Testaria (según MIZZARO-WIMMER Y SALVINI-PLAWEN,

2001). ao: aorta; aob: bulbo del aorta; at: aurícula; em: emunctorio (órgano excretor); exd: conducto

excretor; exs: saco excretorio; gd: gonoducto; gdg: glándula del gonoducto; go: gónada; gpd: comunicación

gono-pericárdica; pc: pericardio; pd: pericardioducto proximal (canal reno-pericárdico); ve: ventrículo.

(central midgut region) and the intestine

is a subsequent narrowed posterior

midgut, in Caudofoveata both the

midgut sack and the midgut duct

(“intestine”) together represent the pos-

terior midgut. These midgut relations in

Caudofoveata and Testaria therefore

cannot be evaluated as being homolo-

gous, and a unifying taxon *Hepagas-

tralia* (HASZPRUNAR 2000) consequently

is misleading.

Though change in nourishment —

such as from herbivory to carnivory —

is followed by reorganising adaptations

in the alimentary tract, such reorganisa-

tion towards carnivory merely involves

a simplification of the existing configu-

ration (with stomach, etc.). As demon-

strated in several conchiferan groups

(e.g. Cephalopoda, Scaphopoda, “Septi-

branchia”; cf. SALVINI-PLAWEN 1988a),

however, the configuration itself

remains unaltered and the genetic infor-

mation for the regionated midgut of Tes-

taria is not suppressed. Despite a regu-

lative plasticity of the alimentary tract

PR comparative view with

cephalopods etc. — a reorganisation of

the midgut relations in Testaria-Pla-

cophora towards those in Solenogastres

and /or Caudofoveata (PELSENEER, 1890)

appears to be an out-dated speculation

(see also NIERSTRASZ, 1910, and HOFF-

MANN, 1930). The main point is that the

uniformly body-filling _midgut of

Solenogastres (and, longitudinally sub-

divided, of Caudofoveata) refers to a

primary configuration rather than a

regressive adaptation from a regionated

testarian midgut.

The excretory system in Testaria typ-

ically performs two different functions.

In Placophora and Conchifera a primary

urine is produced by ultrafiltration

through podocytes at the atrial heart

epithelum (WKLAND, 1980, 1982,

ANDREWS, 1981, 1988, REYNOLDS, 1990).

This primary urine is drained out of the

pericardium by the pericardioducts

(“renopericardial” canals). Here, the

central portion of the two pericardio-

ducts is elaborated to secrete solutes, to

reabsorb organic molecules as well as to

abstract resulting waste material. By an

emunctorial process it produces the sec-

ondary urine and thus represents an

Iberus, 21 (1), 2003

excretory organ (“kidney”) or emuncto-

rium, with the outleading lower pericar-

dioduct — excretory duct or exit canal

(“ureter”) — opening into the mantle

groove or cavity (Fig. 6). In Solenogas-

tres, the production of the primary urine

is likewise performed by the atrial heart

epithelium (REYNOLDS, MORSE AND

NORENBURG, 1993, MORSE AND

REYNOLDS, 1996). In contrast to Testaria,

however, the pericardioducts are

simple, variably ciliated mesodermal

outlets which open into the internalised

mucous tract portions of the ectodermal

mantle cavity (spawning ducts, see

Figure 2); the pericardioducts exhibit no

transformation into an excretory organ

(emunctorium), and no secondary urine

is produced.

In Caudofoveata the primary urine

produced by the atrial heart epithelium

(SALVINI-PLAWEN AND BARTOLOMAEUS,

1995) is drained by the ciliated pericar-

dioducts into the voluminous glandular

ducts, which, in turn, then open into the

mantle cavity (Fig. 2). As the organogen-

esis and morphological significance of

these glandular ducts or sacks are

unknown (see below), no homology can

be ascertained. Crystals or electron-

dense vacuolated inclusions appear to

be present in the cells of the glandular

ducts (WIREN, 1892: 55; HEATH, 1911: 54

and 71; SCHELTEMA, 1978; Tscherkassky

in SCHELTEMA ET AL., 1994: Fig 23G ),

indicating potential formation of excre-

tory products. Yet, these latter organs

themselves (“coelomoducts” in SCHEL-

TEMA, 1978, “lower gametoducts” in

MORSE AND REYNOLDS, 1996) are not

part of the pericardioducts (see below)

and thus appear not to be homologous

to emunctoria. At best, they could repre-

sent analogous organs autapomorphi-

cally adapted to an equivalent or similar

excretory function in caudofoveate fore-

runners.

The organisation of both organ

systems clearly demonstrates that the

axial regionation of the midgut as well

as the elaboration of pericardioducal

excretory organs in Testaria are evolu-

tionary novelties (autapomorphies) that

are not yet differentiated at the apla-

cophoran level; a postulated loss of

excretory organs (SCHELTEMA, 1996 in

contrast to IVANOV, 1996) is not sup-

ported by organogenesis (see above).

On the other hand, the pallial position

of the excretory organs outside the

dorsoventral musculature in Tryblidia

and Bivalvia (in contrast to Placophora;

HASZPRUNAR AND SCHAEFER, 1997b)

could be interpreted such that the elabo-

ration of emunctoria is polyphyletic.

Based on these and other arguments

(SALVINI-PLAWEN, 1988b, SALVINI-

PLAWEN AND BARTOLOMAEUS, 1995), the

excretory organs in Mollusca-Testaria,

though functionally reflecting a so-

called metanephridial system (RUPPERT

AND SMITH, 1988), have organogeneti-

cally (morphologically) nothing to do

with “metanephridia” (or “nephridia”,

“kidneys”) of other animal groups. In

order to avoid unjustified homologisa-

tions and for the sake of accuracy, the

testarian organs should preferably be

termed emunctoria (see HOFFMANN,

1934, 1937).

3. RADULA CONFIGURATIONS

The Solenogastres, with their

retained conservative midgut, exhibit a

remarkable diversity in the mantle char-

acters, in the types of radula, and in the

elaboration of the accessory genital

organs. The conservative level of

Solenogastres, classified according to

their mantle cover as Pholidoskepia, is

clearly reflected by a thin cuticle with

solid, scaly sclerites. This conservative

level is also evidenced by the same con-

dition in Caudofoveata and Placophora

(see below). The basal mantle configura-

tion in Solenogastres, ¡.e. scales, is sup-

ported by the change of the primary

scales to hollow spicules during meta-

morphosis in Rhopalomenia aglaopheniae

(Kow. and Mar., 1887) and Epimenia

babai (Pruvot, 1892, Okusu, 2002), both

belonging to the advanced level of Cavi-

belonia. Also, Pholidoskepia either

possess poorly outlined ventral foregut

glandular organs (comparable to those

in Caudofoveata) or a paired duct with

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

de AX a

Figure 7. EM cross section through left part of most anterior radula of Wirenia argentea, with

dorsal tooth (ra above) and ventrally turned tooth (ra below), with interconnecting radular mater-

¡al (ic) forming symphysis to teeth of right side (photo: G. Haszprunar).

Figura 7. Corte al microscopio de la parte izquierda de la zona anterior de la rádula de Wirenia argen-

tea, con el diente dorsal (ra, arriba) y el diente torcido ventralmente (ra, debajo), con material de inter-

conexión (1c) formando una sínfisis con los dientes del lado derecho (fotografía G. Haszprunar).

subepithelial follicles (so-called type A),

whereas among Cavibelonia — apart

from organs of type A — four additional

types are elaborated (SALVINI-PLAWEN,

1972b, 1978). With regard to the radula,

Pholidoskepia possess either monoserial

or biserial types, whereas Cavibelonia

also have polyserial/polystichous

radulae. :

Investigations on biserial-distichous

radulae in some species revealed partic-

ular conditions. PRUVOT (1900) points to

the development of the distichous

radula of Pruvotina impexa out of a

monoserial anlage. SALVINI-PLAWEN

(1978) describes the developmental

sequence of the biserial radula of Plawe-

nia schizoradulata: it begins with a sym-

physis (as a U-shaped, monoserial

organ), which divides towards the

pharynx to become truly biserial.

SALVINI-PLAWEN (1988a) demonstrates

that the distichous radula of Wirenia

argentea Odhner, 1921 (syn. Aesthoherpia

glandulosa S.-Plawen) is interconnected

by radula material forming a symphysis

(Fig. 7), thus being in truth a monoserial

organ without differentiating a real

radular membrane (ribbon). In contrast,

a basally separated membrane can light-

optically be distinguished in Caudo-

foveata (SCHELTEMA, 1978, SALVINI-

PLAWEN, 1981b, DEIMEL, 1982, SCHEL-

TEMA ET AL., 1994). WOLTER (1992) con-

firms for the investigated Solenogastres

(Genitoconia sp., Wirenia argentea,

Micromenia fodiens (Schwabl,1955) and a

species of Cavibelonia) the lack of a real

ribbon, but states that the Caudofoveata

(Scutopus robustus S.-Plawen,1970) like-

wise do not differentiate a true, sepa-

rated radular membrane, but rather a

type of pre-ribbon. The rudimentary

ribbon in Solenogastres, however, is

Iberus, 21 (1), 2003

attached to the basis of the teeth by

microfibrils that extend into the under-

lying epithelium. These are lacking in

Caudofoveata and Placophora, and the

rudimentary membrane of Caudo-

foveata is linked to older teeth only by

fibrillous hbundles (WOLTER, 1992,

SCHELTEMA ET AL., 1994), as is the ribbon

in Placophora. In connection with scle-

rotisation, this condition distinctly

simulates in Caudofoveata a light-opti-

cally separated ribbon (SALVINI-PLAWEN,

1981b, Wolter, 1992).

All these conditions point to the fact

that a monoserial radula, including the

biserial-distichous radula with sym-

physis, represents the most conservative

type, only retained in Solenogastres.

This is also reflected by computer-

processed analyses (see below), accord-

ing to which the Solenogastres-Pholi-

doskepia with monoserial radulae rep-

resent the basal group. Nearly all

Solenogastres are predators on Cnidaria

(SALVINI-PLAWEN, 1981b), and they

probably retained their conservative

midgut due to their carnivorous habit.

The different types of radulae (family

characters) infer that most of these types

are well-suited or even adapted for

uptake of cnidarian food, particularly

the monoserial and distichous types.

Monoserial radulae (without sym-

physis) are elaborated in Dondersiidae,

- Macellomeniidae, and (?) Sandalomeni-

idae (all Pholidoskepia) as well as in

Acanthomeniidae and Amphimeniidae

(Cavibelonia) (SALVINI-PLAWEN, 1978).

In several species, adaptation has led to

a fused, unpaired outlet of the ventral

foregut glandular organs associated

with the radula. These include Macel-

lomenia (3 spp.), Stylomenia (2 spp.),

Pholidoherpia (2 spp.), Sandalomenia? (2

spp.), two species of Micromenia and

Dondersia festiva Hubrecht, 1888. In con-

trast, other Dondersia (4 spp.), Microme-

nia simplex Leloup,1948, Nematomenia

(N. flavens (Pruvot,1890) and at least five

more species), Lyratoherpia (2 spp.),

Ichthymenia ichthyodes (Pruvot, 1890),

Heathia porosa (Heath,1911), Acanthome-

niidae (3 spp.) and Amphimeniidae (23

spp.) still possess the paired outlet of

the glandular organs (e.g., PRUVOT,

1891, SALVINI-PLAWEN, 1978, SCHEL-

TEMA, 1999 HANDL AND SALVINI-

PLAWEN, 2001, 2003). The distichous

type (in general with paired outlet of

glandular organs) among Pholidoskepia

is elaborated in Gymnomeniidae, Le-

pidomeniidae and Meiomeniidae. The

distichous-biseral type, however, is the

predominant one among Solenogastres

in general; in some members the radula

sheath is even distinctly paired at its

proximal end (e.g., Pruvotina peniculata

in SALVINI-PLAWEN, 1978, Plawenia

sphaera in SCHELTEMA AND SCHANDER,

2000).

All Caudofoveata definitely possess

a distichous radula, though they appear

to be mainly micro-omnivores or micro-

carnivores. In Chaetodermatidae this

nourishment has led to a highly modi-

fied radula apparatus that is correlated

to a kind of stomach region in the

midgut with a protostyle and gastric

shield (SCHELTEMA, 1978, SALVINI-

PLAWEN, 1981b, 1988a). Such a condition

reflects an increasing adaptation to spe-

cialised nourishment, moving away

from micro-omnivory due to a less suit-

able distichous radula. This could point

to an original carnivory in caudofoveate

forerunners. '

SIRENKO AND MINICHEV's (1975)

thesis of a monoserial origin of the pla-

cophoran radula is questioned by

EERNISSE AND KERTH (1988), who under-

line a paired origin with three teeth at

each side. Other molluscs may likewise

show paired anlagen or even an initially

distichous stage of the radula (KERTH;

1983, EERNISSE AND KERTH, 1988); this

led to the assumption that a distichous

type might have been the original

radula of molluscs. All this supports

carnivory as being original in contrast to

the hitherto supposed herbivory.

Accordingly, the primitive radula

served to attack and split open soft-

bodied prey. A monoserial elaboration

of a midventral, serially chitinised pha-

ryngeal cuticle as firm bars or flexible

plates (see symphyses) — at each side

elaborated to a definite cusp, denticle or

hook — meets the conservatively

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

retained types in Solenogastres and

most likely represented the original

radula.

There is no mutuality with respect to

the radula support. All Caudofoveata

investigated possess a pair of elongate,

well-delimited, compact bolsters which

consist of muscular and connective

tissue, frequently interspersed by

turgescent or chondroid cells (e.g.,

WIREN, 1892, SCHELTEMA 1972). In

Solenogastres a radula support fre-

quently is represented only by some

accumulation of muscular and connec-

tive tissue in median or paired arrange-

ment. More differentiated supports

consist of a distinct muscular concentra-

tion which may also include or be asso-

ciated with turgescent cells (e.g.,

Amphimeniidae, Simrothiellidae;

SALVINI-PLAWEN 1978, SCHELTEMA ET

AL., 1994). Occasionally (some Simroth-

iellidae), turgescent cells are very volu-

minous and distincly arranged or may

even be compacted to a pair of muscu-

larly surrounded vesicles. Though such

a latter condition resembles the radula

vesicles of Placophora and Tryblidia (cf.

WINGSTRAND, 1985), the exceptionality

in advanced Solenogastres (Cavibelonia)

clearly implies analogous similarity.

4. SYSTEMATICS OF SOLENOGAS-

TRES

THIELE's — classification of the

Solenogastres (1913) — accepting four

families with respect to the mantle cover

and the presence /absence of respiratory

organs (secondary gills) — had been

abandoned by SALVINI-PLAWEN (1967)

and replaced by a pure alphabetical

order of the genera. Investigating an

extensive material, SALVIN-PLAWEN

(1978) then proposed a new classifica-

tion based on the conditions of the

mantle cover (orders, supraorders) and

the various differentiation of the foregut

glands and the radula (families).

According]ly, the representatives with a

mantle cover of calcareous scales and of

a thin cuticle, the order Pholidoskepia,

are regarded as being conservative.

They are classified together with the

Neomeniamorpha (with thin cuticle and

spearlike, grooved mantle bodies, and

with a highly complex accessory genital

apparatus) as Aplotegmentaria. The

small order Sterrofustia (thick cuticle;

elongate, solid mantle bodies) and the

order Cavibelonia (with acicular, hollow

spicules and mostly with a thick cuticle)

are classified as Pachytegmentaria.

However, some findings raise the possi-

bility that the formation of hollow

needles (Cavibelonia) might be poly-

phyletic (SALVINI-PLAWEN, 1978, SCHEL-

TEMA AND KUZIRIAN, 1991, HANDL AND

SALVINI-PLAWEN, 2003).

To assess the phylogenetic interrela-

tionships and evidence for the system-

atic arrangement of the families of

Solenogastres, two character matrices

for all genera were prepared and sub-

mitted to parsimony searches in PAUP*

4.0b10 (SwOFFORD, 2002). In addition to

the Placophora and Caudofoveata, the

taxon “Solenogastres” is included,

defined by the putative plesiomorphic

character states (when evaluable) or by

“?” entries. Searches were limited to

750.000 shortest trees (using the

command “set maxtrees”) and 22 hours

(using the timelimit option in the

command hsearch), due to the high

number of taxa and resulting trees as

well as the limited computational

resources. All searches hit the set

number of trees, except for the weighted

analysis for 53 characters which hit the

time limit earlier.

(1) A matrix of 35 characters (1-35 in

Tables II and II) at the level of presently

accepted families was analyzed using

unweighted and weighted parsimony

(shortest trees of unweighted analysis:

88 steps, CI=0.3295, RC=0.2792). Both

analyses yield largely unresolved strict

consensus trees (not shown) with three

major polytomies: the genera Wirenia

and Genitoconia (part of Gymnomeni-

idae); the other Pholidoskepia including

the “Solenogastres”; and all remaining

advanced taxa. Among the latter, only

Amphimeniidae and Rhipidoherpiidae

appear monophyletic.

Iberus, 21 (1), 2003

(2) Unweighted parsimony analysis

of an enlarged matrix with 53 characters

(Tables Il and III) returned a similar

strict consensus tree as analysis (1) (not

shown; shortest trees: 161 steps,

CI=0.2609, RC=0.2074). It separates only

the Aplotegmentaria and the

Pachytegmentaria. The basal

Aplotegmentaria consist of a polytomy

of 22 branches of Pholidoskepia (includ-

ing “Solenogastres”) with a common

line for Gymnomeniidae, Acanthomeni-

idae, and monophyletic Neomeniamor-

pha. The advanced level of

Pachytegmentaria represents a poly-

tomy of 43 branches (e.g. Simrothielli-

dae in four lines), including the mono-

phyla Amphimeniidae, Strophomeni-

idae, and Rhipidoherpiidae.

(3) In contrast, the majority-rule con-

sensus tree (Fig. 8 A) is much better

resolved. It separates the “Solenogas-

tres” as the basal-most line from a poly-

tomy of nine branches; seven of these

represent most Pholidoskepia with

monoserial radulae, one unites the

Pholidoskepia with distichous radulae

and the monophyletic Neomeniamor-

pha, and one leads to three dondersiid

genera and to the Pachytegmentaria.

(4) The analysis of the 53-character

matrix attributing the weight 5 to the

characters 18 and 53 (Tables Il and III)

(shortest trees: 174 steps, CI=0.2874,

RC=0.2303) returns an even better

resolved majority-rule consensus tree

(Fig. 8 B). It shows all Pholidoskepia

with monoserial radulae in a basal poly-

tomy, whereas those with distichous

radulae are tied to the genera of the

Sterrofustia, Neomeniamorpha, and

Cavibelonia. Note that in this tree the

Neomeniamorpha appear ¡in an

advanced position together with cavi-

belonian and sterrofustian genera.

The low number of characters avail-

able at the genus-level permits only

limited resolution of the 87 ingroup

taxa. First, the Pholidoskepia represent

indeed the basal level among Solenogas-

tres. Second, the Pholidoskepia with

-82

monoserial radulae appear to be conser-

vative compared with those with disti-

chous radula. Third, the order Neome-

niamorpha is confirmed to be mono-

phyletic, though of doubtful relation-

ship. Fourth, a polyphyly of the Cavi-

belonia is not visible in these trees;

rather, a possible polyphyly is indicated

for the Sterrofustia. Finally, only two

families as presently defined are always

monophyletic, the Rhipidoherpiidae

and Amphimeniidae. Yet, in most

majority-rule-consensus trees of the

larger data set (Fig. 8 A, B), also Acan-

thomeniidae, Gymnomeniidae,

Strophomeniidae, Epimeniidae and Pro-

neomeniidae appear monophy]letic.

It is evident that the “present:

absent” coding (Table III) includes

various convergencies (homoplastic

characters) and most characters in the

trees have low constistency indices (CI <

0.5). In some cases this implies unex-

pected results. Thus, in Fig. 8A, the lack

of the terminal sense organ is a synapo-

morphy for three dondersiid genera

(Helluloherpia, Ichthyomenia, Heathia) and

brings them in a separate position,

whereas the splitting of the

Rhopalomeniidae is due to the foregut

glands and the radula (in Dinomentia).

Only the presence of the radula in

Dinomenia isolates this genus from other

Rhopalomeniidae in Fig. 8 B. The divi-

sion of the Phyllomeniidae refers to the

absence of the terminal sense organ in

Lituiherpia and Ocheyoherpia. The latter

condition unites these with the parar-

rhopaliid Forcepimenia. The three fami-

lies, Phyllomeniidae, Pararrhopaliidae

and Rhopalomeniidae, however, due to

the characters of the mantle cover, the

radula and the foregut glands, appear

not unexpectedly in an intermediate

position between the Pholidoskepia and

_the derived Cavibelonia (see systematic

arrangement in SALVINI-PLAWEN, 1978).

These examples demonstrate that the

available characters are extremely

homoplastic and that homology deci-

sions are far too uncertain to accept the

resulting trees as reflections of the phy-

logeny of the Solenogastres.

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

5. ANAGENETIC LEVELS IN

MOLLUSCA

In view of the two different levels of

elaborations of the midgut and the

excretory system in Mollusca, the apla-

cophoran configurations are clearly

more conservative than those of the Tes-

taria (see also subradular organ, radular

bolster): the respective elaborations

express two subsequent evolutionary

grades of molluscan organisation. This

also holds true for the radula, which in

both aplacophoran groups shows a con-

servative state with a “pre-ribbon” only

(KERTH, 1983, WOLTER, 1992). Within

Testaria, again two clearly different,

subsequent grades are expressed, repre-

senting the polyplacophoran and the

monoplacophoran (conchiferan) levels;

this is also evidenced by cladistic analy-

sis (SALVINI-PLAWEN AND STEINER, 1996,

HASZPRUNAR, 2000). On one hand, the

aplacophoran groups and Placophora

share (as *Aculifera*) several conserva-

tive characters such as the chitinous

cuticle with sclerites, a supra-rectal com-

missure, the heart-ventricle as a middor-

sal invagination of the pericardium and

the paired muscle to roll up, or the ple-

siomorphic organisation of the ciliary

apparatus (LUNDIN AND SCHANDER,

2001). On the other hand, Placophora

and Tryblidia (“monoplacophorans”)

are closely interconnected by the almost

identical organisation of the radular

bolster (WINGSTRAND, 1985) and by the

dorsoventral musculature. The mono-

phyly of the monoplacophoran level

(Conchifera), in turn, is manifested by

the common differentiation of the shell

or concha, of statocysts, of a sub-rectal

commissure, of jaw plate(s), of a style-

sack type of stomach and of cerebrally-

innervated appendages (SALVINI-

PLAWEN, 1988A, SALVINI-PLAWEN AND

STEINER, 1996, HASZPRUNAR, 2000).

The organ system that most clearly

demonstrates the evolutionary pathway,

however, is the mantle cover and its

associated musculature. There is a

name-giving, gradual anagenesis of the

morphologically dorsal integument

(mantle, pallium) from the apla-

cophoran to the polyplacophoran and to

the monoplacophoran (conchiferan)

levels. The mantle in Solenogastres and

Caudofoveata is homogeneously

covered by a chitinous cuticle with

embedded, unicellularly formed arago-

nitic sclerites, such as is also differenti-

ated in the girdle or perinotum of Pla-

cophora. Evaluating the different kinds

of elaborations shows that squamous

elements or scales represent the most

conservative types when compared with

acicular spicules and other types of scle-

rites (SALVINI-PLAWEN, 1972a, HAAS,

1981). This common character in apla-

cophoran and placophoran molluscs is

paralleled by the Musculus longitudi-

nalis along the mantle border; it rolls the

animals up and is present in Solenogas-

tres (M.l.ventralis) and in some lower

Caudofoveata (M.l.ventralis in Scutopus)

as well as in Placophora (M.l.lateralis;

but see HASZPRUNAR AND WANNINGER,

2000). On the other hand, the homoge-

neously elaborated mantle — underlain

by layers of outer transverse, intermedi-

ate oblique and inner longitudinal

muscle fibres (such as in many other

invertebrate groups) — in the apla-

cophoran level is strutted against the

ventral body (foot) by a paired

dorsoventral musculature. This is elabo-

rated in an undefined number of serial

pairs of two bundles, the dorsal-outer

ones intercrossing medioventrally. In

Placophora, the integumental muscula-

ture is restricted to bundles associated

with the eight serial middorsal shell

plates, whereas the dorsoventral muscu-

lature is concentrated (HASZPRUNAR

AND WANNINGER, 2000) and grouped to

16 pairs of three bundles each, the

dorsal-outer ones again intercrossing

medioventrally. An identical configura-

tion of dorsoventral musculature is

present in Tryblidia. Here, however,

every two successive bundle groups of

the paired series are mostly fused into

one common bundle. In the new mono-

placophoran (or conchiferan) condition,

the dorsoventral musculature is thus

represented by eight, in part still subdi-

Iberus, 21 (1), 2003

Placophora

Caudofoveata

Solenogastres

Squamatoherpia

Micromenia

Dondersia

Lyratoherpia

Siylomenia

Nematomenia

Acanthomenia

Dondersiidae

Acanthomeniidae

Amboherpia

Sandalomenia Sandalomeniidae

M. aciculata

Macellom. palifera Í Macellomeniidae

M. adenota

Pholidoherpia incertae sedis

Lepidomenia Lepidomeniidae

Meioherpia ; ,,

Maitena Meiomeniidae

Tegulaherpia , a

Nierstraszia Llepidomeniidae

Wirenia

Gymnomenía Gymnomeniidae

Genitoconia

Hemimenia interm.

H.dorsosulcata

Archaeomenia

Neomenia

Helluloherpia

Ichthyomenia

Heathia

Notomenía

Entonomenía

Pruvotia

Driomenía

Dinomenia

Phyllomenia

Lituiherpia

Ocheyoherpia

Harpagoherpia

Forcepimenia

Lophomenia

Hypomenia

Metameria

Pararrhopalia

Halomenia

Gephyroherpia

Pruvotina

Luitfriedia

Eleutheromenia

Labidoherpia

Scheltemaia

Rhopalomenia

Urgorria

Proneomenia

Dorymenia

Doryherpia

Epimenia

Epiherpia

Rhipidoherpia

Thieleherpia

Strophomenia

Anamenia

Syngenoherpia

Imeroherpia

Heteroherpia

Simrothiella

Birasoherpia

Cyclomenia

Kruppomenia

Aploradoherpia

Biserramenia

Plawenia

Helicoradomenia

Spiomenia

Drepanomenia

Rhabdoherpia

Sialoherpia

Unciherpia

Uncimenia

Plathymenia

Pachymenia

Alexandromenia

Proparamenia

Spengelomenia

loa

Amphimenia

Meromenia

Utralvoherpia

Paragymnomenia

HAHAHA HH HH HA EHH HH HH

Neomeniamorpha

Dondersiidae

Notomeniidae

Rhopalomeniidae

Phyllomeniidae

Pararrhopaliidae

Rhopalomeniidae

Proneomeniidae

Epimeniidae

Rhipidoherpiidae

Strophomeniidae

Syngenoherpiidae

Imeroherpiidae

Heteroherpiidae

Simrothiellidae

Drepanomeniidae

incertae sedis

Pararrhopaliidae

Amphimeniidae

Figure 8. Maximum Parsimony analyses of Solenogastres genera according to 53 morphological

characters (see Tables II and III). Note that due to the incongruency and low CI of most charac-

ters, these trees do not represent the phylogeny (see text). A: Majority-rule consensus tree of

750,000 most parsimonious trees returned by unweighted parsimony analysis of the 53-character

matrix. B: Majority-rule consensus tree of 587.000 most parsimonious trees returned by weighted

parsimony analysis of the 53-character matrix (weight 5 for characters 18 and 53).

Placophora

Caudofoveata

Solenogastres

Squamatoherpia

Micromenia

Dondersia

Lyratoherpia

Stylomenia

Nematomenia

Helluloherpia

Ichthyomenia

Heathía

Acanthomenia

Amboherpia

Sandalomenia

M. adenota

Macellom. palifera

-M. aciculata

Pholidoherpia

Llepidomenia

Meioherpia

Meiomenia

Tegulaherpia

Nierstraszia

Gymnomenia

Wirenia

Genitoconia

lituiherpia

Ocheyoherpia

Forcepimenia

Dinomenia

Pararrhopalia

Lophomenia

Hypomenia

Metameria

Halomenia

Gephyroherpia

Pruvotina

Luitfriedia

Eleutheromenia

Labidoherpia

Scheltemaia

Harpagoherpia

PrlSmenia

Notomenia

Entonomenia

Driomenia

Pruvotia

Rhopalomenía

Urgorria

Epimenia

Epiherpia

Proneomenia

Dorymenia

Donyherpia

Rhipidoherpia

Thieleherpia

Strophomenia

Anamenia

Syngenoherpia

Heteroherpia

Imeroherpia

Neomenia

Archaeomenia

Hemimenia interm.

H.dorsosulcata

Simrothiella

Birasoherpia

Cyclomenia

Kruppomenia

Aploradoherpia

Biserramenia

Plawenia

Helicoradomenia

Spiomenia

Drepanomenia

Rhabdoherpia

Sialoherpia

Unciherpia

Uncimenia

Plathymenia

Pachymenia

Alexandromenia

Proparamenia

Spengelomenia

Sputoherpia

Paragymnomenia

Aena

Meromenia

Utralvoherpia

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

a a o dd d+

tal Ele

Dondersiidae

Acanthomeniidae

Sandalomeniidae

Macellomeniidae

incertae sedis

Llepidomeniidae

Meiomeniidae

lepidomeniidae

Gymnomeniidae

Phyllomeniidae

Pararrhopaliidae

Rhopalomeniidae

Pararrhopaliidae

Phyllomeniidae

Notomeniidae

Rhopalomeniidae

Epimeniidae

Proneomeniidae

Rhipidoherpiidae

Strophomeniidae

Syngenoherpiidae

Heteroherpiidae

Imeroherpiidae

Neomeniamorpha

Simrothiellidae

Drepanomeniidae

incertae sedis

Pararrhopaliidae

Amphimeniidae

Figura 8. Análisis de máxima parsimonia de generos de Solenogastres según 53 caracteres morfológicos

(ver Tablas II y II). Nótese que debido a la incongruencia y al bajo CI de la mayoría de los caracteres,

estos árboles no representan la filogenia (ver el texto). A: Árbol de máximo consenso de entre los 750.000

obtenidos por el análisis de parsimonia (misma importancia para todos los caracteres) de la matriz de los

53 caracteres. B: Árbol de máximo consenso de los 587.000 obtenidos por el análisis de parsimonia (peso

5 para los caracteres 18 a 53) de la matriz.

Iberus, 21 (1), 2003

Table Il. Characters to matrix in Table II (cf. THIELE, 1913; HOFEMANN, 1929; SALVINI-PLAWEN,

1967, 1972b, 1978, 1985; SCHELTEMA, 1999, 2000; SCHELTEMA AND SCHANDER, 2000; GARCÍA-

ÁLVAREZ, URGORRI AND CRISTOBO, 2000; HANDL AND SALVINI-PLAWEN, 2001, 2003).

Table II. Caracteres de la matriz de la Tabla II (cf. THIELE, 1913; HOFFMANN, 1929; SALVINI-

PLAWEN 1967, 1972b, 1978, 1985; SCHELTEMA, 1999, 2000; SCHELTEMA Y SCHANDER 2000;

GARCÍA-ÁLVAREZ, URGORRI Y CRISTOBO, 2000; HANDL Y SALVINI-PLAWEN, 2001, 2003).

co Y Ox UN FE 0 NN —

a 2)

Cuticle: thin: 0, thick: 1

Shell plates: absent: 0, present: 1

Mantle bodies: scales: O, other: 1

Acicular spicules: absent: O, present: 1

Hollow needles: absent: O, present: 1

Mantle bodies: one layer:0, several layers: 1

Mantle bodies: hooks absent: O, hooks present: 1

Mantle bodies : spear grooves absent: O, present: 1

Foot: present: O, absent: 1

Ctenidia: present: O, absent: 1

Other respiratory organs: absent: O, present: 1

Osphradia: present: 1, as diSO: 0

Subradular organ: absent: O, present: 1

Midgut: not regionated: O, regionated: 1

Gonoducts: present: O, absent: 1

Aorta: absent: O, present: 1

Sexes: separate: O, hermaphroditic: 1

Ventral foregut glands: present: O, absent: 1

Ventral foregut glands: type A: O, other: 1

Ventral foregut glands: type B: O, other: 1

Ventral foregut glands: type C: O, other: 1

Ventral foregut glands: type D: O, other: 1

Ventral foregut glands: folicular: O, other: 1

Rodula: present: O, absent: 1

Radula: monoserial: O, other: 1

Rodula: monoserial plates: O, other: 1

Radula: biserial: O, other: 1

Radula: distichous: O, other: 1

Radula: tetraserial: O, other: 1

Rodula: polystichous: O, other: 1

Receptacula seminis: absent: O, present: 1

Receptacula seminis: in bundles: 1, other: 0

Mucous tract glands; epithelial: O, subepithelial: 1

Copulatory-stylet gland: absent: O, present: 1

Commissural sack organ: absent: O, present: 1

Dorsoterminal sense organ: present: O, reduced: 1

Mouth and atrium: separate: O, common/fused: 1

Copulatory stylets: absent: O, present: 1

Secondary genital opening : paired: O, single/fused: 1

Dorsal papilla foregut gland: absent: O, present: 1

Peripharyngeal glandular organs: absent: O, present: 1

Opening of ventral foregut glandular organs: sub-radular: O, pre-radular: 1:

Midgut: without regular constrictions: O, serial pouches: 1

Paired ventral radula sack: missing: O, present: 1

Distally axe-shaped spicules: missing: O, present: 1

Epidermal popillae: not elaborated: O, present: 1

Buccal ganglia: present: O, reduced /replaced: 1 (Epimenia)

gonoducts proper: present: O, not elaborated: 1

Aorta: not elaborated: O, present: 1

Aesthetes: not elaborated: O, present: 1

Subradular sense organ: not present: 0, present: 1

Head-egion: not elaborated: O, separated: 1

Naikformed spicules: absent: O, present: 1

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

vided pairs of bundles (LEMCHE AND

WINGSTRAND, 1959, SALVINI-PLAWEN,

1969, 198la, WINGSTRAND, 1985,

HASZPRUNAR AND SCHAEFER, 1997a).

Beyond this configuration of dorsoven-

tral musculature, the almost identical

organisation of the radular support like-

wise points to the polyplacophoran

level as the direct forerunner of the

monoplacophoran level by fusion of the

eight shell plates towards the concha.

This is also paralleled by the Merismo-

concha (Yu, 1984, SALVINI-PLAWEN,

E: tn higher Conchitera, the

dorsovental musculature is even more

concentrated (see also fossil Cyrtone!l-

ida and Sinuitopsida), resulting in a dif-

ferent number of paired bundles reflect-

ing the shape and functional demands

on the shell in each class.

Altogether, the mantle cover, the

dorsoventral musculature, and the elab-

oration of the alimentary tract present

successive, overlapping sequences of

evolutionary differentiation. Within

Mollusca they reflect a distinct, gradual

anagenesis from the aplacophoran to the

polyplacophoran and monoplacophoran

(conchiferan) levels. These evolutionary

grades also indicate the direction in

which the homology series must be read

and defined. This clearly reveals the

Solenogastres and Caudofoveata to be

early, conservative offshoots of the mol-

luscan organisation (rather than being

derived from polyplacophorans). It like-

wise excludes a monophyly of *Aculif-

era* (e.g. SCHELTEMA,1996, IVANOV,

1996), which would incorrectly assume

the above-presented coincidences in the

alimenary tract and the excretory

system in Testaria to be convergencies; it

also removes most of the potential

synapomorphies from the Solenogastres

and Caudofoveata (revealing them to be

symplesiomorphies).

6. PLESIOMORPHIC CONDITIONS

OF MOLLUSCA

The evaluation of plesiomorphic

characters in Mollusca and of the origi-

nal molluscan organisation yealds two

results: it summarises the smallest

common denominators of features, and

it traces the anagenetic conditions back

to their most basic elaboration.

Within the conservative level of

aplacophoran molluscs, the Caudo-

foveata exhibit, in some characters,

more advanced conditions than

Solenogastres. Of particular interest are

the protostyle formation in the midgut,

an aorta, a radula ribbon and ctenidia —

all also present in Testaria. Are these

characters synapomorphies or merely

convergent autapomorphies in Caudo-

foveata and Testaria (or simply reduced

in Solenogastres) ? Cladistic computer

programs, due to their reflection of

numerical taxonomy, use the parsimony

argument to incorporate them as

synapomorphies and make the

Solenogastres the earliest offshoot (cf.

SALVINI-PLAWEN AND STEINER, 1996,

HASZPRUNAR, 2000). The elaboration of

a midgut section with a protostyle and

gastric shield among Caudofoveata has

been verified in the most advanced

Chaetodermatidae only, but the func-

tional parallelism of this elaboration is

obvious (see also MORTON, 1960, for

similar differentiations in other animal

groups). Conversely, the adaptive flexi-

bility of the circulatory system through-

out Mollusca casts doubt on aorta for-

mation as being a strong synapomor-

phic character rather than a function-

ally-tied, convergent differentiation. As

outlined above, the light-optically rec-

ognized radula membrane in Caudo-

foveata is a kind of pre-ribbon, but

intermediates between the elaborations

in Solenogastres and Placophora. This

may reflect either a synapomorphic or a

convergent differentiation of a true

ribbon and thus weakens the argument.

With regard to the ctenidia and their

characteristic elaborations (see SALVINI-

PLAWEN 198la, 1985, HASZPRUNAR,

1987), they are generally accepted to be

a typical molluscan character and hence

proposed to be already differentiated in

the common forerunners (archimol-

luscs). Their absence in Solenogastres,

either within monophyletic apla-

cophorans or as a sister-group of Tes-

Iberus, 21 (1), 2003

Table III. Matrix for Figure 8.

Tabla II. Matriz para la Figura $.

Placophora

Caudofoveata

Solenogastres

Dondersia

Micromenia

Stylomenia

Lyratoherpia

Nematomenia

Ichthyomenia

Heathia

Squamatoherpia

Helluloherpia

Sandalomenia

Gymnomenia

Wirenia

Genitoconia

Lepidomenia

Tegulaherpia

Nierstraszia

Meiomenia

Meioherpia

Pholidoherpia

Macellom palifera

M aciculata

M adenota

Archaeomenia

Hemimenia interm

H dorsosulcata

Neomenia

Imeroherpia

Phyllomenia

Harpagoherpia

Lituiherpia

Ocheyoherpia

Heteroherpia

Rhabdoherpia

Notomenia

Pararrhopalia

Pruvotina

Labidoherpia

Eleutheromenia

Gephyroherpia

Luitfriedia

Lophomenia

Metameria

Hypomenia

Halomenia

Forcepimenia

Scheltemaia

Unciherpia

Uncimenia

Sialoherpia

Rhopalomenia

Entonomenia

Dinomenia

Driomenia

Pruvotia

Urgorria

123450678 910111213141516171819202122 23242526 272829 303132 33 34 35 36 37 38 39 40 41 42 43 44 45 4647 48 49 5051 5253

010000000001110

000000001000001

00000000010000

0000000001?000

00000000010000

00000000011000

00000000010000

0000000001?000

00000000010000

HOPE

10011

1010?1

1150 10:041

10150004]

101001

1M05150405]

101001

140150:01

101001

1010??

IR0m140+1011

17010141

104.041]

150140:041

101001

0000000001?000101001

00000000011000101001

0000000001?2000101001

0000000001?000101001

AA RIADA MODO AAA

00005000101 1040401011491

0500-0:0:-0 01 011.:0:0:0:1011:11

10010000 150911040501 OMA

10110100011000101010

10010100010000001001

101150125000 15000000 10150005

100101 1"0"0"1-0"0"0"01 0105007

1001 1"0"010*010"0"0 "01000

10110100010000101010

AA AA

1010000.0-0-1?000101001

AAA AI ADA AA AU

1011100000 10000

AA AAA

II A A AA

10 AAA 150-070 140510401

AAA EUA A AAA UA OO

150151 12:0:0:0,1020:0:01 0150504

10011 1400090100 050:150 15001

10111?00010000101001

IO AAA AAA

Or01 1105010" 152200071404 1.-05091

150 MAA ANACO AA NO AO 1

0150-11 12415000 TE1COL0r0 10

0 AO OOOO IA

IAEA EOL

MOI TAO OO OOOO ICOROA

10111100010000101001

LOA 0 PO nO O FOO APOFIAO OA

IOMA OSO PO IO AO Oro O IADIDA

AAA AA

AAA ADA O

VD. —- —= =O€O == —= — ———————— — ll ll ll ll ll Sl ll Sl ll ld lo lo ld ll ld ld dd dl o dl o o a al a a a a a a cd ad al cd cl ad ad ad ad a

A CY dl Al Al ld Al Al dl Al dd dl ld ld ld ld ld dd dl ld dd o al a ad al a cl cl a a cl a cd cl a a a a cd cl a a ca a ad cd cl cd cal cd cl ad dl cd ad ad

O OOOO ooo o

OOOO ooo - —

DN ¡—i —y ld ll dl ld dd

—

== OO0OO0O0O0OO0OO0OO0OOoOOoOoOoOooo Oo

ooo oo——r—— vo» oo

—i

—

Oo oooO

Da a lr ls lr —

—i

—

O OOOO Oo Ooo Oo

A E A E A AN

—

—i

Ed :

A o ano a a ao SS A IN

—

000000000000000?0011110

000000?0000000000010000

200000000000?0000000000

10000000100010000000000

10000000100000000000000

10000001100010000000000

10000000100010000000000

100000101?20000000000000

10000110100010000000000

10000000100000000000000

10000110100000000000000

11000000100000000000000

00001101100010000000000

10001011?00010000000000

10001111100000000000000

20000010100000000000000

20000001100000000000000

10000011100010000000000

2000?011??0000000000000

2000?010??0000000000000

220?001?100000000000000

100000?01000?0000000001

100000101000?0000000001

20000?00?00000000000001

10010011000?10000000000

1001021 711.0-0-21000'0-010:04040%0

10010011000?10000000000

10010011?00?10010000000

10010011000010010000000

20000001000010010100000

10000?001000000?0000000

100001101000100?0000000

100?0101100010000000000

10000011100010010000000

10000000100? 10010000000

10000?200000010010000000

10000001110010010000000

10000010110010010000000

10000011110010000000000

10000010100010000000000

10000000100010010000000

100000101000?0010000000

100000001100?0010000000

10000000110010010000000

20000?00110010010000000

10000000100010010000000

2000010?100010000000000

000010111000?00?0000000

10000010101?10010000000

1000001010120 151-000 40400000

1000000?101?10010000000

10000000100010010000000

10000001100010010000000

10000100100010010000000

10000010100010010000000

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

Table MI. Continuation.

Tabla II. Continuación.

12345678 9 1011121314151617181920212223242526 2728293031 3233343536 37 38:39 40 41 4243 44 45 464] 484950515253

Aconthomenia 000110000100001010011

Amboherpia 04000 0019010000 14051 02415

Amphimenia ORI OOO 1000/0150 15001. 101

Proparamenia LOTA ADO ADA ALDO USA

Pachymenia ¡ONAIAIOnO 051 1050-0101 2001716

Alexandromenia 101111000110001010111

Spengelomenia 101111000110001010111

Paragymnomenia 101111000110001010111

Plathymenia PO AAA AO AA

Sputoherpia PAOLO ALE

Utralvoherpia IIA ON OPOS0 000 A

Meromenia DTO AAA ES.

Simrothiella OO OOOO OA OMAN IO

Kruppomenia AA OA OA A A

Cyclomenia IOMA OOO IO OOOO AO 1

Plawenia OA OI OOO NOR 0

Spiomenia MORO OOO OOOO IAOAIOA

Birasoherpia ATA OO AAA

Aploradoherpia COMWMITDODOOTEO0 AAA OA

Biserramenia O IIAR0 “000 01500500 1501501150 1

Helicoradomenia 1011000001100010101101

Drepanomenia A AO AA A AIDA A OA

Strophomenia OA O TPOCOA0 0010105101

Anamenia OOO OOO IA ONCE

Proneomenia PATA AAA AA O AAA

Dorymenia OOOO OOOO AO AO IA0

Doryherpia IIA OOOO OO OOO

Epimenia A AAA AAA UA

Epiherpio HON IO O OARONO 010 FOO A 1701

Syngenoherpia 105 COJO O, 150:0:0,041,0,1:0:1+01 1

Rhipidoherpia AAA AAA

Thieleherpia AAA OO OOOO AO A OO

taria, was thus regarded as a secondary

loss related to the reorganisation of the

mantle cavity. Yet, computer-processed

cladistics (SALVINI-PLAWEN — AND

STEINER, 1996, HASZPRUNAR, 2000) and

the fairly conservative condition in

several characters (radula, midgut, cir-

culatory and excretory systems, muscu-

lature, larvae) points to the Solenogas-

tres as being an offshoot of the earliest

evolutionary level of molluscs. This con-

servative condition could likewise have

included the lack of ctenidia; this is sup-

ported by the obvious need of many

Solenogastres to differentiate respiratory

organs, which have not been verified as

(recapitulated) ctenidia. The innerva-

tion, however, could provide a counter-

argument: The innervation of the mantle

== OOO _--O0Oo0ooo

== OOOO OOOooo oo

—i

OOO OO Ooo o

011111?0000?21??00000000000000

01111100000010100000000000000

001111?0100110100100010000000

OA OA OLO:050

a O OO OOOO NOOO c0700 0

MIA OO OOOO AO A OnO 040-0000

YAA AA A AAA AO AY

00111110100010100100010000000

2?????10100010100010000000000

OA ARO 0 ONO AAA OOOO 00

OIT AC0 17070010 TONO FIRO FONO 10050000 00

OOOO OO ORAORO O OOOO NOROO

IMAN OOO AO OOOO PO FONO NOROnO

11011110000011100010000000000

11111100000001100000000000000

11011110000011100011000000000

IMA A AO OOOO OOO OOO 00004070

IO OOO OOO OO O ONO Or000

11011120000010100000000000000

11011110000000100000000000000

IMAOA IAAF E OOO ORO RO POC OMOLO

2?????00000110100010010000000

2?????11000010000010010000000

11011111000010000010010000000

11111010000010100010010000000

¡MAA OOO OA OOOO ANA OROROROrO

111110?0000011000010010000000

IMAN ARO ONO ONO RONOFOROrO

111011??000010100010010000000

11101111000011100010010000000

IO OO 00 1 1000.01 0 005150.0-050506040

AAA ALO AAA OA OADADAD DADO

cavity organs from the suprarectal com-

missure is almost identical in Solenogas-

tres, Caudofoveata and Placophora

(SALVINI-PLAWEN, 198la, HASZPRUNAR,

1987). Apart from the dorsoterminal /

osphradial sense organ, this also

includes the ctenidia in Caudofoveata

and — multiplied — in Placophora. A

respective innervation pattern of the

frontal portion of the spawning ducts in

Solenogastres could indicate an original

presence of ctenidia prior to their loss

due to the reorganisation (internalisa-

tion) of the mantle cavity.

A similarly ambivalent character is

the gono-pericardioducts, i.e. apla-

cophoran molluscs lack proper gonod-

ucts. Is this a plesiomorphy, a synapo-

morphy of both groups, or analogy? As

Iberus, 21 (1), 2003

is known from development, the differ-

entiation of gonads as a rostral elonga-

tion of the pericardium, with retroperi-

toneal incorporation of the primary

germ cells, appears to be a key character

of Mollusca (gono-pericardium). The

same conditions are found, e.g., in

Siphonopoda (“cephalopods”) and

Bivalvia-Anodonta, which retain the

interconnection between pericardium

and gonads. In addition, the late anlage

of separate gonducts (apart from peri-

cardioducts) in Placophora and

Siphonopoda-Coleoida (HIGLEY AND

HEaTH, 1912, HOFFMANN, 1937), as well

as the variable gonoduct formation in

Conchifera (RAVEN, 1958), suggest that

the gonoducts are relatively young or

new differentiations (with genetically

weak determination), and may be poly-

phyletic among Testaria. Accordingly,

the configuration in both Solenogastres

and Caudofoveata could represent a

true plesiomorphy (not yet developed

gonoducts) or a paedomorphy (reten-

tion of developmental stage). The pres-

ence of true gonoducts in Solenogastres-

Phyllomenia, due to the much altered

configuration of the pericardioducts and

spawning ducts (cf. SALVINI-PLAWEN,

1978), could represent a specialisation

(autapomorphy) within that genus; it is

thus per se not conclusive (see also

Haszprunar, 1992). Yet, the possible sec-

ondary loss of gonoducts in Solenogas-

tres-Driomenia (HEATH, 1911, HOFFMAN,

1949) lends weight to the Phyllomenia

argument. If, on the other hand, the lack

of gonoducts is truly a secondary loss

(as more generally assumed), due to the

independently narrowed body in

Solenogastres and Caudofoveata, this

reduction would then reflect analo-

gously acquired adaptations or paral-

lelism (rather than a synapomorphy).

The narrow, ciliated pericardioducts

of Caudofoveata often unite abruptly

via a sunken opening into the paired

glandular duct or sack (e.g., HEATH,

1905: 721); only the latter are connected

to the mucous tracts of the mantle

cavity, into which they open ventrolater-

ally by means of a sphincter. These glan-

dular organs often extend beyond the

pericardioduct opening (SALVINI-

PLAWEN, 1972a), be it rostrally (e.g.,

Prochaetoderma californicum Schwabl,

- 1963) or dorsally (e.g., Scutopus robustus;

Fig. 2); in immature Scutopus ventrolinea-

tus S.-Plawen,1968, the glandular sacks

show a dorsoventral extension, with the

sunken pericardioduct openings located

centrally. In contrast, the pericardio-

ducts in many Chaetodermatidae open

axially into the dorsoterminally bent

glandular ducts. Due to our lack of

information on organogenesis, the

homology of these glandular ducts

remains unclear. With respect to their

totally different histology (HEATH, 1905,

1911, SALVINI-PLAWEN, 1972a, SCHEL-

TEMA ET AL., 1994), to the often sunken

opening of the pericardioducts, and to

the anterior extension beyond this

opening, the glandular ducts do not

appear to be part of the mesodermal

pericardioducts. According to their posi-

tion (see Fig. 2), they would more prob-

ably represent a rostral, internalised

portion of a formerly paired mantle

groove (devoid of mucous tracts).

However, their own lateroventral

(instead of frontal) opening into the

mucous tracts, and the elaboration of

these openings as narrow pores with a

distinct sphincter, makes such an inter-

pretation dubious. This also holds true

for the exceptional configuration in

Prochaetoderma californicum, in which the

mucous tracts form tubes being closed

up fram the medial mantle cavity

(SALVINI-PLAWEN, 1972a: Abb. 27F).

Alternatively, the glandular ducts could

be remains of the original gonoducts

into which the pericardioducts opened

to form a common outlet. This interpre-

tation (HOFFMANN, 1937, SALVINI-

PLAWEN, 1972a) suffers from the condi-

tion that such common outlets in

various bivalves and gastropods are

generally characterised by the openings

of gonoducts into lower pericardioducts

(excretory organ or duct), rather than

pericardioducts into lower gonoducts.

Hermaphroditism is characteristic

for Solenogastres and Gastropoda-

Euthyneura, though it also occurs in

other molluscan groups (not known in

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

MEE

Figure 9. Preoral trochi (prototrochs) in Bivalvia. A: pericalymma larva of Yoldia limatula with

three trochi (from DREW, 1901); B stenocalymma larva of Mactra chinensis (ventral view) with

three trochi (from MEDVEDEVA AND MALAKHOV, 1983); C pseudotrochophora (praeveliger) larva

of Pandora inaequivalis (lateral view) with two trochi (after ALLEN, 1961).

Figura 9. Cilios preorales (prototrocas) en Bivalvia. A: larva pericalymma de Yoldia limatula con 3

hileras de cilios (de DREW, 1901); B: larva estenocalymma de Mactra chinensis (vista ventral) con tres

hileras de cilios (tomado de MEDVEDEVA Y MALAKHOY, 1983); C: larva pseudotrocófora (preveliger) de

Pandora inaequivalis (vista lateral) con dos hileras de cilios (de ALLEN, 1961).

Caudofoveata and Siphonopoda/

cephalopods). Among organisms with

separate sexes, free external fertilisation

in the open water appears to be primi-

tive, conservatively also correlated with

so-called ect-aquasperms (“head” of

short nucleus with conical acrosomal

vesicle and 4-5 round proximal mito-

chondria, a diplosome and abruptly

separated flagellum). Internal fertilisa-

tion implies alteration of sperm organi-

sation and shape (introsperm). Once

specialised, no reversal to aquasperm is

possible. This also holds true if separate

sexes are secondarily acquired; a case in

point are the Microhedylidae (Gas-

tropoda-Opisthobranchia-Acochlid-

iomorpha), which use spermatophores

(summarised by E.Wawra in ARNAUD,

POIZAT AND SALVINI-PLAWEN, 1986).

Caudofoveata and Placophora possess

somewhat modified aquasperms,

whereas the hermaphroditic Solenogas-

tres exhibit highly derived introsperms

(SCHELTEMA ET AL., 1994, HEALY, SCHAE-

FER AND HASZPRUNAR, 1995). As evi-

denced by the outgroup comparison (cf.

HEALY ET AL., 1995), the hermaphro-

ditism and introsperms of Solenogastres

clearly appear to be autapomorphies for

this class.

The larval development of lower

groups within molluscan classes (apart

from Siphonopoda/cephalopods) in-

cludes lecithotrophic trochus larvae

(SALVINI-PLAWEN, 1980, 1991; HAsz-

PRUNAR, SALVINI-PLAWEN AND RIEGER,

1995). Among these, Solenogastres show

the pericalymma and stenocalymma

types (SALVINI-PLAWEN, 1991, OKUSU,

2002) and Caudofoveata the stenoca-

lymma type (SALVINI-PLAWEN, 1991,

NIELSEN, 1995). Both these types are also

present in lower Bivalvia (Fig. 9), and

stenocalymma in Scaphopoda, whereas

the larvae of Placophora and most Ar-

chaeogastropoda (without larval shell)

possess the pseudotrochophora or

praeveliger type only (FIORONI, 1982,

SALVINI-PLAWEN, 1991, HASZPRUNAR ET

Iberus, 21 (1), 2003

AL., 1995). Larvae, however, are ecologi-

cally-conditioned and many are pro-

vided with a trochus or other similar

features. Unfortunately, this had led

such larvae to be inaccurately termed

“trochophorae” (ROUSE, 1999). Not only

is this in contrast to the original defini-

tion (Harschek, 1891), but it disregards

the detailed morphological differences

necessary for homologisation (SALVINI-

PLAWEN, 1980), ¡.e. neglecting analogies

(see also HASZPRUNAR, 1995). Note that

all known Solenogastres larvae possess

one preoral trochus only. In constrast,

other free-living pericalymma and

stenocalymma types of Caudofoveata,

Bivalvia and Scaphopoda exhibit three

preoral trochi (NIELSEN, 1995, OkUSU,

2002). Based on the sequence of perica-

lymma (Nematomenia, Neomentia, etc.) to

stenocalymma (Epimenia) in Solenogas-

tres and Bivalvia (Fig. 9), and even an

individual transition from pericalymma

to stenocalymma in Epimenia

(Solenogastres-Cavibelonia; OKUSU,

2002), the pericalymma type appears to

be the conservative one (rather than the

pseudotrochophora type). The outgroup

comparison (SALVINI-PLAWEN, 1980,

HASZPRUNAR ET AL., 1995) supports this

view.

There are only few, more general de-

velopmental coincidences between Mol-

lusca and Coelomata (spiral cleavage,

endomesoblast 4d, trochus larvae of

pericalymma rather than “trochophora”

type; SALVINI-PLAWEN, 1988b); such co-

incidences do not pertain, however, to

the configuration of the body cavity.The

formerly heavily discussed theory of a

coelomate nature and metameric seg-

mentation in Mollusca, renewed with

the one-sided interpretation of Trybidia

organisation (LEMCHE AND WINGSTRAND

1959), is now obsolete (SALVINI-PLAWEN,

1969, 198la, 1988b, SALVINI-PLAWEN

AND BARTOLOMAEUS, 1995, HASZPRUNAR

AND SCHAEFER 1997a, HASZPRUNAR AND

WANNINGER, 2000). The dorsoventral

musculature in Placophora, arranged in

16 pairs of bundles, ontogenetically dif-

ferentiates from a multiple serial condi-

tion similar to that in recent Solenogas-

tres, and the anterior (pre-trochal) mus-

culature recapitulates a “worm-grid”

(HASZPRUNAR AND WANNINGER, 2000).

Moreover, the developmental fate of the

(endo-) mesoblastema in Placophora

(musculature, pericardium) and spi-

ralian Coelomata (secondary body cav-

ity, musculature) is distinctly different

(SALVINI-PLAWEN AND BARTOLOMAEUS,

1995), negating any mutual dependence

of the derivates. Underlined by the or-

ganisation of Solenogastres (SALVINI-

PLAWEN AND BARTOLOMAEUS, 1995), the

molluscan forerunners most logically

originated from epifaunal, ciliary-glid-

ing mesenchymate to pseudo-coelomate

Spiralia (TRUEMAN, 1976).

With regard to the plesiomorphic

conditions of Mollusca, the comparative

consideration of the two diphyletic

recent taxa of -paraphyletic *Apla-

cophora* clearly helps to outline the

probable original organisation of mol-

luscs (archimolluscs). This, in turn, pro-

vides a common base for understanding

the subsequent phylogenetic radiation.

The plesiomorphies include (comp. also

SALVINI-PLAWEN, 1972a, 198la, 1985,

1991, HASZPRUnar, 1992):

e A dorsoventrally constructed body

with separation into a ventral locomo-

tory surface in the form of a ciliary

gliding sole (foot) and a dorsal protec-

tive surface (mantle); the latter secretes a

chitinous cuticle as well as unicellularly

formed, squamous calcareous sclerites

(aragonitic scales).

e Foot and mantle at least in poste-

rior part of body separated by a peri-

pedal groove (pallial cavity) with body

outlets, high glandular mucous tracts

and (?) one pair of ctenidia.

e A paired chemoreceptive (osphra-

dial) sense organ between the terminal

mantle rim and pallial cavity, inner-

vated from the suprarectal commissure.

e An amphineurous nervous system

consisting of a paired cerebral ganglion,

buccal ring and two pairs of medullary

body nerve cords irregularly intercon-

nected by ventral (pedal) commissures

and lateroventral connectives; lateral

cords with suprarectal commissure.

e A subintegumental musculature

below the mantle consisting of outer-

SALVINI-PLAWEN: Significance of aplacophoran Mollusca

transverse, intermediate oblique and

inner longitudinal fibres, the latter rein-

forced lateroventrally below the lateral

rim of the mantle.

e Without segmentation, but with

serially arranged dorsoventral muscula-

ture of two pairs of bundles between

mantle and foot, the dorsal-outer ones

intercrossing medioventrally.

e An alimentary tract only subdi-

vided into foregut and body-filling

midgut without separate gland organ(s).

e A monoserial radula formed of

serially-midventrally chitinised pharyn-

geal cuticle and consisting of firm or

flexible plates, at both sides each with

lateral cusp, denticle or hook; not yet

differentiated radula membrane

(ribbon).

e Associated with the radula, a

paired glandular organ of subepithelial

follicles (without paired duct?).

e A suprarectal pericardium with

middorsal heart invagination having a

single ventricle and a paired auricle

(atrium), the latter also performing

ultrafiltration (for the primary urine).

e A mesenchymate-pseudocoelo-

mate body cavity with an open circula-

tory system without aorta.

e Paired pericardioducts without

excretory function (no formation of sec-

ondary urine). |

e A paired gonad differentiated and

separated as a rostral enlargement of the

pericardium (gono-pericardium); type

of gonadial outlets not yet clarified.

e Sexes separate, free fertilisation by

means of aquasperms.

e Spiral cleavage

* Indirect development through leci-

thotrophic trochus larvae of perica-

lymma type.

e Epibenthic carnivores feeding on

more or less immobile, soft-bodied prey.

CONCLUSIONS

A serious, comparative considera-

tion of the aplacophoran molluscs, de-

spite the limited state of knowledge, re-

veals much important information, par-

ticularly with respect to the

phylogenetic discussion. Their organisa-

tion, especially the different elabora-

tions of mantle cavity organs, clearly re-

flects the presence of two distinct taxa,

the Solenogastres (neomeniomorphs)

and the Caudofoveata (chaetodermo-

morphs). They evolved as diphyletic

clades from the common molluscan bed

(archimolluscan stem group), which

clearly belonged to the aplacophoran

level. Thus, *Aplacophora* is para-

phyletic, also including the common

molluscan ancestors, and the coinci-

dences between the diphyletic

Solenogastres and Caudofoveata appear

to reflect mere symplesiomorphies. The

relative position of both these taxa to the

Testaria, however, still remains to be

clarified. In any case, characters of the

recent Solenogastres and Caudofoveata

must be taken into consideration when

seriously discussing molluscan ple-

siomorphies and the archimolluscan or-

ganisation.

The characters treated above point to

originally mesenchymate-pseudocoelo-

mate organisms with epifaunal, ciliary

locomotion; these ancestors most proba-

bly nourished themselves as carnivores

on soft-bodied, fairly sessile prey (rather

than having been algae-scraping herbi-

vores). Such an acoelomate, ciliary-

gliding organisation clearly points to an

origin from a likewise ciliary-gliding

group of organisms with spiral cleavage

(possibly close to kamptozoan

tholophora larvae; SALVINI-PLAWEN,

1980; HASZPRUNAR ET AL., 1995, HASZ-

PRUNAR, 1996, 2000); these organisations

subsequently differentiated molluscan

characters (dorsal protective cover,

mantle cavity, heart within pericardium,

radula). There is no indication at all that

the molluscan ancestors had a coelo-

mate organisation (i. e. with secondary

body cavity). The pericardium and sub-

sequent gono-pericardium appears to be

an autapomorphic adaptation for the

new heart, which served for body-fluid

circulation and ultrafiltration, i.e. to

compensate a restricted distribution of

metabolites due to the novel mantle

cover and the concentration of gas

exchange at the posterior body (mantle

Iberus, 21 (1), 2003

cavity). In contrast, the common stock of

related coelomate Spiralia most proba-

bly originated as a sister-group by

acquiring a secondary body cavity:

emerging from the same acoelomate

ciliary-gliding forerunners, they

adapted to an infaunal, burrowing habit

(CLARK, 1964). The rounding up of the

body yielded a worm-shaped form with

loss of ciliation, which was subse-

quently followed by the autapomorphic

formation of a body-filling hydrostatic

coelom, i.e. the secondary body cavity.

(Among such monophyletically evolved

coelomate Spiralia, the sipunculids with

their benthosphaera larvae may perhaps

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577

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O Sociedad Española de Malacología

Iberus, 21 (1) 9-104, 2003

The genus Pelycidion (Mollusca: Archaegastropoda) in West

Africa

El género Pelycidion (Mollusca: Archaegastropoda) en Africa occi-

dental

Emilio ROLÁN* and Peter RYALL**

Recibido el 2-V-2002. Aceptado el 20-XI1-2002

ABSTRACT

West African shells of the genus Pelycidion [family Pelycidiidae Ponder and Hall, 1983)

are studied, photographed and compared; the conclusion reached is that only one vari-

able species is present in this area. Some characteristics of the shell of this species are

illustrated.

RESUMEN

Las conchas de Africa Occidental pertenecientes al género Pelycidion [familia Pelycidii-

dae Ponder and Hall, 1983) son estudiadas, fotografiadas y comparadas, llegándose a

la conclusión de que, en esta área, sólo existe una especie morfológicamente muy varia-

ble. Se muestran y comentan algunos caracteres de la concha.

KEY WORDS: Pelycidion, West Africa, shell characters.

PALABRAS CLAVE: Pelycidion, África occidental, caracteres de la concha.

INTRODUCTION

FOLIN AND PÉRIER (1873) described

from Hong-Kong and Senegal the first

known species of the family Pelycidi-

idae Ponder and Hall, 1983, which was

named Pelycidion venustulum Fischer.

PONDER AND HALL (1983) proposed

the family name Pelycidiidae, with Pely-

cidion venustulum as its type species; the

type locality is restricted to Senegal.

This work also mentions the generic

synonyms and lists the previously

known species of this genus, illustrating

most of them.

The position of this family in this

last work is not clear but it is tentatively

placed in Archaegastropoda. Later,

* Cánovas del Castillo, 22, E-36202 Vigo, Spain.

** St, Ulrich 16, A-9161 María Rain, Austria.

BOUCHET AND LE RENARD (1998) placed

it near Pickworthiidae; even the possi-

bility that it may be a synonym of this

family is mentioned (as a personal com-

munication from W. F Ponder).

OKUTANI (2000) places the family Pely-

cidionidae (sic!) between the families

Pickworthiidae and Cingulopsidae.

Two species from West Africa are

mentioned in these works:

-Pelycidion venustulum, represented

in PONDER AND HALL (1983, figs. 1F and

1G) showing shells from Mauritania.

-Pelycidion sp., represented in

PONDER AND HALL (1983, figs. 1E and

3B-3E) showing shells from Dahomey.

Iberus, 21 (1), 2003

MATERIAL AND METHODS

In sediment samples dredged by the

authors in Ghana, several shells of this

genus were found. This material was

noted to be complementary with more

shells from Senegal (J. Pelorce coll.) and

Angola (F. Fernandes coll.). In the

present work, the authors have made

comparison of the shell characteristics

(protoconch, size, sculpture of the last

RESULTS

whorl and microsculpture) of all the

material obtained, comparing it with

those in the above-mentioned literature.

AU the material studied is in collection

of the first author (CER).

Abbreviations:

s Shell

f fragment

] juvenile

Genus Pelycidion Fischer in Folin and Périer, 1873

Type species: Pelycidion venustulum Fischer, 1873. For monotypy.

For synonyms and general characters see PONDER AND HALL (1983).

Pelycidion venustulum Fischer, 1873 |

Pelycidion venustulum Fischer, in Folin and Périer, 1873. Les fonds de la mer.: 182.

Type material: Untraceable.

Material examined: Senegal: 1 specimen, 2s, 1 f, Petit Thouriba, Dakar, 35 m;2s, Grand Thiouriba,

Dakar, 33 m ; 18 s, Madeleines, Dakar, 6-14 m ; 38 s, Madeleines, Dakar, 18 m ; 23 s, Madeleines,

Dakar, 6-14 m;15s, Cap Vert, Epave “Tacoma”, 15 m;15s, Tiwa, Dakar, 35 m . Ghana: 3 s, Miamia,

12m;31s, 6 j, 10 f, Miamia, 35-40 m ; 14 s, Miamia, 45-50 m ; 34 s, 5 f, Cape Three Points, 35-65 m.

Angola: 2 s, Corimba, Luanda, 20 m; 1 s, off Luanda, 60-100 m; 1 s, Namibe, 25m.

Description: Shell (Figs. 1-11) minute,

elongate-pupoid, relatively solid,

whitish or cream-coloured. Protoconch

(Figs. 12-17) brownish in fresh shells,

with about 2 whorls, with a smooth

beginning of almost 1 whorl; the subse-

quent part exhibits an evident sculpture;

in the subsutural area, this is formed at

the beginning by an oblique reticulation

and at the end by axial ribs; in the

middle of the whorl, there are fine spiral

threads, being adapically stronger. At

the end of the protoconch, there is a pro-

nounced subsutural sinus. The teleo-

conch has between 3 */2- 4 */2 whorls

which increase in size gradually being

hardly convex or almost flat. Suture

impressed. The last whorl is large, rep-

resenting almost */2 of the total heigth

of the shell. Some shells indicate a last

whorl uniformly curved towards the

periphery, while others show a strong

100

spiral cord which begins at the end of

the suture and forms a peripheral keel.

The aperture is circular or subcircular

and wweakly thickened externally

towards the apical part. Peristome

simple.

The microsculpture (Figs. 18-23) is

hightly variable; sometimes, the shell

appears to be smooth and only under

magnification the regular spiral striae,

which are formed by one or two rows of

small indentations separated by larger

interspaces, can be observed. In other

cases, these rows of pits are separated

by similar interspaces and appear as a

continuos sculpture. In some shells,

stronger striae appear and the pits or

indentations may or may not be visible.

Even in this last case, one shell we

observed exhibits on the first whorls the

classical pitted sculpture and on the last

whorl this other striated sculpture.

ROLÁN AND RYALL: The genus Pelycidion (Mollusca: Archaegastropoda) in West Africa

Figures 1-11. Shells of 2 venustulum. 1: 1.4 mm, Madeleines, Senegal; 2-4: 1.3, 1.3 and 1.3 mm,

off Luanda, 60-120 m, Angola; 5-11: 1.8, 1.3, 1.4, 1.1, 1.4, 1.1 and 1.3 mm, Miamia, Ghana.

Figuras 1-11. Conchas de P. venustulum. 1: 1,4 mm, Madeleines, Senegal; 2-4: 1,3, 1,3 y 1,3 mm,

fuera de Luanda, 60-120 m, Angola; 5-11: 1,8, 1,3, 14, 1,1, 14, 1,1 y 1,3 mm, Miamia, Ghana.

101

Iberus, 21 (1), 2003

50 um

Figures 12-17. Protoconchs of P venustulum. 12: Madeleines, Dakar, Senegal, 35 m; 13: off Luanda,

Angola, 60-100 m; 14-16: Miamia, Ghana, 20-65 m; 17: La Tacoma, Dakar, Senegal, 15 m.

Figuras 12-17. Protoconchas de P. venustulum. 12: Madeleínes, Dakar, Senegal, 35 m; 13: fuera de

Luanda, Angola, 60-100 m; 14-16: Miamia, Ghana, 20-65 m; 17: La Tacoma, Dakar, Senegal, 15 m.

102

ROLÁN AND RYALL: The genus Pelycidion (Mollusca: Archaegastropoda) in West Africa

E —

20 um

20 um 10 um

Figures 18-23. Microsculpture from shells of P venustulum. 18: Senegal; 19, 20: Angola; 21-23.

Ghana.

Figuras 18-23. Microescultura de conchas de P. venustulum. 18: Senegal; 19, 20: Angola; 21-23.

Ghana.

Dimensions: The shell heigth is vari-

able, the smallest shells of our material

measured 1.0 mm, and the largest 1.8

mm; material of both sizes was collected

in the same place.

Operculum (Fig. 24) is multispiral; it

was examined in only one specimen of

Pelycidion from Dakar, with the opercu-

lum visible at the aperture and the soft

parts dry into.

Radula (Figs. 25, 26) which has the

same appearance to that showed in

PONDER AND HALL (1083) from a species

of West America.

- Distribution: The species is known

from Mauritania (mentioned in PONDER

AND HALL, 1983) and Senegal (type

locality) up to Angola.

- Discussion: After the examination of

the material from several countries and

103

Iberus, 21 (1), 2003

Figures 24-26. P venustulum, Dakar, Senegal. 24: Operculum; 25, 26: radula.

Figuras 24-26. P. venustulum, Dakar, Senegal. 24: Opérculo; 25, 26: rádula.

the graphic material shown in PONDER

AND HALL (1983) we have found that

the morphology of this sepcies is very

variable and there are intergradation

between the different sized shells as

well as in the sculpture of the last whorl

and microsculpture. For these reasons

we conclude that only one species exists

on the West African coast which we

illustrate in this work from several

aspects to show the variability.

BIBLIOGRAPHY

BOUCHET, P. AND LE RENARD, J. 1998. Family

Pickworthiidae, pp. 739-741. In Beesley, P. L.,

Ross, G. J. B. and Wells, A. (Eds.). Mollusca:

The Southern Systhesis. Fauna of Australia, vol

5. CSIRO Publishing, Melbourne, Part B.,

565-1234.

FOLIN, L. DE AND PÉRIER, L. 1867-1886. Les fonds

de la mer, étude internationale sur les particula-

rités nouvelles des régions sous-marins. 4 vol. Pa-

ris.

104

ACKNOWLEDGEMENTS

Our thanks to J. Pelorce for the loan

of material; to Jesús Méndez of the

CACTI of the University of Vigo for the

SEM photographs; also to the PARSYST

project for our estance in the MNHN of

Paris, and the bibliographic information.

This work has been partially sup-

ported by a grant of XUNTA DE

GALICIA PGIDTOOPXI30121PR.

OKUTANI, T. (ed.). 2000. Marine mollusks in Ja-

pan. Tokai University Press, Tokyo. 1173 pp.

PONDER, W. F. AND HALL, S. J. 1983. Pelycidii-

dae, a new family of Archaegastropod mo-

lluscs. The Nautilus, 97 (1): 30-35.

O Sociedad Española de Malacología —_—_—_—_——T— lberus, 21 (1); 105-113, 2003

El conducto excretor de la glándula de la cápsula de Bolinus

brandaris (Gastropoda Prosobranchia): estudio estructural

y ultraestructural

Structural and ultrastructural study of the excretor duct of the cap-

sule gland of Bolinus brandaris (Gastropoda, Prosobranchia)

Ma José AMOR PÉREZ*, Montserrat RAMÓN** y Merce DURFORT I COLL*

Recibido el 4-X-2002. Aceptado el 11-1-2003

RESUMEN

El conducto excretor de la glándula de la cápsula de Bolinus brandaris recorre la glándula

en toda su longitud y está tapizado por un epitelio prismático ciliado y con microvilli que lo

separa de los agregados glandulares adyacentes. Núcleo en posición basal, la heterocro-

matina se distribuye dispersa en el nucleoplasma, y en una banda periférica. El citoplasma

presenta abundante glucógeno y gran cantidad de formaciones vesículo-membranosas; hay

un gran desarrollo del complejo de Golgi y lisosomas, así como formaciones semejantes a

figuras mielínicas, de naturaleza glucídica. El retículo endoplasmático está escasamente desa-

rrollado. La presencia de cilios, microvilli e invaginaciones de la membrana basal corres-

ponde a células con función de reabsorción, transporte y difusión de compuestos, las vesí-

culas de secreción suponen además una función secretora. Se supone que estas células sin-

tetizarían materias primas procedentes de las células glandulares vecinas. Esta hipótesis se

basa no sólo en el gran desarrollo de los sistemas endomembranosos, sino también en la

presencia de vesículas similares a las presentes en las células glandulares vecinas.

ABSTRACT

The excretory duct of the capsule gland of Bolinus brandaris runs along the gland and is co-

vered by a columnar epithelium with microvilli and cilia which separates it from the adjacent

gland complexes. Nucleus located in the base of the cell. Scattered drops of heterochroma-

tin could be seen in the nucleoplasm. An heterochromatin band was also found around the

periphery. The cytoplasm was homogeneously full of glycogen. Vesicle-membranous forma-

tions were well developed (especially, a Golgi body). Associated to the latter were abundant

vesicles, lysosomes, and glucidic myelin-like bodies. The endoplasmic reticulum was not well

developed. The presence of cilia, microvilli and basal membrane invaginations ¡is typical of

cells dedicated to transport, resorption and difussion; the highly- developed membranes also

suggest a secretion role. lt is possible that material previously synthesized in neighbouring

gland cells could be transported for further elaboration in these cells. The presence of vesi-

cles similar to those found in neighbouring gland cells and the well- developed endomem-

branous structures support this hypothesis.

PALABRAS CLAVE: Aparato reproductor femenino, conducto excretor, glándula de la cápsula, epitelio vibrá-

til, endomembranas. ;

KEY WORDS: Female reproductive system, excretory duct, capsule gland, vibratile epithelium, endomembranes.

* Departamento de Biología Celular. Facultad de Biología. Universidad de Barcelona. Avda. Diagonal 645.

08028. Barcelona.

** Institut de Ciencies del Mar (CSIC). Pg. Marítim 37. 08003. Barcelona

105

Iberus, 21 (1), 2003

INTRODUCCIÓN

El aparato reproductor de los molus-

cos presenta diversos grados de comple-

jidad en base al tipo de fecundación: si es

externa el sistema reproductor es muy

sencillo, si es interna es altamente com-

plejo (JONG-BRINK, BOER Y JOOSSE, 1983).

En los moluscos con fecundación interna

se describen tres tipos de glándulas acce-

sorias: la glándula de la albúmina, res-

ponsable de la formación de las cubiertas

oocitarias; glándula ingestiva (receptá-

culo seminal), donde tiene lugar la

fecundación y posterior degradación de

los espermatozoides sobrantes o defec-

tuosos y la glándula de la cápsula,

donde los huevos fecundados son empa-

quetados definitivamente para la puesta.

Estas tres glándulas están comunicadas

entre sí y surcadas por un conducto (ovi-

ducto) que, partiendo del ovario las atra-

viesa. El conducto excretor de la glán-

dula de la cápsula es por tanto una doble

vía: de entrada de los espermatozoides,

y de salida de los huevos (FETTER, 1941;

HYMAN, 1967; GRASSÉ, 1968; OEHLMANN,

STROBEN Y FIORONI, 1988; SCHULTE-

OEHLMANN, FIORONI, (OELMANN Y

STROBEN, 1994 y OEHLMANN, FIORONI,

STROBEN Y MARKERT, 1996).

Al ser interna la fecundación de

Bolinus brandaris, estos tres tipos glandu-

lares se hallan presentes en su aparato

reproductor femenino. Los aspectos

estructurales y ultraestructurales del

conducto excretor de la glándula de la

cápsula, son objeto del presente trabajo.

MATERIALES Y MÉTODOS

Glándulas de la cápsula de hembras

de Bolinus brandaris, procedentes de di-

versos puntos de la costa catalana,

(principalmente Sant Carles de la Ra-

pita, Tarragona), fueron extraídas y pro-

cesadas para su estudio en microscopía

Óptica y electrónica.

Para el estudio en microscopía

óptica, glándulas enteras fueron fijadas

en formol 10%, e incluidas en parafina.

Cortes de 6 pm de grosor, fueron

teñidos con las técnicas de.: hematoxi-

106

lina-eosina, Mallory, y la técnica histo-

química del ácido periódico de Schiff

(PAS) específica para la detección de

carbohidratos.

Para la observación en microscopía

electrónica, pequeñas secciones de glán-

dulas(lmm de grosor), fueron procesa-

das con la técnica convencional de la

doble fijación glutaraldehído 2,5%-para-

formaldehído 3,5% y tetróxido de osmio

al 2%, tamponados con tampón fosfato

(Sorensen). Tras una deshidratación pro-

gresiva, fueron incluidas en resina Spurr.

Cortes semifinos (1 ym de grosor) a fin

de seleccionar el área donde realizar los

cortes ultrafinos (30 nm) se tiñeron con

azul de metileno-bórax. Los cortes ultra-

finos fueron realizados con un ultrami-

crótomo Reichert-Omu provisto de

cuchilla de diamante. Tras ser recogidos

con rejillas de cobre, se procedió al doble

contrastado con acetato de uranilo y

citrato de plomo. En los casos que se

realizó la técnica citoquímica de Thiery,

detectora de carbohidratos, las muestras

fueron recogidas con rejillas de oro y no

contrastadas posteriormente. Las obser-

vaciones se realizaron con los microsco-

pios electrónicos Philips 301 y Hitachi

600, de los Servicios Científico-Técnicos

de la Universidad de Barcelona.

RESULTADOS Y DISCUSIÓN

La glándula de la cápsula de Bolinus

brandaris, tal como está descrito en otras

especies (HYMAN, 1967; GRASSÉ, 1968 Y

OEHLMANN ET AL.,1988) es la mayor de

las glándulas del tracto genital feme-

nino. Macroscópicamente, la glándula

de la cápsula de Bolinus brandaris pre-

senta morfología redondeada, alargán-

dose ligeramente en sus extremos. Exte-

riormente es de color blanco—ocre, y está

dividida en dos lóbulos, derecho e

izquierdo, semejante a las descripciones

de FETTER (1941) y OEHLMANN ET AL.,

1996. Muy desarrollada en la época de

madurez sexual (primavera principal-

mente), en épocas de reposo (resto del

año), está prácticamente atrofiada.

AMOR: Estudio del conducto excretor de la glándula de la cápsula de Bolinus brandaris

Figura 1. Detección de la glándula de la cápsula a nivel de la luz (L) del conducto excretor. Puede

observarse cómo las células del epitelio ciliado (C) son invadidas por material basófilo (Flecha) similar

al contenido en las glándulas adyacentes. Hematoxilina- eosina. Figura 2. Zona colindante entre dos

áreas glandulares diferentes (G1 y G2): Mientras Gl es altamente eosinófila, G2 presenta una clara

basofilia. Obsérvese como los epitelios ciliados correspondientes a estas áreas (C1 y C2), presentan

afinidades tintoriales diferentes entre sí, y similares a la de sus glándulas adyacentes. Hematoxilina-

eosina. Figura 3. Panorámica ultraestructural del epitelio ciliado del conducto excretor donde se apre-

cian los núcleos (N), los cilios (C) así como múltiples formaciones membranosas (asterisco) . En la

luz del conducto (L) pueden apreciarse vesículas de diverso tamaño y electrodensidad (v).

Figure 1. The excretory duct of the capsule gland showing its lumen (L). Ciliate cells (C) are invaded by

basophilic material (arrow). See its similarity of this material with that inside of the neighbouring cells.

Hematoxilin-eosin. Figure 2. Detail of two different areas of glandular masses. While G1 has eosinophil

pH, G2 has intense basophilia. The ciliate epithelia next to these areas (C1 and C2) have varying stain-

ing properties. However, these properties are similar to those of their neighbouring glands. Hematoxilin-

eosín. Figure 3. Ultrastructural observation of the ciliated epithelium of the excretor duct. Observe nu-

clei (N), cilia (C) as well as several membranous formations (asterisk). Vesicles (v) of different size and

electrondensity appear in the lumen (L).

A microscopía óptica, y tal como se

observa en varias especies (HYMANN,

1967; GRASSÉ, 1968; SHULTE-OEHLMANN

ET AL., 1994; FETTER Y GRAHAM, 1994;

OEHLMANN ET AL., 1996) se observa que

ambos lóbulos están formados por masas

glandulares de diferente naturaleza, que

se ponen en evidencia por su diferente

afinidad tintorial, cuyo contenido es

excretado a un conducto irregular que

surca a la glándula en toda su longitud

(Figs. 1, 2). Este conducto excretor está

tapizado por un epitelio prismático

ciliado (75 qm de altura) y con microvilli

como se observa con el microscopio elec-

trónico (Figs. 3-5). La afinidad tintorial de

estas células es similar a la de las células

glandulares vecinas.

La imagen ultraestructural de este epi-

telio corresponde a la descrita en la

107

Iberus, 21 (1), 2003

Pd e ad E E

Figuras 4-6. Zona apical del epitelio cilia

terísticas: el axonema en secciones transversales (a), la zona de transición (necklace) (T), el corpús-

culo basal (cb) y las raíces ciliares próximas a mitocondrias (m). Asimismo pueden detectarse

microvilli (mi) y desmosomas (flecha). Véase la presencia de un cilio en forma de *discocilio” en la

Figura 6 (asterisco). Figura 7. Detalle de la parte basal celular. Puede observarse la lámina basal

(LB) y las múltiples invaginaciones de la membrana plasmática basal (asterisco).

Figures 4-6. View of the apical area of ciliate epithelia. The typical structure of cilia (C) is apparent, i.e.

transversal sections of the axoneme (a), the transition zone (necklace) (D), the basal body (cb) and the

rootlets besides mitochondria (m). Microvilli (mi) and desmosomes (arrow) are also apparent. Modified

cilia, i.e. disk-cilia, can be seen in Figure 6 (asterisk). Figure 7. View of a basal cell. The basal lamina

(LB) and the multiple invaginations of the basal membrane are visible (asterisk).

mayoría de oviductos, tanto de inverte-

brados como de vertebrados (FETTER, 1941;

SHULTE-OEHLMANN ET AL., 1994; OEHL-

MANN ET AL., 1996; PALMER Y GUILETTE,

1988 y HAGIWARA,1995) mostrando células

ciliadas, con núcleo basal, y gránulos de

electrodensidad variable en su citoplasma

que también se detectan en la luz del con-

ducto (Fig. 3). A mayores aumentos, se

108

observa, como se ha citado anteriormente,

la presencia de cilios y microvilli en su

cara apical (Figs. 4 y 5). Estos cilios tienen

un corpúsculo basal de 0,2 um, que se pro-

longa con la característica raíz ciliar de

aspecto estriado (DENTLER, 1981; DUSTIN,

1984; DENTLER, 1990; AMOR, 1992, 1996 ;

HAGIWARA, 1995) de 0,8 um de longitud

y con una periodicidad de 8 nm. En

AMOR: Estudio del conducto excretor de la glándula de la cápsula de Bolinus brandaris

Figura 8. Sección longitudinal de la zona basal es una célula NO Se observa el a (N)

la lámina basal (LB), múltiples repliegues de la membrana plasmática (asterisco) así como lisoso-

mas (L) y diversos orgánulos vesículo-membranosos (v) dispersos por el citoplasma. Figura 9.

Sección del núcleo rodeado de mitocondrias (m), gránulos de glucógeno (g) y vesículas de diferen-

tes tamaños (v). Figuras 10, 11. Imágenes de las diversas morfologías adoptadas por el complejo de

Golgi (G) en sus proximidades al núcleo (N). Obsérvese asimismo la presencia se glucógeno (g),

mitocondrias (m) y vesículas de diferentes tamaños (v).

Figure 8. Longitudinal section of the basal area of a prismatic cell. See the nucleus (N) and the basal la-

mina (LB) as well as invaginations in the basal plasma membrane (asterisk), lysosomes (L) and vesicle

membranous organelles (v) spread through the cytoplasm. Figure 9. Ultrastructure view of the nucleus

surrounded by mitochondria (m), glycogen granules (g) and difjerent- size vesicles. Figures 10, 11. Di-

verse shapes adopted by the Golgi body (G) near the nucleus (N). Glycogen granules (2), mitochondria

(m) and different size vesicles (v) are also be seen.

sentido ascendente se observa el caracte-

rístico necklace o zona de transición (AMOR,

1996) que acaba en el cilio característico.

el tipo discocilia semejantes al gonoducto

masculino de esta especie. (AMOR, 1992,

1996). Aunque ha sido discutida la real

En un 3% de cilios, aparecen dilataciones

en la parte apical del mismo configurando

existencia de este tipo de cilios con anor-

malidades morfológicas, actualmente está

109

Iberus, 21 (1), 2003

ampliamente aceptada su existencia, sig-

nificando una facilitación en la circulación

de material denso (DURFORT, BOZZO,

POQUET, SAGRISTA, FERRER, GARCÍA

VALERO, AMOR Y RIBES, 1990; CAJARAVI-

LLE, MARIGÓMEZ Y ÁNGULO, 1990, AMOR,

1992, 1996; HAGIWARA, OHWADA, AKOI

AND TANAKA, 2000; LUDIN Y SCHAUDER,

2001). En el caso presente, sería lógica su

presencia ya que por este conducto han

de circular tanto las secreciones segrega-

das como los oocitos en todas sus fases.

Es frecuente la presencia de mitocondrias

junto a estas raíces ciliares que, como ha

sido ya descrito, supone aporte energético

(DURFORT, AMOR, FERRER, GARCÍA VALERO,

POQUET, RIBES Y SAGRISTÁ, 1988; DURFORT,

BOzZzZO, POQUET, SAGRISTA, FERRER,

GARCÍA VALERO, AMOR Y RIBES, 1990 y

AMOR, 1992) (Figs. 4, 5). La longitud de

los cilios varía según el tramo del con-

ducto del que se trate. En el inicio del con-

ducto, es decir la parte próximal al ovario,

los cilios son más cortos (12,5 um) que en

la parte distal del mismo (20 um), lo que

podría facilitar la evacuación de los

huevos encapsulados.

La zona lateral de la membrana plas-

mática, en su parte más cercana a la luz,

presenta las características uniones celu-

lares, se trata de largos desmosomas en

banda, de hasta casi 3 qm de longitud

con la característica morfología de anillo

rodeando dos membranas contiguas

(CLIFFE, 2002) (Figs. 4, 5). No se encuen-

tran sin embargo adyacentes a uniones

septadas como se describe en otros

invertebrados (DUVERT, GROS Y SALAT,

1980; AMOR, 1992; KNOX Y BROWN, 2000)

La zona basal de la membrana plas-

mática, se apoya en la lámina basal, y

presenta múltiples invaginaciones que

invaden gran parte del citoplasma (Fig.

7). Esta morfología de invaginaciones

basales asociada a microvilli corres-

ponde a un patrón celular de un epitelio

de absorción y difusión de productos,

citado por AMOR (1992) en el gonoducto

masculino de esta especie.

El núcleo se encuentra en la zona

basal de la célula. Presenta morfología

oval-redondeada (6,2 im) con un patrón

de heterocromatina dispuesto en grumos

dispersos por el nucleoplasma y una

banda de heterocromatina periférica

(Figs. 3, 8-11). Formaciones de tipo gol-

glano son frecuentes en sus proximida-

des (Figs. 10, 11).

El resto del citoplasma se halla inva-

dido por gránulos de glucógeno, distri-

buido de forma homogénea en el interior

celular, mitocondrias, algunas de las cua-

(Página derecha) Figuras 12-15. Diversas morfologías del comportamiento del complejo de Golgi.

Figura 12. Dictiosomas (flechas) rodeados de pequeñas vesículas de secreción (v), y otras de mayor

tamaño, que por su morfología podrían corresponder a mitocondrias degeneradas (asterisco).

Figuras 13, 14. Dictiosomas (flechas) con gran actividad secretora (v). En la Figura 13 se observa la

presencia de vesículas similares a mitocondrias degeneradas (asterisco) y en la Figura 14, puede

observarse parcialmente una vesícula de alta electrodensidad (asterisco). Figura 15. Dos dictioso-

mas concéntricos (G) rodeados de pequeñas vesículas (v), otras medianas cargadas de material elec-

trodenso (+) y unas mayores (asterisco) de posible origen mitocondrial. Se observan asimismo grá-

nulos de glucógeno (g). Figura 16. En esta imagen se aprecia la presencia de un cuerpo multivesi-

cular (CM) y un lisosoma (L), rodeados de vesículas (v) de variado tamaño. Figura 17. Cuerpo

residual (CR) muy frecuente en este tipo de células.

(Right page) Figures 12-15. Detail of different shapes of the Golgi body. Figure 12. Dyctiosomes

(arrows) surrounded by little secretion vesicles (u). Larger vesicles are also detected (asterisk). Their

morphology suggests they are degenerate mitochondria. Figures 13, 14. Dyctiosomes (arrows) with

intense secretory activity (vu). Note the vesicle-like degenerate mitochondria (asterisk) in Figure 13. A

high electrondensity vesicle is partially visible in Figure 14 (asterisk). Figure 15. Concentric dyctiosomes

(G) surrounded by little vesicles (v). Also median vesicles containing electrondense material (+), large

vesicles possibly derived from degenerate mitochondria (asterisk) and glycogen granules are shown. Figure

16. Multivesicular body (CM) and lysosome (L) surrounded by different-size vesicles (v). Figure 17.

Residual body. Typical of' this kind of cells.

110

AMOR: Estudio del conducto excretor de la glándula de la cápsula de Bolinus brandaris

Iberus, 21 (1), 2003

Figura 18. Formaciones membranosas similares a figuras mielínicas, tratadas con la técnica de

Thiery (asterisco), lo que confirma su naturaleza membranosa y no artefactual.

Figure 18. Thierys technique corroborates that myeline-like figures (asterisk) are membranes, not artifacts.

les parecen presentar pérdida progresiva

de crestas (Figs. 12, 15). El sistema mem-

branoso más desarrollado es el complejo

de Golgi (Figs. 12-15). Este está formado

por múltiples dictiosomas de morfología

muy variada, que se rodean de múltiples

formaciones vesículo-membranosas. Es

frecuente encontrar cuerpos multivesicu-

lares y lisosomas en diversas fases de

formación (Figs. 16, 17), así como forma-

ciones membranosas Thiery positivas,

que recuerdan figuras mielínicas (Fig.

18). El hecho de ser Thiery positivas con-

firma su naturaleza glucídica descar-

tando que se trate de artefactos, hecho

que se da también en ciertas fases de la

formación de las plaquetas vitelinas

(AMOR, RAMÓN Y DURFORT, 2001). Algu-

nas de estas vesículas presentan material

de variable electrodensidad (Figs. 12,

14), que pueden visualizarse también

tanto en las células glandulares vecinas

como en la luz del conducto. Se observa

además la presencia de gránulos muy se-

mejantes tanto por su afinidad tintorial

como por las reacciones histo-citoquími-

cas a los gránulos presentes tanto en la

luz del conducto como en las células

glandulares vecinas. Este hecho podría

m1e2

sugerir una actividad elaboradora por

parte de estas células, donde las materias

primas (productos glandulares sintetiza-

dos en las glándulas vecinas) serían

transferidos a estas células que, me-

diante el complejo de Golgi los converti-

ría en gránulos de secreción. Estos grá-

nulos, serían incorporados a vesículas de

origen golgiano y mitocondrias degene-

radas, a manera de las plaquetas viteli-

nas en los oocitos (AMOR ET AL., 2001). El

hecho del gran desfase entre el alto desa-

rrollo del complejo de Golgi y el escaso

desarrollo del retículo endoplasmático

parece apoyar esta hipótesis.

A diferencia de otros Gasterópodos

(VERMEIRE Y HINCH, 1984), en este con-

ducto no se detectan células caliciformes.

AGRADECIMIENTOS

A David Balayla por su asesora-

miento en la lengua inglesa y a Enric

Valencia por su ayuda en el procesado

digital de las fotografías.

Este proyecto está financiado por el

Centre de Referencia d'Aquicultura de

la Generalitat de Catalunya.

AMOR: Estudio del conducto excretor de la glándula de la cápsula de Bolinus brandaris

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O Sociedad Española de Malacología Iberus, 21 (1)115-127, 2003

Four new Euthría (Mollusca, Buccinidae) from the Cape

Verde archipelago, with comments on the validity of the

genus

Cuatro nuevas Euthria (Mollusca, Buccinidae) del archipiélago de

Cabo Verde con comentarios sobre la validez del género

Emilio ROLÁN* António MONTEIRO** and Koen FRAUSSEN***

Recibido el 2-X11-2002. Aceptado el 22-1-2003

ABSTRACT

Four species, collected in the Cape Verde Islands, are described as new and assigned to

the genus Euthria M. E. Gray, 1830. The new species are compared with other taxa from

the Mediterranean Sea and the Cape Verde Archipelago. The genera Euthria, and Buccin-

ulum are compared and the significance of the differences between them is discussed,

leading to the conclusion that both genera are valid and should be kept separated.

RESUMEN

Se describen cuatro especies nuevas del género Euthria M. E. Gray, 1830 recogidas en

aguas circalitorales del archipiélago de Cabo Verde. Las nuevas especies son compara-

das con otros taxones congenéricos existentes en el Mediterráneo y en el propio archip-

iélago. Recientemente, el género Euthria, únicamente conocido del Atlántico oriental, ha

sido sinonimizado con Buccinulum, que posee especies en el Indo-Pacífico, por lo que se

comparan ambos géneros y se discute el valor de las diferencias entre ellos, considerando

finalmente que ambos son válidos, y deben mantenerse separados.

KEY WORDS: Euthria, Buccinulum, Cape Verde Archipelago, new taxa.

PALABRAS CLAVE: Euthria, Buccinulum, Archipiélago de Cabo Verde, nuevos táxones.

INTRODUCTION

During the last few years, the genus

Euthria Gray, 1850 has frequently been

considered as a synonym of Buccinulum

Deshayes, 1830 (BEETS, 1985, SABELLI,

GIANNUZZI-SAVELLI AND BEDULLI, 1990,

POPPE AND GOTO, 1991, BOUCHET, LE

RENARD AND GOFAS, 2001, CHIARELLI,

2002), but regarding the differences

between both genera we suggest to keep

them separate, as commented upon in

“Remarks” of this paper.

The genus Euthria is represented in

the Mediterranean Sea by a single

species, Euthria cornea (Linné, 1758),

which is rather variable. On the con-

trary, several species have been

described, by DAUTZENBERG AND

FISCHER (1906), CoseEL (1982b), COSEL

* Cánovas del Castillo, 22-5%F, 36202 Vigo, España, emiliorolanCinicia.es

** Largo da Princesa 24-1”E, 1400-303 Lisboa, Portugal, a.j.a.monteiroGOnetcabo.pt

** Leuvensestraat 25, B-3200 Aarschot, Belgium, koen.fraussenOskynet.be

Iberus, 21 (1), 2003

Figure 1. Euthria saharica. Original representation of the holotype (from LOCARD, 1897).

Figura 1. Euthria saharica. Representación original del holotipo (de LOCARD, 1897).

AND BURNAY (1983) and ROLÁN (1985),

for the Cape Verde Archipelago, and a

revision of the genus has been pub-

lished by ROLÁN (1987).

In addition to the species from the

Mediterranean and from Cape Verde,

LOCARD (1897) has described Euthria

saharica (Fig. 1), MARCHE-MARCHARD

AND BREBION (1977) have described

Euthriostoma gliberti, both from the

western coast of Africa. These species

have a more fusiform shell, with an

elongated siphonal channel, that it is we

consider it doubtful that they belong to

the genus Euthria at all. BOUCHET AND

WARÉN (1986) have placed these species

in synonym together with Metzgeria

apodema Bouchet and Talavera 1981, con-

sidering it is not an Euthria. GUERREIRO

AND REINER (2000) show some shells

SYSTEMATICS

belonging to this genus that do not

match the ones previously known for

the archipelago and identify them as E.

cf. saharica and E. cornea. ROLÁN (in

press), in a review of the Cape Verde

gastropod fauna, mentions these refer-

ences, but states that it is unlikely that

the proposed identifications are correct.

The present paper is based on mater-

ial obtained during the last few years by

César Fernandes. This material has been

dredged from deep waters in different

points of the Archipelago.

Abbreviations:

MNCN Museo Nacional de Ciencias

Naturales, Madrid

CCF collection of César Fernandes,

Cascais

CER collection of E. Rolán, Vigo

Genus Euthria M. E. Gray, 1850

Type species: Buccinum corneum Linné, 1758. Recent, Mediterranean. Original designation:

“Fusus lignarius Chiaje” (this is Fusus lignarius Lamarck, 1816, a junior synonym of Murex

corneus Linnaeus, 1758).

Description: GRAY (1850): “Teeth,

central 1 large, lateral 1-1, versatile.

Muzzle rectactile. Operculum annular.

Siphon of mantle produced in front” for

ROLÁN £7 4L.: New Euthria from Cape Verde, with comments on the genus

the family; “Operculum ovate, acute;

nucleus apical. Varices of shell rudimen-

tary or none” for the group to which

belong; and, finally “Canal elongate” for

the genus. THIELE (1935): “Shell spindle-

shaped, with high, ribbed spire; aper-

ture oval; canal in most cases fairly long,

somewhat oblique. Operculum with ter-

minal nucleus”.

Remarks: All Recent Euthria species

have a more or less conspicuous subsu-

tural depression or concavity (E. rolani

von Cosel, 1982, ornamented with a

thick and rather bulbous suture, being

an exception); they have some axial

sculpture on the upper whorls, and lack

well developped spiral sculpture but

occasionally show some fine cords or

striae (the only exception being E.

effendyi [see FRAUSSEN, 2002] - of which

we have no radula available - to ensure

the generic placement, an Indonesian

species which is closely related to the

fossil fauna).

The radula of Euthria, consisting of 3

teeth (1 middle, 2 lateral) with each 3

cusps, is typical for Buccinidae. The

form of the middle tooth (triangular

with the cusps on the broad upper

corner) (IREDALE 1917, COOKE 1917,

SHUTO 1978, ROLÁN 1987), is shared

with Euthriostoma Marche-Marchard

and Brebion, 1977 (Fraussen, unpub-

lished) and _Afer Conrad, 1858

(FRAUSSEN and HADORN 1999).

The radula of Buccinulum is similar

in general aspects but differs by a some-

what broader, lesser triangular central

tooth with more rounded margins

(COOKE 1917). We can base this compari-

son on a limited number of species only,

as a more thoroughfull study of the

radula of all New Zealand species is

beyond the limit of the present paper.

Further study probably can bring more

light on the eventual presence of Euthria

species among the buccinulums of New

Zealand.

The radulae of Siphonofusus KURODA

AND HABE, 1954 and Afrocominella

IREDALE, 1918 are similar too, but differ

by the quadrangular middle tooth with

rather sharp corners (COOKEÉ, 1917,

BARNARD, 1959).

The type species of Buccinulum (B.

lineum Martyn, 1784) (Figs. 29-31), and

consequently other Pacific species

included in the same genus, doubtlessly

have a certain morphological resem-

blance to the Atlantic species of Euthria.

Conchological differences between

Euthria and Buccinulum are rather slight,

but we must recognize the presence of

somewhat prominent axial ribs on the

upper whorls and a clear subsutural

concavity in Euthria. The material of

which the shell of Buccinulum is pro-

duced will be an interesting matter for

further study, as the surface is not as

smooth and brilliant as in Euthria, which

is rather glossy, especially in the Indo-

West Pacific.

We agree with BEETS (1986) to con-

sider the Indonesian archipelago as the

geological center of Buccinulum and

Euthria. In the Recent fauna we can rec-

ognize 4 important geographical areas:

the important radiation of Buccinulum

around New Zealand (differentiated

from typical Euthria already early in

history during the Tethys Sea), an

Euthria radiation along the coasts of the

Indian Ocean, another on the lower

shelf around New Caledonia and,

finally, the East Atlantic Euthria -

Euthriostoma - Afer radiation (separated

from the main group during the split of

the Tethys Sea). The species described as

new in the present paper are an addition

to this latter group.

Regarding the difference in radula,

geographic-geological separation and a

number of slight conchological differ-

ences cited above, we consider the

genera Euthria and Buccinulum as being

different and keep them separate.

Euthria soniae spec. nov. (Figs. 2-7, 23B, 25A, 28E)

Type Material: Holotype (Figs. 2-4) in MNCN (15.05/46582). Paratypes in: CCF (1, Fig. 5), CER (1,

Fig. 6), MNHN (1, Fig. 7).

1417

Iberus, 21 (1), 2003

Figures 2-7. Euthria soniae. 2-4: Holotype, 53.3 mm, Cape Verde archipelago (MNCN); 5:

Paratype, 46.3 mm (CCE). 6: Paratype, 47.2 mm (CER). 7: Paratype, 50.2 mm (MNHN).

Figuras 2-7. Euthria soniae. 2-4: Holotipo, 53,3 mm, archipiélago de Cabo Verde (MNCN); 5:

Paratipo, 46,3 mm (CCE). 6: Paratipo, 47,2 mm (CER). 7: Paratipo, 50,2 mm (MNHN).

118

ROLÁN ET 4L.: New Euthria from Cape Verde, with comments on the genus

Figures 8-12. Euthria marianae. 8-10: Holotype, 36.9 mm, between Sáo Vicente and Sáo Nicolau,

CV (MNCN). 11: Paratype, 33.4 mm (CCE). 12: Paratype, 35.3 mm (CER). Figures 13-15.

Euthria cf. marianae. Concha, 34.0 mm, between Sáo Vicente and Sáo Nicolau, CV (CCE).

Figuras 8-12. Euthria marianae. 8-10: Holotipo, 36,9 mm, entre Sáo Vicente y Sáo Nicolau, CV

(MNCN). 11: Paratipo, 33,4 mm (CCE). 12: Paratipo, 35,3 mm (CER). Figuras 13-15. Euthria cf.

maríanae. Concha, 34.0 mm, entre Sáo Vicente y Sáo Nicolau, CV (CCP).

Iberus, 21 (1), 2003

Type locality: Cape Verde Archipelago. The type material was donated by Capeverdian fishermen

without exact information. Itis however likely that it was captured in “barlavento” group of islands.

Derivatio nominis: The new species is named after biologist Sonia Elsy Merino, who has dedicated

herself to the study and conservation of the marine fauna, working in Porto Mindelo, Sáo Vicente,

Cape Verde Archipelago, and who has cooperated with our research.

Description: Shell (Figs. 2-7) ovoid-

fusiform and solid. The protoconch (Fig.

23B, 25A, 28E) is yellowish, with 1 */4 -

2 spiral whorls and a diameter of about

1.7 mm. Teleoconch with 7 rapidly

increasing whorls. The whole spire pre-

sents axial ribs, 9 - 10 in number on first

whorl, 10-12 on last whorl. Subsutural

ramp strongly depressed. Aperture

ovoid, white, with 12-13 internal folds

visible at the internal part of the peris-

tome; it is continued with a narrow and

slightly curved siphonal canal. Yellow-

ish to brown undulated axial flammules,

sometimes coalescing, and even forming

a reticulated pattern, appear on a white

background.

Holotype 53.3 mm x 21.1 mm. The

paratypes are slightly smaller, the small-

est one being 46.3 mm long.

Soft parts unknown.

Distribution area: The species is

known only from the Cape Verde Archi-

pelago. The short protoconch suggests

an insular endemism.

Comparison: The present species

must be compared with E. cornea, which

has a similar profile, but is sometimes

larger, [easily up to 65 mm, but also

larger: 84,7 mm in F. Pusateri collection,

Palermo, and SETTEPASSI (1971) records

the maximum size known at 88 mm]. E.

cornea has axial nodules on the first 4-5

spiral whorls, rarely on the last one. The

main difference is the protoconch,

which in E. cornea (Figs. 23A, 28A) has a

little more than 1 whorl, with about 0.8

mm, and in E. soniae (Fig. 23B, 28E) has

between 1 */4 and 2 whorls, with about

1.7 mm of diameter.

Euthria marianae spec. nov. (Figs. 8-12, 24B, 28C)

Type material: Holotype (Figs. 8-10) in MNCN (15.05 /46583). Paratypes in: CCF (1, Fig. 11), CER

(1/Fig;: 12).

Type locality: Between Sáo Nicolau and Sáo Vicente, at a depth of 75 - 150 m, in the Cape Verde

Archipelago.

Derivatio nominis: The new species is named after Mariana, the grand daughter of César Fernan-

des - a well-known Portuguese collector who has greatly contributed to the knowledge of the mala-

cological faunas of the Cape Verde Islands and of Mozambique - and the daughter of César Fer-

nandes Jr., a keen diver and shell hunter.

Description: Shell (Figs. 8-12) ovoid-

fusiform and solid. The protoconch

(Figs. 24B, 28C) is pink, with between 1

3/4 and 2 spiral whorls and a diameter

of about 1.5 mm. Teleoconch with about

6 rapidly increasing spiral whorls, with

weak undulating axial ribs, present on

the whole shell, 11 on the first whorl, 16

on the last whorl. Subsutural ramp

slightly depressed, suture nodulous.

Aperture ovoid, white, with about 11

teeth in inner part, continued with inter-

nal folds; the aperture extends into a

narrow, elongated and curved siphonal

canal. Background colour white, with a

120

brown band with white blotches on the

last whorl; this band is visible on previ-

ous whorls on a short part on the supra-

sutural area.

The holotype mesures 36.9 x 19.1

mm. The paratypes are of similar size.

Soft parts unknown.

Distribution area: The species is

known only from locations between 5.

Nicolau e S. Vicente, at 75 - 150 m of

water, taken from traps.

The short protoconch suggests an

insular endemism.

Comparison: The present species must

be compared with E. rolani Cosel, 1982

ROLÁN £7 4L.: New Euthria from Cape Verde, with comments on the genus

and E. boavistensis Cosel, 1982, which

have similar aspects. E. rolani was col-

lected near Santa Luzia in shallow water,

and the shell is more globose and solid,

the aperture is pink-violet instead of

white, the subsutural area is not

depressed, the colour pattern is different,

with wide blotches, and the protoconch is

wider and dark brown (Figs. 24A, 27,

28B), about 2.1 mm in diameter. E. boavis-

tensis has a smaller shell, with a different,

more irregular, colour pattern, the aper-

ture constantly pink and the protoconch

a little smaller (about 1.2 mm diameter).

Euthria fernandesi spec. nov. (Figs. 16-18, 28B, 28D)

Type material: Holotype (Figs. 16-18) deposited in MNCN (15.05 /46584).

Type locality: Dredged between the islands of Maio and Boavista, at 75 - 125 m, Cape Verde Arch-

ipelago.

Derivatio nominis: The new species is named after the well-known Portuguese shell collector César

Fernandes.

Description: Shell (Figs. 16-18) ovoid-

fusiform and solid. The protoconch

(Figs. 25B, 28D) is white, with a little

more than 2 spiral whorls and a diame-

ter of 2.2 mm. Teleoconch with 6 rapidly

increasing spiral whorls, with axial ribs

on the whole shell, 10 on the first whorl,

16 on the last whorl; these ribs are

present at the suture, giving it a nodu-

lous aspect. Subsutural ramp strongly

depressed. Aperture ovoid, white, with

13 folds on its outer lip; the aperture

extends into a narrow, slightly elon-

gated and dorsally curved siphonal

canal. The background colour is irregu-

lar light brown, with numerous small

axial white lines; there is a suprasutural

broad dark brown spiral band, inter-

rupted by white blotches, and other

similar but less conspicuous bands

towards the anterior end of the shell.

The holotype is 51.3 mm x 22.6 mm.

Soft parts unknown.

Distribution area: The species is only

known from the type locality

The short protoconch suggests an

insular endemism.

Comparison: Only the holotype is

available, but we decide to describe this

species, because of important differ-

ences with other congeneric species.

E. fernandesí spec. nov. must be com-

pared with E. cornea, and it may be sep-

arated because the latter species does

not have the conspicuous coloured

bands, and the last whorls usually lack

axial and spiral sculpture. Furthermore,

the protoconch has only one whorl, it is

cream or light brown, and its diameter

is smaller (only 0.8 mm in diameter,

versus 2.1 mm in the new species).

E. calypso Cosel and Burnay, 1983 has

a similar size, but the shell pattern is dif-

ferent, with spiral blotches only, more

evident on the suture, and lacking axial

sculpture, except in the first whorls of

the teleoconch. Its spiral sculpture is

formed by spiral threads, which are

evident on the first whorls only, and the

subsutural depression is less evident.

The whorls are less convex.

E. pulicaria Dautzenberg and Fischer,

1906 has a smaller, less elongated shell,

with a small subsutural depression but

prominent suture, a different colour

pattern and a smaller siphonal canal.

E. soniae spec. nov. lacks spiral sculp-

ture, has a more elongated spire, the

colour pattern is formed by axial flam-

mules without any spiral banding, and

the protoconch is smaller and yellowish

(instead of white).

Euthria helenae spec. nov. (Figs. 19-22, 26, 28F)

Type material: Holotype (Figs. 19-21) in MNCN (15.05 /46585). Paratypes in: CCF (1, Fig. 22), CER (1).

Type locality: Between Sáo Nicolau and Sáo Vicente, at 75 - 125 m depth, Cape Verde Archipelago.

121

Iberus, 21 (1), 2003

Figures 16-18. Euthria fernandesi. Holotype, 51.3 mm, between Sáo Nicolau and Sáo Vicente, CV

(MNCN). Figures 19-22. Euthria helenae. 19-21: Holotype, 40.2 mm, between Sáo Nicolau and

Sáo Vicente, CV (MNCN); 22: Paratype, 25.4 mm, (CCE).

Figuras 16-18. Euthria fernandesi. Holotipo, 51,3 mm, entre Sáo Nicolau y Sáo Vicente, CV

(MNCN). Figuras 19-22. Euthria helenae. 19-21: Holotipo, 40,2 mm, entre Sáo Nicolau y Sáo

Vicente, CV (MNCN); 22: Paratipo, 25,4 mm, (CCF).

122

ROLÁN ET AL.: New Euthria from Cape Verde, with comments on the genus

Figures 23-28. Comparison at same magnification between the protoconchs of E. cornea (A) and

E. soniae (B); 24: Comparison at same magnification between the protoconchs of E. rolani (A) and

E. marianae (B); 25: Comparison at same magnification between the protoconchs of E. soniae (A)

and £. fernandesí (B); 26: Protoconch of E. helenae; 27: Larval shells of E. rolaní before hatching;

28: Comparison at same magnification between the protoconchs of E. cornea (A), E. rolani (B), E.

marianae (C), E. fernandesi (D), E. soniae (E) and E. helenae (E).

Figuras 23-28. Comparación al mismo aumento entre las protoconchas de E. cornea (A) y E. soniae

(B); 24: Comparación al mismo aumento entre las protoconchas de E. rolani (4) y E. marianae (B);

25: Comparación al mismo aumento entre las protoconchas de E. soniae (A) y E. fernandesi (B); 26:

Protoconcha de E. helenae; 27: Conchas larvarias de E. rolani antes de la eclosión; 28: Comparación al

mismo aumento entre las protoconchas de E. cornea (A), E. rolani (B), E. marianae (C), E. fernandesi

(D), E. soniae (E) y E. helenae (F).

123

Iberus, 21 (1), 2003

Figures 29-31. Buccinulum lineum, 35.5 mm, New Zealand (CER). Figures 32-36. Euthria cornea.

32: Shell, 51.2 mm, Faro, Algarve, Portugal (CER); 33: Shell, 50.6 mm, Cabo de Palos, Murcia,

Spain (CER); 34: Shell, 37.9 mm, Malta (CER); 35; Shell, 54.7 mm, Sokakagzi, Turkey, 125 m

(CER); 36: Shell, 29.2 mm, Karaade Island, South of Bofrum, Turkey (CER).

Figuras 29-31. Buccinulum lineum, 35,5 mm, Nueva Zelanda (CER). Figuras 32-36. Euthria

cornea. 32: Concha, 51,2 mm, Faro, Algarve, Portugal (CER); 33: Concha, 50,6 mm, Cabo de Palos,

Murcia, Spain (CER); 34: Concha, 37,9 mm, Malta (CER); 35: Concha, 54,7 mm, Sokakagzi,

Turquia, 125 m (CER); 36: Concha, 29,2 mm, isla Karaade, sur de Bofrum, Turquia (CER).

124

ROLÁN ET AL.: New Eutbria from Cape Verde, with comments on the genus

Derivatio nominis: The new species is named after Helena Monteiro, the wife of the second author,

who for many years has endured long hours of malacological discussion at home, long sessions of

shell collecting and bad smelling jars about the house. All shell collectors' wives deserve great tribute.

Description: Shell (Figs. 19-22) ovoid-

fusiform and solid. The protoconch (Figs.

26, 28F) is light brown, with 2 */4 spiral

whorls and a diameter of about 1.6 mm.

Teleoconch with 7 rapidly increasing spi-

ral whorls, with axial ribs on the whole

shell, 9-10 on the first whorl, 12 on the last

one; these ribs do not reach the lower su-

ture and disappear towards the base. The

subsutural ramp is strongly depressed. 4-

5 main spiral threads on the first whorls,

18 on the last one; between them there are

other smaller threads, only visible under

magnification. Aperture ovoid, white, with

11 internal folds at the external part; the

.aperture extends into a narrow slightly

elongated and curved siphonal canal.

Colour whitish cream, except for the pro-

toconch and the first two whorls of the

teleoconch, which are brown. The tip of

the siphonal canal is also dark in the holo-

type. The two paratypes are ornamented

with dark blotches at the base and brown

colour in the spaces between the lower

spiral cords.

REMARKS

As far as the specific separation is con-

cerned, we have had the possibility of

examining large samples of the type

species, E. cornea. This species seems to be

present only in the Mediterranean Sea -

Tanger being the most western location

mentioned by PASTEUR-HUMBERT (1962)

in his study of the fauna from Morocco -

and, to the North, it reaches the Atlantic,

being present at Algarve, on South Por-

tugal (pers. inf.). It is usually found in

shallow waters, but has also been taken

from depths of over 100 m. Within its geo-

graphical range, the species presents a

wide morphological variability (Figs. 32-

36), which even made ROLÁN (1987) con-

sider the possibility that more than one

species could be under hand. However,

and despite the mentioned variability, all

specimens show a short protoconch (Figs.

23A, 28A), with a single spire whorl, of a

Dimensions: Holotype is 40.2 x 17.3

mm.

Soft parts unknown.

Distribution area: The species is known

only from the type locality, collected in

traps.The short protoconch suggests an

insular endemism.

Comparison: The present species has

spiral threads and a deep suture as its

main characteristic, and so we will

compare it with other species with a

similar sculpture.

E. pulicaria has a smallef, more globose

and shorter shell; the suture is prominent.

instead of deep, the subsutural depres-

sion is scarcely noticeable and it is orna-

mented with a brownish banded pattern.

E. adeles Dautzenberg and Fischer,

1906 has a smaller shell, with a shallower

suture, the profile of the last whorl lesser

convex, and a shorter siphonal canal.

E. calderoni has a smaller, narrower,

more elongate shell, with variable

colour, smaller protoconch and variable

apertural colour.

cream colour and a diameter of about 0.8

mm.

Among the species known from Cape

Verde, the great morphological variability

in the shells of E. calderoni has already

been stressed (see ROLÁN, 1987), and it

will be documented in further studies

(ROLÁN, in press). But the examination of

the protoconch and of juveniles - in ages

close to hatching - has shown that a great

intraspecific similitude in the larval shells

exists, as well as a great interspecific sep-

aration, which made comparison with the

newly found species, all of them easy to

distinguish from E. cornea.

So, it is obvious that our knowledge

about this genus in the Cape Verde Islands

has grown a lot in the last few years, from

the three species mentioned by VON COSEL

(1982a) to the ten presently known species.

But some material waiting further study

125

Iberus, 21 (1), 2003

Table I. Morphological characteristics of Euthria species. 1: maximum length for the species, in

mm; 2: axial sculpture on the last whorls, from — to ++; 3: spiral sculpture on the last whorls, from

— to ++; 4: colour of inner aperture; 5: colour of protoconch; 6: diameter of protoconch, in mm;

7: number of whorls in the protoconch.

Tabla I: Características morfológicas de las especies de Euthria. 1: máxima longitud de la especie, en

mm; 2: escultura axial de las últimas vueltas, de

- a ++; 3: escultura espiral de las últimas vueltas, de -

a ++; 4: color de la apertura interna; 5: color de la protoconcha; 6: diámetro de la protoconcha, en mm;

7: número de vueltas de la protoconcha.

l 2 3 4 5 6 /

E.comea 88 + > from white to dark brown light brown 0.8 11/8

E. adeles 30 ++ + whitish brown ? ll

E. pulicaria 36 ++ + white pink ? 1?

E. rolani 4] - - pink to purple brown 2.1 22 1/4

E. boavistensis 31 - - pink to purple pink 172 2

E. calypso 44 - + white ? ? 2

E. calderoni 18 + + colour of shell colour of shell 13 2

E. sonige 53 white yellowish 17 13/42

E. marianae 37 white pink 155 13/42

E. fernandesi 5] ++ ++ white white DE) 2

E. helenae 10 + + white brown 1.6 21)

can involve the discovery of even more

undescribed species.

For an easier grasp of the Euthria

species, we schematically present a few

morphological characteristics, in Table 1.

Although all the species were col-

lected in the Cape Verde Islands, it is

probable that their distribution range

does not include the whole archipelago,

and also that they have different habi-

tats and depth ranges. Table II shows a

summary of this information.

ACKNOWLEDGEMENTS

The authors thank Jesús S. Troncoso,

from the Universidad de Vigo, in whose

Departamento de Ecología the photos of

the present work were made.

Table II. Known distribution area and depth range of the studied species of the genus Euthria.

Tabla II. Área de distribución conocida y rango de profundidad de las especies de Euthria estudiadas.

Known distribution area Depth range

E. .corea Mediterranean and South Portugal shallow water

E. adeles CV, Santa Luzia 52m

E. pulicaria CV, Boavista 91 m

E rolani CV, Santa Luzia shallow water

E. boavistensis CV, Boavista shallow water

E. calypso CV, between Boovista 8. Maio —100 to — 200 m

E. colderoni CV, all of the Archipelago shallow water to — 30 m

E. soniae CV, Barlovento group — 100 m?

E. marianae CV, between Súo Nicolau 8. Súo Vicente —715to—150m

E. fernandesi CV, between Maio 8. Boovista —75t0-125m

E. helenae CV, between Súo Nicolau 8. Súo Vicente —75t0—150m

126

ROLÁN ET AL.: New Euthria from Cape Verde, with comments on the genus

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SHUTO, T., 1978. On the genera Siphonofusus

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Contribución al conocimiento de la fauna malacológica

miocénica de La Pedrera, Sant Llorenc d'Hortons (NE de la

Península Ibérica)

Contribution to the knowledge of the Miocenic malacological fauna

from La Pedrera, Sant Llorenc d"Hortons (NE of the Iberian

Peninsula)

Josep A. MORENO*, Anselmo PEÑAS** y Emilio ROLÁN***

Recibido el 3-VI-2002. Aceptado el 24-1-2003

RESUMEN

Se presenta una lista de 160 especies de moluscos marinos fósiles (138 gasterópodos, 19

bivalvos y 3 escafópodos), recolectados en el yacimiento del Mioceno medio de La Pedrera,

en Sant Llorenc d'Hortons (Barcelona, NE de la Península Ibérica). De estas especies, 40

son citas nuevas para la Península Ibérica y, además, 29 primeras citas para el yacimiento.

Se describen las siguientes nuevas especies: Obtusella orisparvi, Caecum verai, Eulimella

penedesensis, Noemiamea batllorii y Turbonilla olivellai. Proponemos dos nuevos nombres

para Eulimella concinna Sorgefrein, 1958 non Jeffreys, 1884 y Odostomia modesta Sor-

gefrein, 1958 non Stimpson, 1851. Se presta especial atención a los moluscos de pequeño

tamaño que se han estudiado con mayor detalle. Se incluyen asimismo comentarios sobre

algunos de los taxones mencionados y se muestran muchos de ellos al microscopio electró-

nico de barrido (MEB), con especial atención a la familia Pyramidellidae.

ABSTRACT

A list of 160 species of fossil marine molluscs (138 Gastropoda, 19 Bivalvia and 3

Scaphopoda) is presented. They were collected in the middle Miocene deposit of La Pedr-

era, Sant Llorenc d'Hortons (Barcelona, NE of the Iberian Peninsula). From these species,

40 are first record for the Iberian Peninsula, and 29 are first record for this deposit. The

following new species are described: Obtusella orisparvi, Caecum verai, Eulimella

penedesensis, Noemiamea batllorii and Turbonilla olivellai. We propose two new names

for replacing: Eulimella concinna Sorgefrein, 1958 non Jeffreys, 1884 and Odostomia

modesta Sorgefrein, 1958 non Stimpson, 1851. Special attention is paid to the smaller

molluscs, studied detailly. Comments are done on some taxa previously mentioned and

some of them are represented at SEM, mainly those of the Pyramidellidae family.

PALABRAS CLAVE: Moluscos marinos, Mioceno medio, La Pedrera, Sant Lloreng d"Hortons, NE Península

Ibérica, micromoluscos, Pyramidellidae, taxonomía, nuevas especies.

KEY WORDS: Marine molluscs, middle Miocene, La Pedrera, Sant Lloreng d'Hortons, NE Iberian Peninsula,

micromolluscs, Pyramidellidae, taxonomy, new species.

* Carrer Josep Pers i Ricart, 1, 2%. 08800 Vilanova i la Geltrú (Barcelona).

** Carrer Olérdola, 39, 5%. 08800 Vilanova i la Geltrú (Barcelona). ansppGwanadoo.es

*** C/ Cánovas del Castillo, 22, 5. 36202 Vigo (Pontevedra). emiliorolanGinicia.es

129

Iberus, 21 (1), 2003

INTRODUCCIÓN

El presente trabajo trata sobre los

moluscos marinos que los autores han

encontrado en el yacimiento miocénico

del Burdigaliense superior-Langhiense

(BATLLORI Y MARTORELL, 1992) de La

Pedrera, dedicando una especial aten-

ción a los micromoluscos, cuya abun-

dancia ofrece material suficiente para

realizar este trabajo. Por este motivo se

ha utilizado una metodología especial-

mente dedicada a la recolección y

estudio de este material.

Se trata de uno de los yacimientos

más estudiados del Mioceno medio

catalán, desde ALMERA (1896). Entre los

numerosos trabajos realizados sobre este

yacimiento, tanto desde el punto de

vista geológico como paleontológico,

destacamos los de TRUYOLS Y CRUSA-

FONT (1951), CRUSAFONT Y TRUYOLS

(1954), CRUSAFONT (1959), CASANOVAS,

CALZADA Y SANTAFÉ (1972), SALAJ

(1972), MAGNÉ (1978), MARTINELL Y

PORTA (1981), LÓPEZ-CivIT (1984), BAT-

LLORI (1990), BATLLORI Y MARTINELL

(1992), BATLLORI (1995) y NAVAS, MARTI

NELL, DOMENECH, Y BATLLORI (1996).

TRUYOLS 1 CRUSAFONT (1951) hablan

de la sedimentación transgresiva-regre-

siva del Mioceno del Valles-Penedes;

estudian una serie estratigráfica en la

confluencia de la riera Rierussa con el

río Anoia; y consideran que la edad del

yacimiento de la Pedrera es Helve-

ciense. CRUSAFONT (1959) estudia la

denominada segunda fase transgresiva,

atribuyendo La Pedrera al período Tor-

toniense; la presencia de Mytilus y la

ausencia de corales indicaría un

ambiente de tipo estuario con fondos

fangosos; también ofrece una lista de

moluscos marinos; y mantiene que el

yacimiento de La Pedrera es claramente

Tortoniense. CASANOVAS ET AL. (1972)

estudian la litología, sedimentología y

paleontología de dos series de la Rie-

russa; admiten la entrada del mar por el

Besós y que el Garraf sumergido actúa

de barrera y favorece la sedimentación

de elementos finos; y sugieren la exis-

tencia de una estructura tipo deltáica de

edad Tortoniense. SALAJ (1972) en un

130

breve estudio microbioestratigráfico

sobre las series de la Rierussa y La

Pedrera, hechas por TRUYOLS Y CRUSA-

FONT (1951), atribuye las capas basales

de la primera localidad al Aquitaniense

superior y el resto de capas de ambas

localidades las atribuye al Burdigaliense

y Tangiense inferior y superior. MAGNÉ

(1978) ofrece la aportación quizá más

importante en el campo de la cronología

del Mioceno del Valles-Penedes; estudia

los foraminíferos y ostrácodos de nume-

rosas localidades de la Depresión; y

establece una nueva edad para el

Mioceno de esta cuenca en la que los

sedimentos marinos de la Pedrera perte-

necerían al Langhiense. MARTINELL Y

PORTA (1981) comentan la fauna encon-

trada en La Pedrera y basándose en el

hallazgo de Vaginella austriaca atribuyen

este yacimiento al Langhiense-Serrava-

liense inferior; consideran, además, que

debido a la abundante presencia de

Potamididae y Batillarididae el

ambiente de La Pedrera es típico

marino. LÓPEZ-CivIT (1984) estudia los

ostrácodos del Mioceno del Alt Penedés.

BATLLORI (1990) realiza un estudio siste-

mático de los moluscos de La Pedrera.

BATLLORI Y MARTINELL (1992, 1993) estu-

dian la actividad predadora en molus-

cos y consideran, además, en su compa-

ración con el Mioceno salobre de Cer-

danyola del Valles, que los niveles de La

Pedrera corresponden a unas facies

marinas con claras influencias continen-

tales. BATLLORI (1995) abunda especial-

mente en el estudio sistemático sobre los

moluscos gasterópodos marinos del Alt

Penedes. NAVAS ET AL. (1996), en base a

los moluscos fósiles y criterios sedimen-

tológicos, realizan una reconstrucción

paleoambiental del sector de Sant Pau

d'Ordal y de La Pedrera y deducen la

equivalencia lateral de ambas localida-

des, a las que atribuyen una edad Lang-

hiense-Serravaliense inferior. La presen-

cia del opistobranquio Vaginella austriaca

les induce a afirmar que durante este

período existió una buena comunicación

entre el golfo del Penedes y el mar

abierto.

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

Ma PA Ñi

dden__9 HR

< AN

Il ajjr r ag

S Es

Ss ES

E Badalona

Barcelona

ES VHospitalet de Llobr:

Bói de Llobr

St. Llorenc

d'Hostons

Sant Sadurní d'Anoia

Figura 1. Situación geográfica de Sant Loreng d'Hortons.

Figure 1. Geographical situation of Sant Lloreng d"Hortons.

ZONA DE ESTUDIO

El yacimiento de La Pedrera está

situado al oriente de la localidad de Sant

Llorenc d“Hortons, provincia de Barce-

lona, junto al torrente de La Rierussa,

cerca de la masía de La Pedrera (Fig. 1).

Descripción de los niveles: MARTI-

NELL Y PORTA (1981) diferencian por

primera vez los niveles A y B de este yaci-

miento. El nivel A está formado por

margas amarillas fosilíferas, asentadas

sobre un nivel calizo arenoso. Sobre el

nivel A se sitúa el nivel B que está

formado por margas grisáceas, el cual se

presenta más compactado. En el nivel A

la fauna aparece concentrada levemente

en finos estratos, caracterizados por la

presencia de conchas de Pinnidae. Dentro

del nivel A, se ha muestreado especial-

mente un estrato de una potencia

máxima de 15 cm de morfología lenticu-

lar (Fig. 2). El material aparece en buen

estado, aunque ofrece evidencias de

haber sido levemente compactado, lo que

hace que la fauna se presente fisurada en

el interior del sedimento y aparezcan las

conchas fragmentadas una vez lavado.

En el nivel B el material se encuentra

mejor conservado, presentando alguna

vez trazas de la coloración original. Este

nivel se encuentra más cementado, pro-

bablemente debido a la precipitación de

carbonato procedente de la disolución de

las conchas que contenían los niveles are-

nosos superiores. En este nivel el material

aparece concentrado en lentejones.

MATERIAL Y MÉTODOS

La mayor parte del material estu-

diado procede del lavado de varias

muestras, obtenidas mediante muestreo

aleatorio sencillo, en un transecto de

unos dos metros cuadrados aproxima-

damente. En el curso de diversos mues-

treos se han lavado un total de 265 kilos.

Todas las muestras, excepto 5 kilos

recogidos en el nivel B, se tomaron en el

nivel A. El nivel B del yacimiento clásico

quedó enterrado, en Julio del 2000, tras

unas lluvias torrenciales.

Una vez lavado el material y pasado

por tres tamices, siendo el más fino de

300 micras de luz de malla, se recogieron

unos 6,5 litros de sedimento conchífero y

de otra fauna marina. La fracción fina,

inferior a 2 mm, alcanzaba unos 3,5 litros.

Predominaban en ese sedimento

conchas, valvas y restos de moluscos.

131

Iberus, 21 (1), 2003

arenas

] metros

oo margas

=jP98 margas

calcarenitas

Figura 2. Posición estratigráfica de las muestras estudiadas en la sección de La Pedrera.

Figure 2. Stratigraphical position of the studied samples in La Pedrera section.

También eran comunes restos de cirrípe-

dos (Balanus) y en menor medida restos

de otra fauna marina: otolitos, espinas,

vértebras y dientes de peces, pinzas de

cangrejos, radiolas y placas de erizos,

placas de asteroideos, el briozoo Schizopo-

rella unicornis (Johnston in Wood, 1844),

tubos de anélidos, ostrácodos, escasos

foraminíferos y algún fragmento de coral.

También se han estudiado, aunque

en escasa cantidad, ejemplares dejados

al descubierto por la lluvia, tanto en los

niveles A como B.

Se ha tratado de representar algunas

de las especies no citadas anteriormente

en este yacimiento y también aquellas

sobre las que se aporta nueva informa-

ción que consideramos de interés.Y se

ha procurado fotografiar al MEB la

mayoría de los micromoluscos. Las foto-

grafías fueron realizadas por Jesús

Méndez en el CACTI (Centro de Apoyo

Científico y Tecnológico a la Investiga-

ción) de la Universidad de Vigo.

Para la descripción de la protocon-

cha de las especies pertenecientes a la

familia Pyramidellidae se utilizará la

nomenclatura de AARTSEN (1987) con las

modificaciones adoptadas por LINDEN Y

EIKENBOOM (1992) y SCHANDER (1994), y

cuyas formas generales A-I, A-II, B y C,

pueden verse representadas en PEÑAS,

TEMPLADO Y MARTÍNEZ (1996) con pun-

tualizaciones en su medición en PEÑAS Y

ROLÁN (1999). De forma esquemática se

pueden comentar: protoconchas de tipo

A, con un ángulo alrededor de 90-120"

de desviación entre su eje y el de la

concha, siendo A-I cuando es planispiral

132

y A-II cuando es helicoidal; de tipo B,

cuando este ángulo tiene entre 130-160

y de tipo C cuando tiene aproximada-

mente 180".

El listado de especies, a nivel supra-

específico, ha sido confeccionado, bási-

camente, siguiendo el orden taxonómico

del CLEMAM (Check List of European

Marine Mollusca) http: / /www.mnhn.fr

/cgi-bin /mamlist.

La mayor parte del material aquí

tratado, así como el sedimento conchí-

fero y de otra fauna, ha sido depositado

en la Sección de Paleontología del

Museu del Vi, de Vilafranca del Penedes

(Barcelona). Todo él está a disposición

para ulteriores estudios.

En los pies de figura de las láminas,

las medidas que se mencionan para los

ejemplares allí representados se refieren

a la máxima dimensión de cada concha.

Abreviaturas utilizadas:

c concha

f fragmento

v valva aislada

h altura de la última vuelta

H altura total de la concha

D diámetro de la concha

= aproximadamente igual

MGSB Museo Geológico del Seminario

de Barcelona

MMP Museo Municipal Paleontológico,

Estepona, Málaga

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MVV Museu del Vi, Vilafranca del

Penedes, Barcelona

MORENO £7 42.: Fauna malacológica miocénica de La Pedrera

RESULTADOS

Se presentarán de la siguiente forma:

en primer lugar un listado general de

las especies aquí estudiadas. A continua-

ción, en una segunda parte, se hará

estudio más pormenorizado de algunas

de estas especies, las cuales serán

comentadas y muchas veces representa-

das. Finalmente, se hacen unos comen-

tarios finales sobre la información exis-

tente previamente y la aportada en este

trabajo y se termina con unas conclusio-

nes.

1. Listado de especies

Se presenta en la Tabla 1. En ella se

indican las nuevas especies y/o citas, las

figuras donde se representan en este

trabajo y su abundancia.

2. Descripción, comentarios y repre-

sentación de algunas especies nuevas o

sobre las que se aporta nueva informa-

ción.

De la mayoría de especies citadas

previamente para La Pedrera existen

fotografías y descripciones actualizadas

en la literatura. En este apartado nos

hemos limitado a referir solamente

algunos de los taxones de la lista ante-

rior que precisaban comentarios, ya

fuese por su rareza, la escasa informa-

ción existente sobre ellos, para aclarar

anteriores determinaciones discutibles,

O para describir aquellas nuevas espe-

cies para la ciencia. De algunas de estas

especies aportamos fotografías, la

mayoría de ellas al MEB.

Clase GASTROPODA

Teinostoma sp. (Figs. 14, 15)

Material examinado: 1 c.

Descripción: Concha diminuta, con

un diámetro de 1,0 mm, sólida, muy bri-

llante. Forma discoidal, muy aplanada,

con la espira que queda oculta bajo la

última vuelta. Escultura formada por

tenues estrías espirales concéntricas en

ambas caras. Abertura pequeña suboval.

Débil callosidad basal. No umbilicada.

Comentarios: El ejemplar fotografiado

procede del yacimiento miocénico

cercano de “Can Rosell”, Subirats, Bar-

celona, en el que la especie es común.

Bittium multiliratum Brusina, 1877

Bittium multiliratum Brusina, 1877. Journ. Conchi., 25: 380.

Cerithidium sculptum Seguenza, 1880. Mem. Cl. Sc. Fis., Mat. e Nat. R. Acc. Lincei, 53, 6 (6): 109,

lám. 11, fig. 16.

Material examinado: 3 c.

Descripción: En BALUK (1975) y en

MORONI Y RUGGIERI (1988).

Dimensiones: 4,5 mm en 7 vueltas de

la teloconcha.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Korytnika, Polo-

nia (BALUK, 1975). Mioceno medio:

Cuenca de Viena, Moravia y Hungría

(BRUSINA, 1877); Mioceno medio (Lang-

hiense): La Pedrera, Barcelona. Mioceno

superior: Sicilia, Italia (MORONI Y RUG-

GIERI, 1988).

Comentarios: Nuestros ejemplares se

corresponderían a la variedad B. multili-

ratum saheloparvulum (Moroni, 1955),

caracterizada por tener 6-7 cordones

espirales en la penúltima vuelta y la

ausencia de escultura axial.

133

Iberus, 21 (1), 2003

Tabla I. Listado general de las especies estudiadas en el presente trabajo. Especies en negrita son

objeto de comentarios en el texto; (*): primera cita para el área de estudio; (**): primera cita para la

Península Ibérica; (***): nueva especie para la ciencia o nombre específico nuevo (no se consideran

citas ni especies nuevas para la zona o la Península aquellas que sólo se mencionan con el nombre

del género sin denominación especifica); Fig: figura de este trabajo donde se representa esa especie

(casi todo el material fotografiado procede de La Pedrera, en el texto se indican sólo las conchas

fotografiadas que procedan del yacimiento cercano de Can Rosell, Subirats); f: sólo fragmentos; +:

1-2 ejemplares, ++: 3-10 ejemplares, +++: 11-100 ejemplares, ++++: más de 100 ejemplares; B:

especie encontrada solamente en el nivel B.

Table I. General list of the species studied in this paper. Species in bold are commented in the text; (*):

first record in the studied area; (**): first record in the Iberian Peninsula; (*%): new species or new spe-

cific name (those species only mentioned by its genre are not cosidered either new species in the area or

the Iberian Peninsula); Fig: figure in this paper where the species is shown (almost all the photographed

material comes from La Pedrera, in the text only the shells from the nearby Can Rosell deposit are indi-

cated); f: only fragments; +: 1-2 specimens, ++: 3-10 specimens, +++: 11-100 specimens, ++++: more

than 100 specimens, B: species found only in level B.

Species Fig Abundancia

Clase GASTROPODA

Familia NERITIDAE

* Smaragdia picta (Férussac, 1825) +

Familia SKENEIDAE

** Teinostoma defrancei (Basterot, 1825) Figs. 3-6 +++

leinostoma minimum Boettger, 1907 Figs. 7-10 ++

Teinostoma woodi (Hórmes, 1856) ) Figs. 11-13 ++

Teinostoma sp. Figs. 14-15 + B

Familia CERITHIIDAE

Cerithium bronni Dollfus y Dautzenberg, 1886 Fig. 16 +

Cerithium vulgatum Bruguiére, 1792 f +

** Bittium multiliratum Brusina, 1877 +

** Bittium spina (Hórnes, 1855) ++

Familia SCALIOLIDAE

Sandbergeria perpusilla (Grateloup, 1827) HH

Familia POTAMIDIDAE

Potamides theodiscus (Rolle in Hilbert, 1879) +HH++

Familia BATILLARIDIDAE

Granulolabium pictum (Basterot, 1825) +++

Familia TURRITELLIDAE

Turritella bicarinata Eichwald, 1830 ++

lurritella cf. terebralis Lamarck, 1822) +.

*Turritella turris Basterot, 1825 f + B

* Turritella subangulata (Brocchi, 1814) f +

Protoma proto (Basterot, 1825) ne

Protoma rotifera (Lamarck, 1804) +

** Protoma vasconiensis Cossmann y Peyrot, 1917 +

Familia TRIPHORIDAE

Monophorus perversus (Linné, 1758) H

Familia CERITHIOPSIDAE

** Cerithiopsis minima (Brusina, 1865) Figs. 17-19 +++

** Cerithiopsis lottae Wienrich, 2001 Fig. 20 +

Seila sp. +

134

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

Tabla I. Continuación.

Table [. Continuation.

Species Fig Abundancia

Familia ACLIDIDAE

** Graphis albida (Kanmacher, 1798) f +

** Alis cf. ascaris (Turton, 1819) se

Familia EPITONIIDAE

Epitonium Sp. ++

Acrilloscala sp. +

Familia EULIMIDAE

* Eulima glabra (Da Costa, 1778) +

* Balcis alba (Da Costa, 1778) ++

* Niso terebellum (Chemnitz, 1788) +

Eulima sp. +

Familia RISSOIDAE

Rissoa Sp. ++

** Obtusella orisparvi spec. nov. Figs. 21-24 +++

Obtusella sp. +

Familia ADEORBIDAE

* Circulus striatus (Philippi, 1836) ++

Familia ELACHISINIDAE

** Elachisina moravica (Rzehok, 1893) Figs. 25-26 ++

Familia CAECIDAE

*** Caecum veraí spec. nov. Figs. 27-30 +++

Familia IRAVADIIDAE

** Chevallieria ct. peculiaris (Boettger, 1901) +

Familia HYDROBIIDAE

** Hydrobia peregrina Boettger, 1901 Figs. 31-33 +++

Familia TORNIDAE

* Tornus belgicus belgicus Baluk, 1975 +

* Tornus planorbillus (Dujardin, 1837) +

** Discopsis belgicus Glibert, 1949 Figs. 34-36 +

** Discopsis canvi (Morgan, 1920) Figs. 37-39 +++

* Discopsis trigonostoma (Bosterot, 1825) Figs. 40-42 ht

Familia VERMETIDAE

* Vermetus granulatus (Gravenhorst, 1831) dE

Serpulorbis arenarius (Linné, 1767) H

Familia STROMBIDAE

Tibia dentata ordalensis Almera y Bofill, 1886 Fig. 43 +

Dientomochilus decussatus (Defrance in Basterot, 1825) ze

Familia CALYPTRAEIDAE

Colyptraea chinensis (Linné, 1758) US

Colyptraea ornata Bosterot, 1825 H+

Crepidula unguiformis Lamarck, 1822 --

Familia CYPRAEIDAE

Cypraea sp. f +

Fomnilia NATICIDAE

Criptonotica sp. +H

** Eunaticina striatella (Groteloup, 1828) Figs. 44-45 ++

Lunotía helicina (Brocchi, 1814) +H

135

Iberus, 21 (1), 2003

Tabla 1. Continuación.

Table [. Continuation.

Species

Notica millepunctata Lamarck, 1822

Neverita josephina Risso, 1826

** Tectonatica miopusilla (Kautsky, 1925)

Familia CASSIDAE

Cassidaria Sp.

Familia BURSIDAE

* Bursa cancellata (Gratelup, 1840)

Familia MURICIDAE

Murex aquitanicus (Grateloup, 1833)

** Murex aturensis (Cossmann y Peyrot, 1923)

Murex sp.

* Ocinebrina avitensis (Cossmann y Peyrot, 1923)

Ocinebrina sp.

Hadriania craticulata boecki (Hómes y Auinger, 1885)

Purpura duvergieri (Cossmann y Peyrot, 1924)

** Purpura vindobonensis altispirata Venzo y Pelosio, 1963

* Tritonalia polimorpha (Brocchi, 1814)

Familia BUCCINIDAE

Melongena comuta (Agassiz, 1843)

Phos poligonus (Brocchi, 1814)

Familia FASCIOLARIIDAE

Euthriofusus burdigaliensis (Defrance, 1820)

Familia NASSARIIDAE

Amyclina aff. Karreri (Hórnes y Avinger, 1822)

Nassarivs cf. aturensis (Peyrot, 1928)

Nassarivs girondicus (Peyrot, 1927)

** Nassarius impar (Bellardi, 1878)

Nassarius schoenni (Hómes y Avuinger, 1882)

Cyllene vulgatissima Peyrot, 1927

Familia COLUMBELLIDAE

Pyrene fallax (Hóres y Avinger, 1880)

Familia OLIVIDAE

Oliva sp.

Familia VASIDAE

* Tudicla rusticula (Basterot, 1825)

Familia CANCELLARIIDAE

Cancellaria sp.

Narona varicosa (Brocchi, 1814)

Narona dufoiri (Grateloup, 1832)

Trigonostoma scrobiculatum (Hómes, 1856)

Familia CONIDAE

* Conus dujardini Deshayes, 1845

Familia TEREBRIDAE

Strioterebrum basteroti (Nyst, 1845)

Subula fuscata (Brocchi, 1814)

lerebra acuminata Borson, 1820

Familia TURRIDAE

136

** Agathotoma hautensis (Csepreghy-Meznerics, 1953)

Bela sp. |

Fig

Fig. 46

Fig. 47

Figs. 48-49

Fig. 50

Figs. 51-52

Figs. 53-55

Fig. 56

Abundancia

+++

MORENO £7 ALz.: Fauna malacológica miocénica de La Pedrera

Tabla I. Continuación.

Table I. Continuation.

Species

Bela sp. 2

Clavatula spinosa (Grateloup, 1832)

Clavatula jovanneti (Desmoulins, 1872)

Gemmula denticula (Basterot, 1825)

Gemmulo sp.

Raphitoma ct. hispidula (Jan in Bellardi, 1847)

Familia PYRAMIDELLIDAE

Pyramidella unisulcata Dujardin, 1837

** Chrysallida cimbrica (Koutsky, 1925)

Chrysallida sp.

* Eulimella acicula (Philippi, 1836)

** Eulimella iusta nom. nov.

*** Eulimella penedesensis spec. nov.

* Eulimella scillae (Scacchi, 1835)

Eulimella sp. |

Eulimella sp. 2

** Syrnola hoernesi (Koenen, 1882)

** Syrnola laevissima (Basquet, 1859)

Syrnola sp.

* Megastomia pallidaeformis (Sacco, 1892)

** Odostomia degrangei (Cossmann y Peyrot, 1917)

** Odostomia aff. desuefacta Peñas y Rolún, 1999

** Odostomia vera nom. nov.

** Odostomia perstricta (Sacco, 1892)

Odostomia polysarcula (Cossmann y Peyrot, 1917)

* Odostomia turritella (Groteloup, 1832)

Odostomia sp. |

Odostomia sp. 2

Ondina sp.

*** Noemiamea batllorii spec. nov.

** Turbonilla pliomagna Sacco, 1892

** Turbonilla sallomacensis Cossmann y Peyrot, 1917

** Turbonilla olivellai spec. nov.

Turbonilla sp. |

Turbonilla sp. 2

Familia EBALIDAE

** Anisocycla cf. bezanconi Cossmonn y Lambert, 1884

** Anisocycla eulimoides Fekih, 1969

Familia AMATHINIDAE

* Clathrella sulcosa (Brocchi, 1814)

Familia ACTEONIDAE

* Acteon semistriatus (Basterot, 1825)

Ácteon sp.

Familia RETUSIDAE

** Retusa obtusa (Montagu, 1803)

** Retuso semisulcato (Philippi, 1836)

Retuso truncatula (Bruguiére, 1792)

Fig

Figs. 57-60

Fig. 61

Figs.62-66

Figs. 67-68

Figs. 69-73

Fig. 74

Figs. 75-77

Figs. 78-79

Figs. 80

Figs. 81-82

Fig. 83

Figs. 84-87

Figs. 88-92

Fig. 93

Figs. 94-96

Fig. 97-98

Figs. 99-101

Fig. 102-103

Figs. 104-107

Figs. 108-109

Abundancia

23%

137

Iberus, 21 (1), 2003

Tabla I. Continuación.

Table I. Continuation.

Species Fig

Familia RINGICULIDAE

* Ringicula ventricosa (Sowerby, 1824)

Ringicula sp.

Familia CYLICHNIDAE

Acteocina lajonkaireana (Basterot, 1825)

Familia CAVOLINIDAE

Vaginella austriaca Kittl, 1886

Familia ELLOBIIDAE

Ovatella sp.

Clase BIVALVIA

Familia ARCIDAE

(Lamarck, 1805)

Familia NOETIDAE

Noetiella rollei (Húmes, 1870)

Familia MYTILIDAE

Myrilus cf. michelin; Matheron, 1843

Familia PINNIDAE

Átrina pectinata (Linné, 1767)

Familia ANOMIIDAE

Anomia ephippivm Linné, 1758

Familia OSTREIDAE

Ostrea edulis Linné, 1758 Figs. 110-113

Familia CHAMIDAE

* Chama gryphoides linné, 1758

Familia CARDIIDAE

Cardium cf. hians Brocchi, 1814

* Aconthocardia paucistriata (G.B. Sowerby, 1841)

Fomilia TELLINIDAE

* Gastrana lacunosa (Chemnitz, 1782)

Familia DONACIDAE

Donax sp.

Familia SEMELIDAE

Ervilia . pusilla (Philippi, 1836)

Familia KELLIELLIDAE

** Alveidus nitidus (Reuss, 1867) Figs. 114-120

Familia VENERIDAE

Callista sp.

Venus sp.

Pitar rudis (Poli, 1795)

lapes sp.

Familia CORBULIDAE

* Corbula revoluta (Brocchi, 1814)

* Corbula gibba (Olivi, 1792)

Close SCAPHOPODA

Familia DENTALIIDAE

* Antolis mutabilis (Hómes, 1856)

* Fustiaria jani (Hórmes, 1856)

Familia GADILINIDAE

* Gadilina triquetra (Brocchi, 1814)

138

Abundancia

MORENO £7 4L.: Fauna malacológica miocénica de La Pedrera

200 um

Figuras 3-6. Teinostoma defrancel. 3-5: conchas, 2,5, 2,0 y 1,7 mm; 6: protoconcha. Figuras 7-10.

Leimostoma minimum. 7-9: conchas, 1,5,1,5 y 1,9 mm; 10: protoconcha.

Figures 3-6. Teinostoma defrancei. 3-5: shells, 2.5, 2.0 and 1.7 mm; 6: protoconch. Figures 7-10. Tei-

nostoma minimum. 7-9: shells, 1.5, 1.5 and 1.9 mm, 10: protoconch.

189

Iberus, 21 (1), 2003

Cerithiopsis minima (Brusina, 1865) (Figs. 17-19)

Cerithium minimum Brusina, 1865. Verhandl. Zool. Bot. Gessell., Wien, 15: 17.

Bittium reticulatum (Da Costa, 1778) in BATLLORI (1995). Tesis Doct. Univ. Barcelona, lám. 7, figs.

25.1-25.3.

Material examinado: 70 c.

Descripción: Concha (Figs. 17, 18) di-

minuta, pupoide; protoconcha (Fig. 19)

lisa, subcilíndrica, de 4 vueltas; Escultu-

ra de la teloconcha formada por tres cor-

dones espirales desde la primera vuelta,

granulosos y un cordón basal espiral en

ejemplares adultos. No hemos apreciado

diferencias en las dimensiones, tanto de

la concha como de la protoconcha, entre

la especie fósil y la actualmente vivien-

te, ni tampoco hemos observado dife-

rencias en la relación H/D.

Dimensiones medias: 2,5 x 0,6 mm en 6

vueltas de la teloconcha.

Comentarios: No hemos encontrado

cita de esta especie ni en el Mioceno ni

en el Plioceno, y sí de C. tubercularis

(Montagu, 1803) y de C. nana (Wood,

1848), especies parecidas. HARMER

(1915) cita C. minima para el Crag britá-

nico, pero según MARQUET (1997), con el

que estamos de acuerdo, la determina-

ción es incorrecta ya que se trata de C.

nana.

Obtusella orisparvi spec. nov. (Figs. 21-24)

Material tipo: Holotipo (Fig. 21) y un paratipo depositados en el MNCN (n*% MNCNI-37741 y

37742). Otros paratipos: MVV (3), MGSB (3) y MMP (3)

Otro material examinado: Más de 200 c.

Localidad tipo: Mioceno medio (Langhiense), La Pedrera, Sant Llorenc d'Hortons (Barcelona).

Etimología: El nombre específico deriva de las palabras latinas os-oris, boca, y parvus-a-um, pequeño,

que hacen referencia a la pequeñez de la abertura de esta concha.

Descripción: Concha (Figs. 21, 22) dimi-

nuta, pero sólida, conoidea, algo pupoide.

Apice obtuso, con la protoconcha lisa (Fig.

24), dificil de definir su longitud por no

presentar un claro límite con la telocon-

cha, pero presentando un cordón spiral

próximo a la sutura. Espira total de la '

concha con unas 3,5-4 vueltas convexas.

Sutura no profunda, impresa (Fig. 23). Hay

indicios de algunas estrías espirales sub-

suturales y las líneas de crecimiento son

ligeramente prosoclinas. Abertura circu-

lar, pequeña. Sin claro ombligo, pero en

algunas conchas con una estrecha fisura

umbilical. Labro externo delgado.

Dimensiones del holotipo: 0,96 x 0,56

mm en 4 vueltas.

Comentarios: Esta diminuta especie

presenta claras diferencias con las dos

especies de este género actualmente

vivientes en el Mediterráneo y también

con O. sabelliana Tabanelli, 1991 de la

que hemos observado varios topotipos

140

(col. Tabanelli). Esta especie vivió en

aguas profundas del Plioceno inferior,

asociada a Korobkovia oblonga y a Limea

strigillata (Tabanelli com. pers.). Nuestra

especie, que vivió en aguas someras

tiene un perfil menos cónico, con la

espira más elevada, las vueltas más con-

vexas y la protoconcha es lisa, mientras

que en O. sabelliana es esculturada.

O. macilenta (Monterosato, 1880),

actual, que vive en aguas profundas del

Mediterráneo, también tiene la proto-

concha lisa, pero la concha tiene un

perfil más cónico, con la última vuelta

mucho más ancha, tiene la sutura pro-

funda, la abertura mucho mayor y tiene

un claro ombligo.

O. intersecta (S. W. Wood, 1857), del

Plioceno y actual, tiene una concha

mucho mayor, globosa, con clara escul-

tura espiral y umbilicada.

O. lata Rolán y Rubio, 1999, actual, y

que vive en el archipiélago de Cabo

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

Figuras 11-13. Teímostoma wood. 11, 12: conchas, 2,1 y 1,5 mm; 13: protoconcha. Figuras 14, 15.

Teinostoma sp. Concha, 1,0 mm, Can Rosell.

Figures 11-13. Teinostoma woodi. 11, 12: shells, 2.1 and 1.5 mm; 13: protoconch. Figures 14, 15.

Teínostoma sp. Shell, 1.0 mm, Can Rosell.

Iberus, 21 (1), 2003

Verde, es más globosa y más cónica,

mientras que O. orisparvi es mas alargada

y con tendencia a la forma cilíndrica.

O. communis Boettger, 1906, del Tor-

toniense de Kostej, Hungría, también

carece de escultura espiral, pero tiene

una forma más cónica, casi piramidal,

con la última vuelta más ancha; H/D =

1,5 frente a 1,7 en O. orisparvi; y la aber-

tura es mayor: 45% H, frente al 35%. Ver

en ZICH (1934: lám. 5, fig, 78) ilustración

del holotipo.

Circulus striatus (Philippi, 1836)

Valvata striata Philippi, 1836. En. Moll. Soc., 1: 147, lám. 9, figs. 3a-c.

Tornus planorbillus (Dujardin, 1837) in BATLLORI, 1995. Tesis Doct. Univ. Barcelona, lám. 3, fig. 7.

Material examinado: 3 c.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Cuenca del

Loira, Francia (GLIBERT, 1952); Mioceno

medio (Langhiense): La Pedrera, Barce-

lona. Mioceno superior: Tortoniense de

Calabria, Italia (SEGUENZA, 1880). Plio-

ceno: Kallo, Cuenca del Mar del Norte,

Bélgica (MARQUET, 1997), Crag británico

(WooD, 1848); Plioceno piamontés, Italia

(PAvIA, 1975); Tabiano modenense, Italia

(Coprr, 1880); Estepona, Málaga (VERA-

PELÁEZ ET. AL., 1996). Actual: Mediterrá-

neo y costa occidental de África (ADAM

Y KNUDSEN, 1969).

Dimensiones: de 2,3 a 3,5 mm.

Comentarios: Las conchas encontra-

das en el yacimiento de La Pedrera no

presentan diferencias apreciables con las

actuales. Se trata de la primera cita para

el yacimiento.

Elachisina moravica (Rzehak, 1893) (Figs. 25, 26)

Littorina moravica Rzehak, 1893. Fauna der Oncophora Schichten Maehrens: 30, lám. 2, figs. 2, 3.

Material examinado: 80 c.

Descripción: Concha (Fig. 25) muy

pequeña naticiforme, con las vueltas

convexas, de crecimiento muy rápido,

la última vuelta globosa. Protoconcha

(Fig. 26) obtusa, lisa. Escultura de la

teloconcha formada por finas estrías

espirales en toda la superficie, distri-

buidas de forma regular. Abertura

grande, subcircular, con el peristoma

continuo. Columela delgada, a veces

replegada hacia la zona umbilical.

Fisura umbilical estrecha pero profun-

da.

Dimensiones medias: 1,8 x 1,4 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio de Moravia, Rep.

Checa (COSSMANN, 1921). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Esta es la primera cita

para el Mioceno de la Península Ibérica.

Ver PONDER (1985), en el que des-

cribe la nueva familia Elachisinidae

(Rissoacea).

Actualmente está en estudio la revi-

sión de las Elachisina del África Occiden-

tal, las cuales podrían ser descendientes

de esta especie miócenica, la única que

conocemos.

Caecum verai spec. nov. (Figs. 27-30)

Material tipo: Holotipo (Fig. 27) y un paratipo, depositados en el MNCN (n*% MNCNLI- 37743 y

37744). Otros paratipos: MVV (1), MGSB (2) y MMP (1).

Otro material examinado: 3 c.

142

MORENO ET 4L.: Fauna malacológica miocénica de La Pedrera

100 um

Figura 16. Cerithium bronni. Concha, 6,1 mm. Figuras 17-19. Cerithiopsis minimum. 17, 18:

conchas, 2,5 y 2,2 mm; 19: protoconcha. Figura 20. Cerithiopsis lottae. Concha, 4,1 mm. Figuras

21-24. Obtusella orisparvi. 21: holotipo, 0,96 mm; 22: concha, 0,8 mm; 23: detalle de la sutura;

24: protoconcha.

Figure 16. Cerithium bronni. Shell, 6.1 mm. Figures 17-19. Cerithiopsis minimum. 17, 18: shells,

2.5 and 2.2 mm; 19: protoconch. Figure 20. Cerithiopsis lottae. Shell, 4.1 mm. Figures 21-24. Obtu-

sella orisparvi. 21: holotype, 0.96 mm; 22: shell, 0.8 mm; 23: detail of the suture; 24: protoconch.

143

Iberus, 21 (1), 2003

Localidad tipo: Mioceno medio (Langhiense), La Pedrera, Sant Llorenc d'Hortons, (Barcelona).

Etimología: El nombre específico se dedica al Dr. José Luis Vera, del MMP, de Estepona, por su

ayuda en la determinación de varias especies.

Descripción: Concha (Figs. 27, 28)

diminuta, cilíndrica. Escultura anular

(Fig. 29) conspicua formada por estrías y

cordoncillos irregularmente distribui-

dos. Zona apical (Fig. 30) más atenuada,

con el septum muy obtuso, suavemente

arqueado en su parte anterior y ligera-

mente anguloso en la posterior. El perfil

de la concha se va haciendo más estre-

cho en la zona de la apertura, la cual es

circular, con el borde afilado.

Dimensiones del holotipo: 1,7 x 0,45 mm.

Discusión: Difiere de C. banoni

Benoist, 1874, del Mioceno inferior

(Aquitaniense) francés, que tiene unas

dimensiones similares, pero con un

perfil más anguloso; carece de escultura

anular; la abertura sigue el perfil de la

concha y en su borde interior tiene un

delgado anillo; finalmente, tiene el

septum prominente. Las ilustraciones de

esta especie en LOZOUET, LESPEROT Y

RENARD (2001) presentan un septum

redondeado, mientras que en GLIBERT

(1952a) es triangular.

C. subanulatum De Folin, 1870,

viviente en el Mediterráneo y citada en

el Plioceno, tiene unas dimensiones y un

septum parecidos, pero carece de escul-

tura anular y en la abertura tiene un

anillo bien desarrollado. PORTA, MARTI-

NELL Y GONZÁLEZ DELGADO (1993, lám.

3, fig. 4) muestran una concha proce-

dente del Mioceno Langhiense de Sant

Sebastiá dels Gorchs, Barcelona, que

determinan como GC. subanulatum,

aunque nos parece más bien la especie

que aquí describimos como C-. veral.

C. glabrum (Montagu, 1803), viviente

en el Atlántico europeo, citada por HAR-

MER (1921) para el Crag británico y por

GLIBERT (1952a) para el Mioceno medio

de la Cuenca del Loira, tiene unas di-

mensiones mayores, carece de escultura

anular, tiene la abertura menos ate-

nuada y su septum es muy prominente.

CHIRLI (1995) en su revisión del

género Caecum en el Plioceno Toscano

describe e ilustra C. crispum Cerulli-

Irelli, 1912 y C. nysti De Stefani y Panta-

nelli, 1878. Ambas especies tienen una

concha mayor; además, la primera tiene

un septum globoso, muy prominente y

la segunda lo tiene triangular.

Discopsis belgicus (Glibert, 1949 ) (Figs. 34-36)

Adeorbis belgicus Glibert, 1949. Mem. Inst. Roy. Sci. Nat. Belgique, 121: 113, lám. 6, fig. 14.

Material examinado: 1 c.

Descripción: En GLIBERT (1949, 1952a)

y en MARQUET (1997). En el presente tra-

bajo se representa un ejemplar no

adulto.

Dimensiones: 1,2 mm de diámetro.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Cuenca del

Loira, Francia (GLIBERT, 1949); Mioceno

medio (Langhiense): La Pedrera, Barce-

lona. Plioceno: Kallo, Cuenca del Mar

del Norte, Bélgica (MARQUET, 1997).

Comentarios: Esta es la primera cita

para el Mioceno de la Península Ibérica.

De las tres especies encontradas en La

Pedrera pertenecientes a este género, ésta

144

se diferencia a primera vista por su espira

saliente y sus dos carenas basales.

Las especies actuales de África Occi-

dental que se habían incluido por ADAM

Y KNUDSEN (1969) en el género Cochliole-

pis no parecen tener una real similitud en

sus partes blandas con la especie tipo de

este género del Caribe Cochliolepis parasi-

tica Stimpson, 1859. En un reciente trabajo

(ROLÁN Y RUBIO, 2002) este grupo de espe-

cies actuales, que morfológicamente se

parecen a las fosiles aquí estudiadas, se

han considerado en el género Discopsis De

Folin, 1869 y por ese motivo utilizamos

este género para la ubicación de las mismas

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

100 um

Figuras 25, 26. Elachisina moravica. 25: concha, 1,5 mm; 26: protoconcha. Figuras 27-30. Caecum

verai. 27: holotipo, 1,7 mm; 28: concha; 29: detalle de la escultura anular; 30: septum. Figuras 31-

33. Hydrobia peregrina. 31, 32: conchas, 1,8 y 1,9 mm; 33: protoconcha.

Figures 25, 26. Elachisina moravica. 25: shell, 1.5 mm; 26: protoconch. Figures 27-30. Caecum

verai. 27: holotype, 1.7 mm; 28: shell, 1.8 mm; 29: detail of the annular sculpture; 30: septum. Figures

31-33. Hydrobía peregrina. 31, 32: shells, 1.8 and 1. 9 mm; 33: protoconch.

145

Iberus, 21 (1), 2003

Discopsis canui (Morgan, 1920) (Figs. 37-39)

Adeorbis woodi Ivolas y Peyrot, 1900: Contr. Paléont. Tour. (1900): 154 (non Hórnes).

Tornus canui Morgan, 1920. Bull. Soc. Géol. France, 19: 343, fig. 40.

Material examinado: 14 c.

Descripción: En MORGAN (1920) y en

GLIBERT (1952a). Se ilustra la concha y la

microescultura de la base.

Dimensiones medias: 1,1 mm de diá-

metro.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Faluns de la Tou-

raine (MORGAN, 1920) y Cuenca del Loira,

Francia (GLIBERT, 1952a). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Esta es la primera cita

para el Mioceno de la Península Ibérica.

Esta especie se caracteriza por tener

la espira más deprimida, ausencia de

carena basal y tener la superficie muy

tenuemente estriada.

Discopsis trigonostoma (Basterot, 1825) (Figs. 40-42)

Delphinula trigonostoma Basterot, 1825. Mem. Soc. Hist. Nat. París, 2: 28, lám. 4, fig. 10.

Material examinado: 36 c.

Descripción: En COSSMANN Y PEYROT

(1918), GLIBERT (1952a) y JANSSEN (1984).

En el presente trabajo se representa un

ejemplar no adulto y la escultura de la

base.

Dimensiones medias: 1,3 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Aquitaniense

de Saucats, Francia (LOZOUET ET AL.

2001); Burdigaliense de Léognan,

Francia (BASTEROT, 1825 y COSSMANN Y

PEYROT, 1918). Mioceno medio: Cuenca

del Loira (GLIBERT, 1952); Winterswijk-

Miste, Holanda (JANSSEN, 1984); Cuenca

del Rhing, Kevelaer, Alemania (WIEN-

RICH, 2001). Mioceno medio (Lang-

hiense): La Pedrera, Barcelona. Mioceno

superior: Tortoniense de Kostej,

Hungría (BOETTGER, 1906).

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibé-

rica.

Eunaticina striatella (Grateloup, 1828) (Figs. 44, 45)

Natica striatella Grateloup, 1828. Bull. Hist. Nat. Soc. Lin. Bordeaux, 2: 158 n* 146.

Sinum sp. in BATLLORI (1995). Tesis Doct. Univ. Barcelona.

Material examinado: 3 c.

Descripción: En COSSMANN Y PEYROT

(1919) y en JANSSEN (1984). En este

trabajo se muestran concha y protocon-

cha.

Dimensiones medias: 9,8 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Aquitaniense

de Saucats y de las Landas y Burdiga-

liense de St-Jean-de-Marsacq (LOZOUET

ET AL., 2001). Mioceno medio: Winters-

wijk-Miste, Holanda (JANSSEN, 1984).

146

Mioceno medio

Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Creemos que es la única especie fósil

de este género encontrada en Europa, la

cual es muy cercana a E. africana Burnay

y Fernandes, 1984, especie actual que

vive en Angola y Gabón. Varias especies

de este género, típico de aguas tropica-

les, viven también en el Indo-Pacífico.

(Langhiense): La

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

100 um

Figuras 34-36. Discopsis belgicus. 34, 35: concha, 1,2 mm; 36: detalle de la escultura. Figuras 37-

39. Discopsis canui. 37, 38: conchas, 1,3 y 1,0 mm; 39: detalle de la escultura basal. Figuras 40-42.

Discopsis trigonostoma. 40, 41: conchas, 1,1 y 1,0 mm; 42: detalle de la escultura basal.

Figures 34-36. Discopsis belgicus. 34, 35: shell, 1.2 mm; 36: detail of the sculpture. Figures 37-39.

Discopsis canui. 37, 38: shells, 1.3 and 1.0 mm; 39: detail of the basal sculpture. Figures 40-42. Dis-

copsis trigonostoma. 40, 41: shells, 1.1 and 1.0 mm; 42: detail of the basal sculpture.

Iberus, 21 (1), 2003

Nassarius cf. aturensis (Peyrot, 1928) (Figs. 48, 49)

Nassa (Telasco?) aturensis Peyrot, 1928. Act. Soc. Lin. Bordeaux, 79 suppl.: 132, lám. 3, figs. 100-105.

Material examinado: más de 300 c.

Descripción: En PEYROT (1928) y en

BATLLORI (1995). Concha (Fig. 48)

pequeña, conoidea, robusta. Concha lar-

varia (Fig. 49) de 3 vueltas, planctotró-

fica, con un diámetro de unas 750 qm.

Teloconcha de espira poco elevada, con

unas 6 vueltas ligeramente convexas.

Unas 16 costillas axiales robustas, redon-

deadas, ligeramente escalonadas, mucho

más anchas que sus interespacios. Escul-

tura espiral débil formada por estrechos

surcos en los interespacios, más aprecia-

ble en la base de la última vuelta. Aber-

tura subcircular, con el canal sifonal

corto. Peristoma continuo. Labro externo

varicoso en su exterior y en ejemplares

adultos con seis dientes en su interior.

Dimensiones: se han medido 100

conchas presentando unas dimensiones

medias de 3145 x 1,75 mm. y unas

dimensiones máximas de 5,75 x 2,25

mm, el ejemplar ilustrado mide 4,5 x 2,2

mm. La relación H/D media es de 1,97.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Aquitaniense,

Francia (PEYROT, 1928). Mioceno medio

(Langhiense): Alt Penedes, Barcelona

(BATLLORI, 1995).

Comentarios: BATLLORI (1995) consi-

dera que no existen diferencias aprecia-

bles con la forma típica descrita por

PEYROT (1928), salvo su menor tamaño,

que atribuye a las condiciones paleoe-

cológicas especiales de La Pedrera. En

el material recolectado por nosotros, no

sólo las diferencias de tamaño son muy

considerables (85 x 3,55 mm en la

especie aquitánica frente a 5,75 x 2,25

mm como máximo en la especie de La

Pedrera), sino que las relaciones H/D

son muy diferentes: > 2,4 en la especie

aquitánica, frente a < 2 en la de La

Pedrera. Por otra parte los ejemplares

de La Pedrera tienen un perfil casi esca-

lonado, y la escultura axial es siempre

conspicua.

Nassarius schoenni (Hoernes y Auinger, 1882) (Figs. 51, 52)

Buccinum schónni Hoernes y Auinger, 1882. Die Gasteropoden.. 125, lám. 15, figs. 18-20.

Nassarius saccoi (Trentanove, 1911) in BATLLORI (1995). Tesis Doct. Univ. Barcelona: 347-350, lám.

18, figs. 59.1-59.3.

Material examinado: 40 c.

Descripción: En BALUK (1970 y 1997).

Se ilustra aquí un ejemplar adulto (Figs.

o):

Dimensiones: 10 x 6,7 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Cuenca de

Viena, Austria (HÓRNES, 1856); Koryt-

nika, Polonia (BALUK, 1997); Cuenca

mediterránea austro-húngara (STRAUSZ,

1966); Mioceno medio (Langhiense):

Depresión del Valles ( (VILLALTA Y

ROSELL, 1966 y CALZADA ET AL., 1972);

La Pedrera, Barcelona; Colli turinés,

148

Italia (BELLARDI, 1882). Mioceno supe-

rior: Tortoniense polaco (BALUK, 1970).

Comentarios: Los ejemplares de La

Pedrera presentan un tamaño menor

que los de la forma típica; también

tienen la última vuelta más cuadrangu-

lar y la callosidad labial muy desarro-

llada en la zona adapical. Esta forma

coincide con las ilustradas por BALUK

(1970: lám. 12, fig. 13 y 1997: lám. 1, fig.

1) y también con la descrita como N.

dujardini (Deshayes, 1844) por PEYROT

(1928: 141, lám. 3, figs. 124-126).

MORENO E£7 4L.: Fauna malacológica miocénica de La Pedrera

200 um

Figura 43. Tibia dentata ordalensís. Protoconcha. Figuras 44, 45. Eunaticina striatella. 44: concha,

9,8 mm; 45: protoconcha. Figura 46. Hadriania craticulata boecki. Concha, 15,2 mm. Figura 47.

Phos poligonus. Concha, 11,1 mm. Figuras 48, 49: Nassarius cf. aturensis. 48: concha, 4,5 mm; 49:

protoconcha. Figura 50. Nassarius impar. Concha, 6,0 mm.

Figure 43. Tibia dentata ordalensis. Protoconch. Figures 44, 45. Eunaticina striatella. 44: shell, 9.8

mm; 45: protoconch. Figure 46. Hadriania craticulata boecki. Shell, 15.2 mm. Figure 47. Phos poli-

gonus. Shell, 11.1 mm. Figures 48, 49. Nassarius cf. aturensis. 48: shell, 4.5 mm; 49: protoconch.

Figure 50. Nassarius impar. Shell, 6.0 mm.

Iberus, 21 (1), 2003

Clavatula spinosa (Grateloup, 1832) (Figs. 57-60)

Pleurotoma spinosa Grateloup, 1832. Actes Soc. Linn. Bordeaux, 6e article: 5, 30: 324, lám. 19, fig. 24.

Material examinado: Varios fragmentos.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno de Dax y Burdeos,

Francia (GRATELOUFP, 1832). Mioceno me-

dio (Langhiense): La Pedrera, Barcelona.

Comentarios: BATLLORI (1995) des-

cribe C. asperulata (Lamarck, 1822) para

el Mioceno del Penedes, especie muy

citada durante el Mioceno en diferentes

cuencas europeas, la cual se caracteriza

por el desarrollo de espinas desde las

vueltas juveniles. Por el contrario, en los

ejemplares de La Pedrera y también en

los del yacimiento cercano de Can

Rosell no ocurre así. Según VERA (com.

per.) nuestros ejemplares pertenecen a la

especie C. spinosa, la cual se caracteriza

por presentar una doble carena muy

pronunciada.

Las conchas fotografiadas (Figs. 57-

59) pertenecen al yacimiento cercano de .

Can Rosell, Subirats, Barcelona, donde

esta especie es abundante. La protocon-

cha (Fig. 60) es lisa con 2,5 vueltas de

espira.

Chrysallida cimbrica (Kautsky, 1925) (Figs. 62-66)

Pyrgulina cimbrica Kautsky, 1925. D. Mioc. v. Hemmoor u. Busbeck-Osten, 73: lám. 6, figs. 27, 28.

Material examinado: Más de 500 c.

Descripción: Concha muy pequeña

(Figs. 62-65), algo cónica. Protoconcha

(Fig. 66) obtusa, del tipo C, con un diá-

metro de unas 270 um. Escultura axial

de la teloconcha formada por entre 20 y

30 costillas, delgadas, flexuosas, algo

más anchas que sus interespacios. Escul-

tura espiral formada por 1-2 cordones

en las vueltas anteriores y 2-3 en la

última, muy juntos y situados justo

sobre la sutura. Abertura oval, con un

pliegue columelar débil y atrasado.

Dimensiones medias: 1,8 x 0,7 mm.

Distribución cronoestratigráfica y geográ-

fica: Mioceno medio: Cuenca del Mar del

Norte alemán y formación Arnum de

Dinamarca (SORGENFREI, 1958). Mioceno

medio (Langhiense): La Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

El gran número de ejemplares estu-

diados nos ha permitido observar que se

trata de una especie muy variable, tanto

en el perfil de las vueltas, como en la

mayor o menor flexuosidad de las

mismas, O la más o menos conspicua

escultura espiral, a veces obsoleta. La

forma extrema sería la ilustrada en las

Figuras 64 y 65, más cercana a C. longula

(Boettger, 1906 sensu Cossmann, 1921).

Chrysallida sp. (Figs. 67, 68)

Material examinado: 2 c.

Descripción: Concha (Figs. 67, 68)

muy pequeña, subcilíndrica. Protocon-

cha obtusa, del tipo C. Teloconcha con

unas 5 vueltas ligeramente convexas.

Sutura profunda. Escultura axial

formada por unas 18 costillas muy fle-

150

xuosas, de igual anchura que sus inte-

respacios; unas y otros se interrumpen

brúscamente en la periferia de la última

vuelta. Sin escultura espiral. Abertura

oval, con un débil pliegue columelar.

Dimensiones: 2,4 x 0,7 mm.

MORENO ET 4L.: Fauna malacológica miocénica de La Pedrera

500 um

Figuras 51, 52. Nassarius schoennií. Conchas. Figuras 53-55. Trigonostoma scrobiculatum. 53, 54:

conchas, 22,7 mm; 55: protoconcha. Figura 56. Strioterebrum basteroz. Concha, 11,9 mm. Figuras

57-60. Clavatula spinosa. 57-59: conchas, 24,4 y 25,0 mm, Can Rosell; 60: protoconcha.

Figures 51-52. Nassarius schoenni. Shells. Figures 53-55. Trigonostoma scrobiculatum. 53, 54:

shells, 22.7 mm; 55: protoconch. Figure 56. Strioterebrum basteroi. Shell, 11.9 mm. Figures 57-60:

Clavatula spinosa. 57-59: shells, 24.4 and 25.0 mm, Can Rosell; 60: protoconch.

Iberus, 21 (1), 2003

Comentarios: Esta especie tiene un

gran parecido con la pliocénica y actual

C. sigmoidea (Monterosato, 1880), pero

ésta última es mayor, tiene estrías espi-

rales y carece de pliegue columelar.

Chemnitzia curvicostata S. Wood, 1848,

descrita para el Crag británico, es muy

parecida pero tiene un perfil más cónico

y carece de pliegue columelar así como

de escultura espiral, aunque WooD

(1848) reconoce que su material es de ma-

la calidad. Pavia (1976), que la cita para el

Plioceno italiano de Monteu Roero, la

describe con finísimas estrías espirales.

Eulimella iusta nom. nov.

Eulimella concinna Sorgenfrei, 1958 (non Odostomia praelonga concinna Jeffreys, 1884). Geol. Surv.

Denmark, IL, 79: 321, lám. 69, figs. 238a, b; lám. 70, fig. 238.

Material examinado: 1 c no adulta.

Etimología: El nombre especifico proviene de la palabra latina ¡ustus, que significa justo, legítimo.

Descripción: En SORGENFREI (1958).

La protoconcha de Eulimella iusta es

muy característica: del tipo B, planispi-

ral, relativamente grande, que sobresale

del perfil de la primera vuelta de telo-

concha. Este tipo de protoconcha es

común en varias especies que viven en

aguas cálidas del África Occidental,

pero no se encuentra en especies actua-

les del Mediterráneo y Atlántico

europeo.

Dimensiones: 2 x 0,6 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Cuenca del Mar

del Norte danés, formación Arnum

(SORGENFREI, 1958). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Esta es la primera cita

para el Mioceno de la Península Ibérica.

En la descripción de Odostomia prae-

longa, JEFFREYS (1884: 350) menciona que

esta especie podría ser incluida en el

género Eulimella y previamente dentro

de esta descripción comenta lo

siguiente: “From the Adventure Bank

Station I have an imperfect specimen and

some fragments of apparently another

species allied to the present. It was a thick

shell with an angular periphery and a

straight pillar, a slighter suture, and a

smaller tooth. If more specimens should be

found this may be named concinna.”

WARÉN (1980) refiere este taxon como O.

praelonga var. concinna, aunque en la

descripción de Jeffreys se hace referen-

cia a una especie y no a una variedad;

además, menciona sus caracteres dife-

renciales de una forma clara por lo que

puede ser considerada una descripción

válida, aunque limitada a un único

ejemplar. Posteriormente, el nombre

Eulimella praelonga se considera nombre

no válido por estar ocupado por Odosto-

mia praelonga Deshayes, 1861, y por

tanto Eulimella cerullii (Cossmamn, 1915)

sería el nombre válido para la especie.

Pero el nombre concinna, descrito antes

de 1961 y sin que su autor le diese

expresamente categoría infrasubespecí-

fica, debe ser considerado válido incluso

aunque el material tipo no haya apare-

cido, siendo además evidente que se

trata de una Eulímella. NORDSIECK (1972)

menciona el taxon Eulimella concinna

como sinónimo de E. pyramidata (Desha-

yes, 1864) y SABELLI, GIANUZZI-SAVELLI

Y BEDULLI (1990) como sinónimo de E.

praelonga. Por todas las mencionadas

razones, el nombre de Eulimella concinna

debe considerarse utilizado y E. con-

cinna Sorgenfrei, 1958 debe ser reempla-

zado.

Eulimella penedesensis spec. nov. (Figs. 69-73)

Material tipo: Holotipo (Fig. 69) y 3 paratipos depositados en el MNCN (n2? MNCNL- 37745, y 37746-

48). Otros paratipos: MGSB (3), MVV (3) y MMP (3).

152

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

ís

; d

:. E

Figura 61. Pyramidella unisulcata. Concha, 3,1 mm. Figuras 62-66. Chrysallida cimbrica. 62-65:

conchas, 2,1, 1,8, 1,7 y 1,5 mm; 66: protoconcha. Figuras 67, 68. Chrysallida sp. Conchas, 2,4 y

1,5 mm.

Figure 61. Pyramidella unisulcata. Shell, 3.1 mm. Figures 62-66. Chrysallida cimbrica. 62-65: shells,

2.1, 1.8, 1,7 and 1.5 mm; 66: protoconch. Figures 67, 68. Chrysallida sp. Shells, 2.4 and 1.5 mm.

153

Iberus, 21 (1), 2003

Otro material examinado: 40 c.

Localidad tipo: Mioceno medio (Langhiense), La Pedrera, Sant Llorenc d'Hortons (Barcelona).

Etimología: El nombre específico hace referencia a la comarca, El Alt Penedes, donde está ubicado

este yacimiento.

Descripción: Concha (Figs. 69-71)

muy pequeña pero sólida, subcilíndrica.

Protoconcha (Figs. 72, 73) del tipo A-Il,

pequeña, bastante obtusa pero con la

espira de 1,5 vueltas casi totalmente

emergida, con un diámetro de 200 um.

Espira de la teloconcha no muy elevada,

con 5-6 vueltas algo convexas, la última

oval redondeada, h = 50% H. Relación

H/D = 3. Sin otra escultura que la

formada por las líneas de crecimiento

(Figs. 72, 73), que es bien conspicua y

característica: a partir de la sutura infe-

rior las líneas de crecimiento son muy

arqueadas y opistoclinas hasta un tercio

antes de la sutura superior, donde pasan

a ser muy prosoclinas, también arquea-

das; el cambio de dirección de las líneas

de crecimiento forma una especie de

cinturón, visible en la mayoría de ejem-

plares. Abertura oval-piriforme. Colu-

mela arqueada, opistoclina, replegada

sobre sí misma. Sin diente ni pliegue

columelar.

Dimensiones del holotipo: 2,6 x 0,9 mm

en 6 vueltas de teloconcha.

Discusión: Esta especie tiene un

cierto parecido con Eulimella (Liostomia)

hydrobiopsis Cossmann y Peyrot, 1917

del Mioceno inferior (Aquitaniense)

francés, sin embargo esta especie tiene

la superficie lisa, incluso a gran

aumento, mientras que en E. penedesen-

sis se observa a simple vista la curiosa

escultura, salvo en ejemplares muy

deteriorados; pero incluso en ellos se

puede apreciar una especie de cinturón

o banda oscura que separa las líneas de

crecimiento de muy opistoclinas a muy

prosoclinas.

No conocemos ninguna especie fósil

o actual, de los mares europeos y oestea-

fricanos, con esa escultura tan caracte-

rística.

Eulimella sp. 1 (Fig. 74)

Material examinado: 1 c.

Descripción: Concha muy pequeña,

cónica regular. Protoconcha del tipo A-

IT, con el núcleo emergido y con un diá-

metro de unas 280 um. Teloconcha de

espira elevada, con 5 vueltas planas, de

crecimiento lento en altura, la última

muy angulosa en la periferia. Sutura

somera, pero bien marcada. Abertura

pequeña, romboide. Profundamente

umbilicada.

Dimensiones: 2,1 x 0,9 mm.

Comentarios: Se trata de una concha

muy parecida a S. hoernesi (ver Figs. 75-

77) de la que se diferencia por tener un

claro y profundo ombligo; y carecer apa-

rentemente de pliegue columelar.

Eulimella sp. 2

Material examinado: 1 c, juvenil.

Comentarios: El ejemplar encontrado,

bastante deteriorado, es un juvenil

formado por la protoconcha y una sola

vuelta de teloconcha, insuficiente para

154

su descripción; sin embargo la protocon-

cha se asemeja en su forma y dimensio-

nes a la de E. unifasciata (Forbes, 1844)

viviente en el Mediterráneo y Atlántico.

MORENO £7 4L.: Fauna malacológica miocénica de La Pedrera

200 um

Figuras 69-73. Eulimella penedesensis. 69: holotipo, 2,6 mm; 70, 71: conchas, 1,6 y 1,4 mm; 72,

73: protoconchas y detalle de la escultura. Figura 74. Eulimella sp. Concha, 2,1 mm. Figuras 75-

77. Syrnola hoernesí. 75: concha, 2,8 mm; 76: protoconcha; 77: detalle de la columela.

Figures 69-73. Eulimella penedesensis. 69: holotype, 2.6 mm; 70, 71: shells, 1.6 and 1.4 mm; 72,

73: protoconchs and detail of the sculpture. Figure 74. Eulimella sp. Shell, 2.1 mm. Figures 75-77.

Syrnola hoernesi. 75: shell, 2.8 mm; 76: protoconch; 77: detail of the columella.

Iberus, 21 (1), 2003

Syrnola hoernesi (Koenen, 1882) (Figs. 75-77)

Turbonilla hoernesi Koenen, 1882. Abhan. Geol. Spec. Preussen Thuringischen Staaten, 2: 263, lám.

6, fig. 1.

Material examinado: 10 c.

Descripción: En GLIBERT (1949,

1952b), ANDERSON (1964), NORDSIECK

(1972) y JANSSEN (1972, 1984).

Concha (Fig. 75) cónica, pequeña.

Protoconcha (Fig. 76) del tipo A-IL, con

un diámetro de 250 ym. Espira elevada.

Sutura estrecha, pero muy profunda.

Líneas de crecimiento ortoclinas. Aber-

tura con un fuerte pliegue columelar

(Fig. 77).

Dimensiones: 2,8 x 0,8 mm en 6

vueltas de la teloconcha.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Helveciense de

la Cuenca del Loira, Francia y Anver-

siense belga (GLIBERT, 1949, 1952);

Cuenca del Norte y Oeste alemán

(ANSERSON, 1964); Winterswijk-Miste,

Holanda (NORDSIECK, 1972 y JANSSEN,

1984). Mioceno medio (Langhiense): La

Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

GLIBERT (1952b) compara esta

especie con E. neumayeri (Koenen, 1882),

de la que se distingue por su mayor

tamaño, vueltas más numerosas, eleva-

das y bien planas, su perfil más regular-

mente cónico y estrecho.

Ver comentarios en Eulimella sp. 1.

Syrnola laevissima (Bosquet, 1859) (Figs. 78, 79)

Turbonilla laevissima Bosquet, 1859. Natuurk. Verh. Akad. Amsterdam, 7: 18, lám. 2, figs. 5a-c.

Material examinado: Más de 150 c.

Descripción: En JANSSEN (1963) y R.

JANSSEN (1979).

Concha (Fig. 78) alargada, pupoide,

muy pequeña. Protoconcha (Fig. 79) del

tipo A-I[, con un diámetro de 220 qm, con

el núcleo semisumergido. Teloconcha de

espira poco elevada, las vueltas casi

planas, la última oval alargada, aproxi-

madamente la mitad de la altura total de

la concha. Sutura somera, muy inclinada.

Líneas de crecimiento ortoclinas. Abertura

pequeña, romboide, con el peristoma con-

tinuo en ejemplares adultos. Columela

opistoclina, con un diente columelar

oblicuo, bien conspicuo y adelantado.

Dimensiones medias: 2,7 x 0,75 mm en

5 vueltas de la teloconcha.

Distribución cronoestratigráfica y geo-

gráfica: Oligoceno de Belgica, Holanda y

Alemania. Mioceno medio (Langhiense):

La Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Ver comentarios en Syrnola sp.

Syrnola sp. (Fig. 80)

Material examinado: 6 c.

Descripción: Concha cónica, estrecha,

alargada. Protoconcha del tipo A-II, con

una vuelta del núcleo emergida en su

totalidad. Teloconcha con las vueltas

156

plano-cóncavas. Sutura estrecha, somera,

muy inclinada. Abertura pequeña, estre-

cha. Columela replegada sobre sí misma,

con un diente columelar muy oblícuo.

MORENO ET A4L.: Fauna malacológica miocénica de La Pedrera

Figuras 78, 79. Syrnola laevissima. 78: concha, 2,5 mm; 79: protoconcha. Figura 80. Syrnola sp.

Concha, 2,8 mm. Figuras 81, 82. Odostomia degrangez. 81: concha, 2,3 mm; 82: protoconcha. Figura

83. Odostomia aft. desuefacta. Concha, 1,4 mm. Figuras 84-87. Odostomia vera. Conchas, 1,0, 1,3,

1,0 y 1,3 mm..

Figures 78, 79. Syrnola laevissima. 78: shell, 2.5 mm; 79: protoconch. Figure 80. Syrnola sp. Shell, 2.8

mm. Figures 81, 82. Odostomia degrangei. 81: shell, 2.3 mm; 82: protoconch. Figure 83. Odostomia

aff. desuefacta. Shell, 1.4 mm. Figures 84-87, Odostomia vera. Shells, 1.0, 1.3, 1.0 and 1.3 mm.

157

Iberus, 21 (1), 2003

Dimensiones: 2,8 x 0,7 mm en 6

vueltas de la teloconcha.

Comentarios: JANSSEN (1984: 356, lám.

16, figs. 3a-c) describe una Syrnola sp. de

similares características.

Especie parecida a S. laevissima, pero

la espira de Syrnola sp. es más elevada,

el perfil más cónico, las vueltas son

plano-cóncavas y la protoconcha es

mayor y con la espira emergida.

Megastomia pallidaeformis (Sacco, 1892)

Odontostomia pallidaeformis Sacco, 1892. I. Moll. ter. terz. Piem. e Lig., 11: 35, lám. 1, fig. 70bis.

Material examinado: 7 c.

Descripción: En Sacco (1892) y en

COSSMANN Y PEYROT (1917). Representa-

ción del holotipo en FERRERO MORTARA

ET AL. (1984, lám. 10, fig. 7).

Dimensiones medias: 2,7 x 1,2 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Colli turinés,

Italia (SAcco, 1892); Helveciense de

Sallespisse, Saucats y Orthez, Francia

(COSSMANN Y PEYROT, 1917); Cuenca del

Loira, Francia (GLIBERT, 1952); Winters-

wijk-Miste, Holanda (NORDSIECK, 1972).

Mioceno medio (Langhiense): La

Pedrera, Barcelona. Mioceno superior:

Tortoniense de Cacella, Portugal

(PEREIRA DA COSTA, 1866).

Comentarios: Ésta es la primera cita

para el Mioceno medio de la Península

Ibérica.

JANSSEN (1972) considera que la des-

cripción e ilustración de SORGENFREI

(1958) no corresponde a esta especie

sino que se trata de O. aff. mutinensis

(Sacco, 1892).

Odostomia degrangei (Cossmann y Peyrot, 1917) (Figs. 81, 82)

Odontostomia (Auristomia) degrangei Cossmann y Peyrot, 1917. Conch. Néog. de l' Aquitaine: 330,

lám. 10, figs. 50, 51.

Material examinado: 8 c.

Descripción: En COSSMANN Y PEYROT

(1917):

Concha (Fig. 81) muy pequeña, oval

cónica. Protoconcha (Fig. 82) del tipo B,

en forma de casquete, con un diámetro

de 220 ym. Espira de la teloconcha corta,

h = 70% H, con las vueltas convexas, la

última grande, oval-redondeada. Sutura

profunda. Superficie sin escultura, salvo

las líneas de crecimiento que son ortocli-

nas a ligeramente prosoclinas. Abertura

grande, oval. Columela arqueada, opis-

toclina, algo replegada sobre sí misma.

Sin diente ni pliegue columelar visibles.

No umbilicada.

Dimensiones: 2,5 x 1mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Burdigaliense

y Mioceno medio: Orthez, Francia

(COSSMANN Y PEYROT, 1917). Mioceno

medio (Langhiense): La Pedrera, Barce-

lona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Odostomia aff. desuefacta Peñas y Rolán, 1999 (Fig. 83)

Odostomia desuefacta Peñas y Rolán, 1999. Iberus, suplemento 5: 52-54, figs. 117-119.

Material examinado: 1 c.

158

MORENO £7 ALz.: Fauna malacológica miocénica de La Pedrera

Descripción: Concha diminuta, cónica

escalonada. Protoconcha más bien

obtusa, del tipo A-IT, con un diámetro de

unas 200 ym, con la espira de 1,0 vueltas

sumergida más de la mitad. Teloconcha

con la espira muy corta, h > 65% H, con

unas 3 vueltas convexas bajo la sutura y

planas sobre ella, escalonadas. Sutura

estrecha pero profunda, con un estrecho

hombro bajo ella. Abertura pequeña,

oval, estrecha. Peristoma continuo.

Columela opistoclina, con pliegue colu-

melar conspicuo, muy oblicuo.

Dimensiones: 1,0 x 0,5 mm.

Comentarios: Esta especie tiene un

gran parecido con O. desuefacta Peñas y

Rolán, 1999, que vive en aguas del África

Occidental, en su forma escalonada, tipo

de protoconcha, peristoma continuo y

pliegue columelar oblicuo. Sin embargo,

O. desuefacta es más alargada, H/D = 3,

frente a 2,1 en la especie fósil; las vueltas

crecen más deprisa, la repisa subsutural

es angulosa y la abertura es mayor.

O. sempert (Bosquet, 1859), del Oligo-

ceno holandés, también tiene las vueltas

escalonadas, pero su perfil es más

cónico, sus vueltas crecen más deprisa

en anchura, el ápice es más agudo, la

abertura mayor y el diente columelar es

perpendicular a la columela. Ver ilustra-

ción de esta especie en JANSSEN (1963).

Odostomia vera nom. nov. (Figs. 84-87)

Chrysallida modesta Sorgenfrei, 1958. Geol. Surv. Denmark, 5, 79: 309, lám. 67, figs. 228a, b. (non

Odostomia modesta Stimpson, 1851).

Material examinado: 10 c.

Etimología: El nombre especifico proviene de la palabra latina verus que significa verdadero.

Descripción: En SORGENFREI (1958).

Concha (Figs.84-87) minúscula, sub-

cilíndrica. Protoconcha obtusa, del tipo

C. Unas 4 vueltas de teloconcha conve-

xas, la última oval redondeada. Escul-

tura axial obsoleta formada por las

líneas de crecimiento que son flexuosas.

Escultura espiral formada por finísimas

estrías en toda la vuelta entre suturas.

Abertura oval. Sin diente columelar,

sólo se observa un muy débil pliegue

columelar.

Dimensiones medias: 0,8 x 0,3 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Cuenca del Mar

del norte danés, formación Arnum (Sor-

GENFREI, 1958). Mioceno medio (Lang-

hiense): La Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Creemos que esta especie, descrita

como Chrysallida modesta Sorgentfrei,

1958, debe ubicarse en el género Odosto-

mia, ya que carece de escultura axial

conspicua. La fina escultura espiral es

común en muchas especies de este

género. Por tanto, al ser encuadrada en

el género Odostomia se produce una

sinonímia con Odostomia modesta Stimp-

son, 1851, especie actual del Atlántico

americano. Por ese motivo el nombre

está ocupado y debe ser ser reempla-

zado.

El ejemplar de la figura 84 presenta

un tamaño algo mayor, y las estrías espi-

rales son más conspicuas y espaciadas;

sin embargo, al no haber encontrado

más material para comparar, creemos

que esas diferencias pueden formar

parte del rango de variabilidad.

Odostomia perstricta (Sacco, 1892)

Odontostomia (Macrodostomia) perstricta Sacco, 1892. I Moll. ter. terz. Piemonte e Liguria, 11: 44, lám.

1, fig. 98bis.

Material examinado: 1 c.

159

Iberus, 21 (1), 2003

Descripción: FERRERO MORTARA ET AL.

(1984: lám. 9, figs. 10a, b) ilustran el

holotipo de esta especie.

Concha pequeña, alargada, conoidea,

que parece una Syrnola. Apice agudo.

Teloconcha con las vueltas planas a ligera-

mente convexas. Sutura somera. Abertura

suboval con un pliegue columelar oblicuo.

Dimensiones: 2,5 x 1 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Colli turinés,

Italia (SACCO. 1892). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibé-

rica.

Odostomia turritella (Grateloup, 1828) (Figs. 88-92)

Auricula turritella Grateloup, 1828. Bull. Hist. Nat. Soc. Lin. Bordeaux, 2 (9): n* 72, 105.

Acteon bulimoides Grateloup, 1838. Actes Soc. Lin. Bordeaux, 10, 53: 275, lám. 6, figs. 44, 45.

Material examinado: 150 c.

Descripción: En COSSMANN Y PEYROT

(1917). Ilustración en LOZOUET ET AL. (2001).

Concha (Figs. 88-91) muy pequeña,

oval-conidea. Protoconcha (Fig. 92) obtusa,

del tipo C. Teloconcha con las vueltas poco

convexas, la última oval-alargada. Líneas

de crecimiento ortoclinas. Abertura oval,

aguda hacia arriba. Columela opistoclina

replegada sobre sí misma, formando a

menudo una fisura umbilical. Diente colu-

melar inclinado, bien conspicuo.

Dimensiones medias: 3 x 1 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno inferior: Aquitaniense y

Burdigaliense francés (COSssSMANN Y

PeYrRO1,1917 y LOZOUET EN AO

Mioceno medio (Langhiense): Depresión

del Vallés (VILLALTA ET AL., 1968); La

Pedrera, Barcelona.

Comentarios: LOZOUET ET AL. (2001)

retoman el nombre más antiguo atri-

buido a Grateloup, pues contrariamente

a lo que opinan COSSMANN Y PEYROT

(1917) “turritella” Grateloup, 1828 nunca

pudo estar pre-ocupada por “turrite-

llata” Deshayes, por ser un nombre dife-

rente y además haber sido descrita en

1864 (1).

Odostomia sp.1

Material examinado: 1 c, ligeramente inmadura.

Descripción: Concha diminuta

tronco-cónica. Protoconcha del tipo A-II,

con una espira de dos vueltas, promi-

nente, que sobresale bastante del perfil

de la primera vuelta de teloconcha.

Vueltas planas, la última angulosa en la

periferia. Sutura somera. Abertura rom-

boide, con un diente columelar conspi-

cuo, perpendicular a la columela.

Dimensiones: 0,8 x 0, 4 mm.

Comentarios: Esta especie es muy

parecida a O. verhoeveni Aartsen, Gitten-

berger y Goud, 1998, especie actual que

vive en aguas del Africa Occidental, de

la cual se diferencia casi únicamente en

que ésta tiene una sutura profunda y

canaliculada (ver ilustración en PEÑAS Y

ROLÁN (1999).

Odostomia sp. 2 (Fig. 93)

» Material examinado: 1 c.

Descripción: Concha pequeña,

cónico-alargada, que parece una Syrnola.

160

Ápice agudo con una protoconcha

pequeña, del tipo B. Espira de la telo-

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

Figuras 88-92. Odostomia turritella. 88-91: conchas, 2,8, 2,2, 2,4 y 2,5 mm; 92: protoconcha.

Figura 93. Odostomia sp. 2. Concha, 2,9 mm. Figuras 94-96. Noemiamea batlloriz. 94: Concha, 1,6

mm; 95: holotipo, 1,3 mm; 96: protoconcha del holotipo.

Figures 88-92. Odostomia turritella. 88-91: shells, 2.8, 2.2, 2.4 and 2.5 mm; 92: protoconch. Figure

93. Odostomia sp. 2. Shell, 2.,9 mm. Figures 94-96. Noemiamea batllorii. 94: Shell, 1.6 mm; 95:

holotype, 1.3 mm; 96: protoconch of the holotype.

161

Iberus, 21 (1), 2003

concha elevada, con las vueltas planas,

la última muy angulosa en la periferia, h

< 50% H. Sutura somera. Sin escultura

aparente. Abertura romboide, con un

diente columelar estrecho, pero conspi-

cuo, perpendicular a la columela.

Ombligo estrecho pero profundo.

Dimensiones: 2,8 x 1,2 mm.

Comentarios: Esta especie tiene un

gran parecido con Syrnola lanotensis

(Lozouet, 1998) del Mioceno inferior

(Aquitaniense), pero ésta última tiene

una concha mayor y el pliegue colume-

lar oblicuo.

Ondina sp.

Material examinado: 1 c juvenil.

Comentarios: La protoconcha de esta

especie es cercana a la especie actual O.

obligua (Alder, 1844), pero no es posible

describir la concha ni comparararla

porque se trata de un juvenil con una

sola vuelta de teloconcha

Noemiamea batllorii spec. nov. (Figs. 94-96)

Material tipo: Holotipo (Fig. 95) depositado en el MNCN (n? MNCNI- 37749). Paratipos en las

siguientes colecciones: MVV (1) y MGSB (1).

Otro material estudiado: 5 c (Fig. 94) (una de ellas perdida durante el estudio).

Localidad tipo: Mioceno medio (Langhiense), La Pedrera, Sant Llorenc d'Hortons, (Barcelona).

Etimología: El nombre específico se dedica al Dr. Jordi Batllori, de la Universidad de Barcelona,

por su colaboración en la realización de este trabajo.

Descripción: Concha (Figs. 94, 95) di-

minuta, ovoide. Protoconcha (Fig. 96) del

tipo B, en forma de casquete. Espira de la

teloconcha muy corta, con 2,0-2,5 vueltas

que son ligeramente convexas, algo esca-

lonadas, de crecimiento muy rápido, la

última oval, muy grande, h = 90% H. Su-

tura estrecha pero profunda. Escultura es-

piral formada por numerosas estrías equi-

distantes, unas 10 entre las suturas de la

última vuelta, siendo más numerosas y

apretadas en la base. Sin escultura axial.

Abertura oval, muy grande, > 70% H. Pe-

ristoma continuo. Columela arqueada, con

un pliegue oblicuo, atrasado, conspicuo,

aunque no prominente. No umbilicada.

Una de las conchas tiene la espira

más corta y las estrías espirales son más

numerosas, pero una única concha nos

parece insuficiente material para darle

nombre.

Dimensiones del holotipo: 1,3 x 0,7 mm.

Dimensiones máximas: 1,6 x 0,9 mm.

Discusión: No hemos encontrado

ningún género en el que esta especie

pudiese ser claramente ubicada y

creemos que Noemiamea es el más

162

cercano, por sus caracteres genéricos:

concha muy pequeña, globular-ovoide,

con 2-3 vueltas de crecimiento rápido y

perfil algo escalonado, con escultura

espiral conspicua y sin escultura axial.

Por otra parte, presenta diferencias

con las siguientes especies afines:

N. dolioliformis (Jeffreys, 1848), que

vive en el Atlántico europeo y en el

Mediterráneo, es más globosa, tiene la

protoconcha del tipo A, es umbilicada y

tiene un diente columelar prominente.

Acteopyramis merignacensis Cossmann

y Peyrot, 1917, del Mioceno inferior (Bur-

digaliense y Aquitaniense) francés, tiene

una concha mayor, con la espira más

elevada, la protoconcha es obtusa, del tipo

C y tiene la escultura espiral formada por

surcos profundos más espaciados.

Acteopyramis elatus (Von Koenen, 1882),

del Oligoceno alemán y Kleinella ivolasi

(Mayer, 1900) del Mioceno medio belga y

holandés, tienen la concha mayor, con la

espira más elevada, el ápice obtuso y

tienen una escultura espiral diferente, no

formada por estrías sino por surcos más

anchos con estrías verticales en su interior.

MORENO £7 AL.: Fauna malacológica miocénica de La Pedrera

Turbonilla olivellai spec. nov. (Figs. 99-101)

Material tipo: Holotipo (Fig. 99) y dos paratipos, depositados en el MNCN (n*% MNCNLI- 37750 y

37751-2). Otros paratipos: MMP (2), MGSB (3), y MVV (2).

Otro material examinado: 20 c.

Localidad tipo: Mioceno medio (Langhiense), La Pedrera, Sant Llorenc d'Hortons, (Barcelona).

Etimología: El nombre específico se dedica a Rosendo Olivella, paleontólogo del Museu del Vi de

Vilafranca del Penedes, por poner a nuestra disposición la colección paleo-malacológica del Museo.

Descripción: Concha (Figs. 99, 100) muy

pequeña, cónica. Protoconcha (Fig. 101)

algo obtusa, del tipo A-IL, con un diáme-

tro de 250 ym, y una espira de 1,5 vueltas,

emergida. Unas 5-6 vueltas de teloconcha

casi planas, algo angulosas en la zona

sutural, la última angulosa en la periferia,

h = 40% H. Sutura estrecha pero profunda,

inclinada. Escultura axial no varicosa,

formada por unas 12 costillas rectas, opis-

toclinas, delicadas, algo más estrechas que

sus interespacios; unas y otros se atenuan

hasta desaparecer en la periferia de la

última vuelta. Sin escultura espiral. Aber-

tura pequeña, romboide. Columela recta,

vertical, sin pliegue aparente.

Dimensiones del holotipo: 2,1 mm x 0,7

mm. en 5 televueltas.

Discusión: Concha parecida a T.

obliqua Degrange-Touzin, 1895, del

Mioceno Helveciense y Burdigaliense

francés, y del Crag británico, pero ésta

tiene un mayor tamaño, la espira más

elevada con vueltas más convexas y

estrechas, las costillas más robustas, la

sutura más profunda y ondulada, y

tiene un pliegue columelar.

Nuestra especie es cercana a T. incog-

nita Degrange-Touzin, 1895, del

Mioceno inferior (Aquitaniense) francés,

que fue insuficientemente descrita en

base a un solo fragmento sin ápice ni

primeras vueltas, y redescrita por Coss-

MANN Y PEYROT (1917: 352, lám. 10, figs.

72, 73). Se trata de una especie mucho

mayor (H = 9 mm), con las vueltas aún

más planas, la sutura más somera y las

costillas más robustas, casi ortoclinas.

T. dertogracilis (Sacco, 1892), del

Mioceno superior (Tortoniense) italiano

y húngaro tiene una concha mayor, las

costillas axiales son más anchas y obso-

letas, algunas varicosas, que se desvane-

cen antes de la periferia de la última

vuelta, la cual es redondeada; y tiene

cordoncillos espirales visibles en el inte-

rior del labro externo.

T. astensidelicata Sacco, 1892, del Plio-

ceno medio-superior italiano, tiene el

ápice más agudo, la espira más elevada,

y un mayor número de costillas (16-20),

las cuales son robustas y más anchas

que sus interespacios.

Tanto T. costellatoides Sacco, 1892, del

Mioceno medio-superior italiano y del

Plioceno italiano, como T. costellata Gra-

teloup 1827, del Mioceno inferior (Aqui-

taniense y Burdigaliense) francés y del

Mioceno medio de Dinamarca y Alema-

nia, tienen una concha mucho mayor,

con las vueltas bien convexas y la sutura

más profunda, ambas tienen pliegue

columelar y T. costellata tiene, además,

una especie de cordón subsutural.

Turbonilla plivmagna Sacco, 1892

Turbonilla postacuticostata? var. pliomagna Sacco, 1892. I Moll. ter. terz. Piem. e Lig., 11: 76, lám. 2,

fig. 60.

Material examinado: 1 c, no adulta, 1 f.

Descripción: En SACco (1892). Ilustra-

ción en FERRERO MORTARA ET AL. (1984:

lám. 12, figs. 7a, 7b).

Dimensiones: 2,8 x 0,7 mm.

Distribución cronoestratigráfica y geo-

gráfica: Plioceno de Masserano, Italia

(Sacco, 1892). Mioceno medio (Lang-

hiense): La Pedrera, Barcelona.

163

Iberus, 21 (1), 2003

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica

Según MICALI (com. per.) esta especie

debe ser redescrita, ya que la considera

válida y no una variedad de T. postacuti-

costata Sacco, 1892 nom. nov. pro T. acu-

ticostata Jeffreys, 1884 non O. Speyer,

1870. No realizamos la redescripción en

este trabajo debido al escaso material

disponible.

Turbonilla sallomacensis Cossmann y Peyrot, 1917 (Figs. 97, 98)

Turbonilla (Pyrgolidium) sallomacensis Cossmann y Peyrot, 1917. Conch. Néog. de I' Aquitaine: 364,

pl. 10, fig. 89, 90.

Material examinado: 3 c.

Descripción: Concha (Fig. 97) cónica,

estrecha. Protoconcha (Fig. 98) pequeña,

del tipo B, planispiral, con un diámetro de

unas 200 um. Teloconcha con las vueltas

casi planas, que crecen rápidamente.

Numerosas costillas axiales, rectas, orto-

clinas, delicadas, atenuadas bajo la sutura,

más anchas que sus interespacios. Unas y

otros se prolongan en la base más allá de

la periferia. Sin escultura espiral. Aber-

tura pequeña, romboide. Columela algo

opistoclina, replegada.

Dimensiones: 2 x 0,7 mm.

Distribución cronoestratigráfica y geo-

gráfica: Mioceno medio: Helveciense de

Salles, Francia (COSSMANN Y PEYROT,

1917). Mioceno medio (Langhiense): La

Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Turbonilla sp.1

Material examinado: 1 c

Descripción: Concha muy pequeña,

subcilíndrica. Protoconcha del tipo A-IL,

con un diámetro de unas 230 ym, con

una espira de 1,5 vueltas totalmente

emergida y prominente. Teloconcha con

las vueltas apenas convexas. Sutura

somera. Escultura axial formada por

unas 20 costillas estrechas, algo arquea-

das, que se atenuan en la periferia de la

última vuelta hasta desaparecer. Nume-

rosas estrías espirales visibles con

aumento. Abertura subcuadrangular.

Columela vertical, sin pliegue visible.

Dimensiones: 2,4 x 0,6 mm.

Comentarios: Esta especie tiene gran

parecido con T. templadoi Peñas y Rolán,

1997, que vive en el África Occidental,

en su tamaño, forma, tipo de protocon-

cha y microescultura espiral. Se diferen-

cian en que ésta última tiene las vueltas

más convexas, la sutura más profunda y

la escultura espiral más conspicua.

Turbonilla sp. 2 (Figs. 102, 103)

Material examinado: 1 c.

Descripción: (Concha (Fig. 102)

pequeña, conoidea, algo pupoide. Proto-

concha (Fig. 103 ) del tipo B, con un diá-

metro de unas 200 ym, con el núcleo y

más de media vuelta visible. Teloconcha

con las vueltas muy convexas, la última

164

grande, h= 50% H. Sutura muy pro-

funda. Escultura axial formada por unas

14 costillas lamelosas, 3-4 veces más

estrechas que sus interespacios; unas y

otros se prolongan en la base hasta la

zona umbilical. Escultura espiral sola-

MORENO £7 4L.: Fauna malacológica miocénica de La Pedrera

50 um

Figuras 97, 98. Turbonilla sallomacensis. 97: concha, 2,0 mm; 98: protoconcha. Figuras 99-101.

Turbonilla olivellai. 99: holotipo, 2,1 mm; 100: concha, 1,7 mm; 101: protoconcha. Figuras 102,

103. Turbonilla sp. 2. 102: concha, 1,75 mm; 103: protoconcha. Figuras 104-107 Anisocycla cf.

bezanconi. 104, 105: conchas, 1,5 y 1,6 mm; 106: protoconcha; 107: detalle de la escultura.

Figures 97, 98. Turbonilla sallomacensis. 97: shell, 2.0 mm; 98: protoconch. Figures 99-101. Turbo-

nilla olivellai. 99: holotype, 2,1 mm; 100: shell,1,7 mm; 101: protoconch. Figures 102, 103. Turboni-

lla sp. 2. 102: concha, 1.75 mm; 103: protoconcha. Figures 104-107. Anisocycla cf. bezanconi. 104,

105: shells, 1.5 and 1.6 mm; 106: protoconch; 107: detail of the sculpture.

Iberus, 21 (1), 2003

mente en los interespacios, formada por

pocas estrías espirales equidistantes

que, en la base, son mucho más numero-

sas y apretadas. Abertura oval. Colu-

mela opistoclina, muy replegada hacia

la zona umbilical.

Dimensiones: 1,75 x 0,76 mm.

Comentarios: Esta especie pertenece

al grupo de T. jeffreysii (Jeffreys, 1848) y

T. rufescens (Forbes, 1846) por su forma y

escultura axial lamelosa; sin embargo,

difiere de ellas en la forma de la proto-

concha, en la escultura espiral y en la

forma de la columela.

Anisocycla cf. bezanconi Cossmann y Lambert, 1884 (Figs.104-107)

Anisocycla bezanconi (Cossmann y Lambert) sensu Gougerot, 1991. Bull. Nat. Parisiens, 47 (1): 8.

Material examinado: 5 c.

Descripción: Concha diminuta (Figs.

104, 105), ligeramente subcilíndrica,

alargada, muy frágil. Protoconcha (Fig.

106) del tipo B, planispira, con un diá-

metro de unas 180 ym, con el núcleo

visible, y la sutura de separación con la

teloconcha muy angulosa. Espira

elevada, con las vueltas convexas, de

crecimiento regular, la última oval

redondeada. Sutura profunda. Escultura

espiral (Fig. 107) delicada formada por

débiles surcos relativamente equidistan-

tes. Sin escultura axial salvo las líneas

de crecimiento, ligeramente arqueadas,

algo opistoclinas bajo la sutura. Aber-

tura oval. No umbilicada.

Dimensiones: 1,6 x 0,5 mm.

Distribución cronoestratigráfica y geo-

gráfica: Oligoceno de la Cuenca de París

(GOUGEROT, 1991). Mioceno inferior

(Aquitaniense) Saucats, Francia

(LOZOUET ET AL., 2001). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

GOUGEROT (1991) cree que esta

especie, procedente del Oligoceno, muy

probablemente no difiera de A. scalarina

(Deshayes, 1861) del Eoceno parisino e

incluye en el mismo taxon 2 ejemplares

del Aquitaniense de Las Landas. LE

RENARD (1998) designa un neotipo de A.

scalarina, el cual presenta claras diferen-

cias con A. bezanconi. Por otra parte,

LOZOUET ET AL. (2001) ponen en duda la

sinonimia de ambas especies, opinión

que compartimos. Creemos que se trata

de dos especies diferentes: A. bezanconi

sería antecesora de la actual A. nitidis-

sima (Montagu, 1803), mientras que A.

scalarina sería antecesora de A. striatula

(Jeffreys, 1856).

Anisocycla eulimoides Fekih, 1969 (Figs. 108, 109)

Anisocycla eulimoides Fekih, 1969. Notes du Service Geologique, Tunis, 29: 57, lám. 11, fig. 15.

Material examinado: 2 c.

Descripción: En FEKIH (1969).

Mostramos concha (Fig. 108) y pro-

toconcha (Fig. 109), del tipo B, planis-

pira, con la sutura muy inclinada. Resal-

tamos el rápido crecimiento de las

vueltas, tanto en anchura como en

altura, siendo la última muy grande,

con más del 50% de la altura total. Se

aprecia en la zona subsutural que las

líneas de crecimiento son muy numero-

166

sas, prosoclinas, y en la última vuelta se

observa solamente a gran aumento una

tenue escultura espiral.

Dimensiones: 1,35 x 0,45 mm.

Distribución cronoestratigráfica y geo-

gráfica: Plioceno de Oued el Galea,

Túnez (FeEkIH, 1969). Mioceno medio

(Langhiense): La Pedrera, Barcelona.

Comentarios: Esta es la primera cita

para el Mioceno de la Península Ibérica.

—

MORENO £7 4Z.: Fauna malacológica miocénica de La Pedrera

200 um

Figuras 108, 109. Anisocycla eulimoides. 108: concha, 1,3 mm; 109: protoconcha. Figuras 110-

113. Perla. 110: perla, diámetro de 0,69 mm; 111-113: detalle de la estructura laminar.

Figures 108, 109. Anisocycla eulimoides. 108: shell, 1.3 mm; 109: protoconch. Figures 110-113.

Pearl. 110: pearl, diameter of 0.69 mm; 111-113: detail of the laminar structure.

Iberus, 21 (1), 2003

Esta especie, descrita para el Plio-

ceno de Túnez, presenta unas claras

diferencias con el resto de especies de

este género, sean fósiles o vivientes: su

relación H/h = 2 y su relación H/D =

Clathrella sulcosa (Brocchi, 1814)

Nerita sulcosa Brocchi, 1814. Conch. fos. subap., 298, lám. 1, figs. 3a, b.

Material examinado: 1 f.

Comentarios: Ver en PEÑAS Y ROLÁN

(2001) ilustración del holotipo y su

comparación con la especie actual-

mente viviente en aguas del África

Occidental, Clathrella volumen Peñas y

Rolán, 2001.

Clase BIVALVIA

Anadara cf. diluvii (Lamarck, 1805)

Arca diluvii Lamarck, 1805. Ann. Mus. Nation. Hist. Nat. Paris, 219.

Material examinado: 40 c, 220 v.

Dimensiones máximas: 30 x 20 mm.

Distribución cronoestratigráfica y geo-

gráfica: Esta especie ha sido citada en

todas las cuencas europeas desde el Bur-

digaliense hasta la actualidad.

Comentarios: Especie polimorfa.

CALZADA (1982) denominó a este

bivalvo como Scapharca (Cunearca) sp.,

después del estudio de material proce-

dente de Can Rosell, Sant Pau d'Ordal

y La Pedrera. Este autor comparó este

taxon con Anadara fichteli, Scapharca

turonica, Anadara numida y Anadara

diluvii, concluyendo que era diferente a

las especies citadas, aunque recono-

ciendo una cierta similitud con Anadara

diluvii var. pertransversa. Ciertamente, la

especie actualmente viviente presenta

claras diferencias con el fósil miocé-

nico.

Ostrea edulis Linné, 1758 (Figs. 110-113)

En la fracción fina del sedimento

tamizado se halló un diminuto cuerpo

esférico, de aspecto nacarado (Fig. 110),

con un diámetro de 687,5 qm. Creimos

que podía tratarse de una perla produ-

cida probablemente por algún ejemplar

de Ostrea edulis, especie común en el

yacimiento, y se procedió a fotografiarla

al MEB (Figs. 111-113) con 600, 4.810 y

9.621 aumentos. Las fotografías revelan

la estructura concéntrica e irregular de

las capas laminares semiabiertas típica

de las perlas. En las de mayor aumento

se Observa la estructura laminar y pun-

tualmente la forma subexagonal de los

cristales de aragonita característica de las

perlas. Nogués (com. per.) nos confirma

que se trata de una perla, a pesar de

tener su superficie bastante deteriorada.

Ver estructura microscópica de las

capas de la perla en NOGUÉS, VENDRELL

Y ARBUNIES (1980).

Alveidus nitidus (Reuss, 1867) (Figs. 114-120)

Spaniodon nitidus Reuss, 1867. Sitzungsber. D. Kais. Akad. D. Wiss., 55 (1): 135-136, lám. 8, figs. 3a, b.

168

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

100 um d 100 um

Figuras 114-120. Alveidus nitidus. 114: valva derecha; 115: interior de la valva derecha; 116: inte-

rior de la valva izquierda; 117: charnela de la valva derecha; 118: charnela de la valva

izquierda.119: detalle de la charnela de la valva derecha; 120: detalle de la charnela de la valva

izquierda.

Figures 114-120. Alveidus nitidus. 114: right valve; 115: inner part of the right valve; 116: inner

part of the left valve; 117: hinge of the right valve; 118: hinge of the left valve; 119: detail of the hinge

of the right valve; 120: detail of the hinge of the left valve.

Iberus, 21 (1), 2003

Lutetia girondica Benoist in coll. Cossmann y Peyrot, 1912. Actes Soc. Lin. Bordeaux: 610-612, lám.

26, figs. 16-19.

Material examinado: Más de 300 c, más de 5.000 v.

Descripción: En COSSMANN Y PEYROT

(1912) y en A W. JANSSEN (1984). Se ilus-

tran aquí la concha y la charnela en

detalle.

Dimensiones: 1,2 x 1,2 mm.

Distribución cronoestratigráfica y geo-

gráfica: Oligoceno superior (ANDERSON,

1964). Mioceno inferior: Burdigaliense y

Aquitaniense de Francia (COSSMANN Y

PEYROr, 1912). Mioceno medio: Winters-

wijk-Miste, Holanda (JANSSEN, 1984);

Badeniense del sur de Polonia (STU-

DENCKA, 1986). Mioceno medio (Lang-

hiense): La Pedrera, Barcelona. Mioceno

superior: Tortoniense de Saubrigues,

Francia (COSSMANN Y PEYROT, 1912).

Comentarios: Ésta es la primera cita

para el Mioceno de la Península Ibérica.

Se trata de la especie predominante

de este yacimiento, tanto en número de

ejemplares como en volumen de sedi-

mento estudiado. De esta minúscula es-

3. Comentarios finales

Especies previamente citadas en La

Pedrera, no encontradas por nosotros: Se

relacionan seguidamente las especies

citadas anteriormente en este yaci-

miento, indicando a continuación el

autor o autores de la cita en negrita. No

se incluyen las que consideramos sino-

nimias ni los moluscos determinados a

nivel genérico: CRUSAFONT (1959): C;

CASANOVAS, CALZADA Y SANTAFÉ (1972):

CCS; MARTINELL Y PORTA (1981): MP;

BATLLORI (1995): B; NAVAS ET AL. (1996):

Teinostoma degrangei

Pleurotoma monilis

Turritella cestacensis

Bittium reticulatum

Cerithium olivaeformis

Cerithium turonicum

Setla turritellata

Opalia branderburgi

Calyptraea cf. costaria

ZEN AS ZOOZ

HF Y

170

pecie se han contado en la fracción fina

del nivel A cerca de 1.000 valvas por

centímetro cúbico, además de unos dos

ejemplares completos (las dos conchas

unidas) de media y gran cantidad de

fragmentos. Teniendo en cuenta que el

volúmen estudiado de esta fracción es

de unos tres litros y medio, calculamos

que en él se encuentran unos tres millo-

nes de valvas, sin contar los fragmen-

tos. Además, hemos constatado que

más del 85% de esa fracción fina del se-

dimento resultante del lavado está for-

mado por valvas y fragmentos de esta

especie. En el nivel B el volumen de

esta especie es mucho menor que en el

nivel A.

STUDENCKA (1986) discute la ubica-

ción genérica de esta especie.

CASANOVAS ET AL. (1972) la conside-

ran en otro género, como Polimesoda sp.

(Calzada, com. per.).

Pereiraea gervaisi

Xenophora testigera

Euspira protracta

Ficus cingulatus

Ficus conditus

Sinum haliotoideum

Triton abbreviatus

Trunculariopsis turonensis

Murex delbosianus

Murex vindobonensis

Columbella curta

Nassarius borsoni

Nassarius brugnont?

Nassarius elatus

Nassarius hoernessi?

Nassarius orthezensis

Nassarius prismaticus

Nassarius rectus

Babylonia brugadina

Athleta ficulina

Pusionella pseudofusus

Eulimella subumbilicata

Spondylus deshayesi

Thracia dollfusi

Meretrix ericina

OZZWVORBIRZZINO O ZW ZW ZUR ZZZ

MORENO ET AL.: Fauna malacológica miocénica de La Pedrera

Nuevas citas para la Península ibérica y

nuevas especies para la ciencia: Como se

señaló anteriormente, en el listado de

especies se identifican con un asterisco

aquellas que, de acuerdo con la biblio-

grafía, se citan por primera vez para este

yacimiento, que son 29 en total. Con dos

asteriscos se señalan las que son

primera cita para la Península Ibérica,

40 en total. De ellas son nuevas para la

ciencia las siguientes: Obtusella orisparvi,

Caecum verai, Eulimella penedesensis, Noe-

miamea batllorii y Turbonilla olivellas.

Debemos resaltar el gran número de

taxones citados a nivel genérico: 33, bas-

tantes de ellos difícilmente determinables

por la escasez y mal estado de conserva-

ción del material. Sin embargo, varios de

ellos (como Rissoa sp., Bela sp.2, Syrnola sp.

o Venus sp.), de los que se ha encontrado

suficiente material, precisan de una revi-

sión para la cual es necesaria la consulta

de numerosos trabajos muy dispersos,

actualmente no accesibles para nosotros.

Abundancia relativa: Tan solo doce de

las especies encontradas en este yaci-

miento representan más del 95% del

total de ejemplares hallados en nuestros

muestreos. Se relacionan a continuación,

por orden de mayor a menor abundancia

en número de ejemplares: Alveidus

nitidus, Sandbergeria perpusilla, Acteocina

lajonkairena, Nassarius cf. aturensis, Obtu-

sella orisparvi, Potamides theodiscus, Gra-

nulolabium pictum, Hydrobia peregrina;

Chrysallida cimbrica, Nassarius girondicus,

Odostomia turritella y Cyllene vulgatissima.

Como ya se comentaba anteriormente

Alveidus nitidus debe ser considerada la

especie predominante de este yacimiento,

tanto en número de ejemplares como en

volúmen conchífero. Entre los gasterópo-

dos, Sandbergeria perpusilla es la especie

más abundante en número de ejemplares

y Potamides theodiscus la que representa el

mayor volúmen conchífero.

CONCLUSIONES

El presente trabajo confirma la

riqueza malacológica de este yacimiento

de La Pedrera, para el cual, el número

total de especies de moluscos marinos

fósiles conocidos actualmente es de 194

(169 Gasterópodos, 22 Bivalvos y 3 Esca-

fópodos). En este trabajo, recolectadas

por los autores, se citan 160 especies

(138 Gasterópodos, 19 Bivalvos y 3 Esca-

fópodos).

En nuestra opinión la fauna malaco-

lógica hallada pertenece al nivel infrali-

toral somero de un ambiente típico

marino, equivalente a la fauna actual-

mente existente en ese nivel, con una

cierta influencia de agua dulce, dada la

abundante presencia de hidróbidos.

Se constata asimismo la escasa infor-

mación ofrecida sobre micromoluscos,

aunque no de otros microfósiles, en

anteriores trabajos paleontológicos refe-

ridos a este yacimiento y a la Península

Ibérica en general, que es casi nula si se

trata del período miocénico.

AGRADECIMIENTOS

Nuestro agradecimiento a los

siguientes profesores de la Universidad

de Barcelona: al Dr. Jordi Batllori, por su

ayuda en la determinación de algunas

especies y por permitirnos consultar su

colección; al Dr. Carles Martín, que nos

mostró el yacimiento y nos facilitó infor-

mación sobre el mismo; al Dr. Carles Gili

por su ayuda en la revisión de la familia

Nassariidae; al Dr. Jaume de Porta por su

ayuda en la determinación de la familia

Ostreidae; al Dr. Salvador Reguant y a su

ayudante Yael Díaz por la determinación

del briozoo; al Dr. Joaquín María Nogués

por su asesoramiento y ayuda en la iden-

tificación de la perla; a Nuria Martínez

por su ayuda en la determinación de

algunos bivalvos. A los padres del

primero de los autores y a José L.

Cañadas por acompañarnos durante la

recogida de muestras. A Laia Fonolla,

quien nos mostró la zona de estudio. A

Pasquale Micali, de Fano, Italia, por su

ayuda en la clasificación de varias espe-

cies, especialmente de Pyramidellidae, y

en la obtención de bibliografía. Al Dr.

Pierre Lozouet, del MNHN, por su

ayuda y revisión de las familias Potami-

didae y Batillariidae y cesión de mate-

171

Iberus, 21 (1), 2003

rial. Al Dr. Sebastián Calzada, director

del MGSB y al Dr. Antonio Abad por sus

comentarios, sugerencias y ayuda en la

obtención de bibliografía. Al Dr. José

Luis Vera Peláez del Museo Municipal

de Paleontología de Estepona, por su

ayuda y revisión de las familia Turridae

y la clase Scaphopoda. Al Dr. Gonzalo

Giribet, de la Universidad de Harvard,

por el envío de bibliografía. A la Dra.

Bárbara Studenka del Muzeum Ziemi de

Polonia por su ayuda en la determina-

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VERA-PELÁEZ, J.L., LOZANO-FRANCISCO, M.C.,

MUNIZ-SOLIS, R., GILI, C., MARTINELL, J.,

DOMENECH, R., PALMOVIST, P. Y GUERRA-

MERCHÁN, A., 1996. Estudio preliminar de la

malacofauna del Plioceno de Estepona (Má-

laga, España). Iberus, 13 (2): 93-117.

VILLALTA, J.F. Y ROSELL, J., 1966. Aportaciones

al estudio del Mioceno marino de la comarca

del Valles. Acta Geológica Hispánica, 1: 5-8.

VILLALTA, J.F., ROSELL, J. Y OBRADOR, A,, 1968.

Una nueva aportación al conocimiento del

Mioceno marino del Vallés. Acta Geológica

Hispánica, 3 (1): 19-21.

WARÉN, A., 1980. Marine Mollusca described by

John Gwyn Jeffreys, with the location of the

type material. Conchological Society of Great Bri-

tain anbd Ireland, Special Publication 1: 1-60, 8

láms.

WIENRICH, G., 2001. Die Fauna des marinen

Miozáns von Kevelaer (Niederrhein). Band 3.

Gastropoda bis Cancellariidae. Backhuys Pu-

plishers BV, Leiden,: 387-639.

WOOD, S.V., 1848. A monograph of the Crag Mo-

llusca. Part 1. Univalves. Paleontographical

Society, London: 208 pp., 21 láms.

Z1CH, A., 1934. Zur fauna des Mittel-Miócans

von Kostej (Banat), Typus-Bestimmung und

Tafeln zu O. Boettger's Bearbeitungen. Senc-

kenbergiana, 16 (4-6): 193-302.

a e

¡BN De E 7

br 1 sh coins

dd px ammm

bis no del Pa Ea

1144 MS 43 A

ria

O Sociedad Española de Malacología Iberus, 21 (1): 177-189, 2003

Adiciones a la fauna malacologica del litoral del Garraf (NE

de la Península Ibérica)

Additions to the malacological fauna of El Garraf (NE of the Iberian

Peninsula)

Anselmo PEÑAS* y Gonzalo GIRIBET**

Recibido el 15-VII-2002. Aceptado el 27-1-2003

RESUMEN

Se presenta una segunda lista de 54 especies de moluscos marinos (47 gasterópodos y 7

bivalvos), no citados previamente, recolectados en aguas del Garraf (Barcelona, NE de la

Península Ibérica). De estas especies, 18 se citan por primera vez en el Mediterráneo

español, siendo una de ellas, Pseudosetia ficaratiensis, la primera cita, probablemente

actual, en el Mediterráneo. Además, se cita por primera vez en el Mediterráneo el género

Bathycrinicola (Eulimidae), describiendo una nueva especie. Se incluyen, asimismo,

comentarios sobre algunos de los taxones mencionados y se ilustran al microscopio elec-

trónico de barrido (MEB) varios de ellos.

ABSTRACT

A complementary list of new citations of marine mollusks from waters of El Garraf (Barce-

lona, NE Iberian Peninsula) is presented. The new list includes 54 species of marine

mollusks (47 gastropods and 7 bivalves). From these, 18 are first reports for the Spanish

Mediterranean, including the first recent specimens of Pseudosetia ficaratiensis from the

Mediterranean. The genus Bathycrinicola (Eulimidae) is also reported for the Mediterra-

nean for the first time, including a new species. Some of the taxa are discussed and illus-

trated at the scanning electron microscopy (SEM).

PALABRAS CLAVE: Moluscos marinos, Garraf, NE Península Ibérica, Mar Mediterráneo, nueva especie.

KEY WORDS: Marine mollusks, Garraf, NE Iberian Peninsula, Mediterranean Sea, new species.

INTRODUCCIÓN

En el trabajo de GIRIBET Y PEÑAS

(1997), en el que se citaron 622 especies

de moluscos marinos (7 poliplacóforos,

417 gasterópodos, 190 bivalvos y 8 esca-

fópodos), quedó patente la gran riqueza

malacológica de los fondos marinos

situados frente a la pequeña comarca del

Garraf, En dicho trabajo, al cual nos remi-

timos, se delimitó la zona de estudio,

incluyendo un mapa, y se describió la

variedad y particularidad de sus fondos.

Se constató también que, probable-

mente, el Garraf sea la zona del Medite-

rráneo español más exhaustivamente

estudiada hasta la fecha, pues se citaron

53 especies por primera vez en él,

siendo dos de ellas primera cita para

todo el Mediterráneo.

* Carrer Olérdola, 39-5%. 08800 Vilanova i la Geltrú, Barcelona.

** Museum of Comparative Zoology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA.

Iberus, 21 (1), 2003

Posteriormente, GIRIBET Y PEÑAS

(1999) describieron en la zona una

nueva especie para la ciencia: Epilepton

parrusetensis. PEÑAS Y ROLÁN (2000)

citaron también en esta zona Turbonilla

postacuticostata Sacco, 1892.

El presente trabajo es una continua-

ción de los anteriores, y el resultado del

estudio de nuevo material. Se han

encontrado 255 especies, 201 de las

cuales ya fueron citadas anteriormente,

en fondos de similares características.

Por tanto, en este trabajo se citan sola-

mente las 54 especies que son nuevas en

la zona (47 gasterópodos y 7 bivalvos).

En total, se conocen hasta la fecha en el

litoral de la comarca del Garraf 678

especies de moluscos marinos.

La base de este trabajo ha sido el

estudio de nuevos sedimentos obteni-

dos en la zona, por tanto referido a

conchas, si bien en varias de ellas se

encontraron restos de partes blandas.

MATERIAL Y MÉTODOS

La mayor parte del material objeto

del presente trabajo fue recolectado

entre octubre y diciembre de 1996 por la

embarcación de pesca de arrastre “Joven

Mateo”, del puerto de Vilanova i la

Geltrú (Barcelona).

El 22 de octubre de 1996 se obtuvie-

ron unos 10 litros de detrito en el cala-

dero denominado “Mar de Nacra”, a

7,25 millas de la costa, frente a Sitges

(entre las pedanías de Vallcarca y

Garraf), a una profundidad de 105

metros. El lugar es un fondo arenoso-

fangoso, con una densa población de

Leptometra phalangium (Muller, 1841) y

donde hace años era muy abundante

Atrina pectinata (Linnaeus, 1767). Los

equinodermos fueron pescados vivos,

aunque se fragmentaron por efecto del

arrastre, y representaron casi la mitad

del volumen obtenido. Una vez lavado

el sedimento y pasado por una serie de

tamices, siendo el más fino de 0,3 mm

de luz de malla, se obtuvieron 0,8 litros

de arenas silíceas y materia orgánica (sin

contar los restos de crinoideos), rica en

micromoluscos, especialmente los gaste-

178

rópodos, muchos de ellos con restos de

partes blandas.

El 12 de diciembre de 1996 se obtu-

vieron unos 12 litros de sedimento en el

lugar denominado “El Vinyet”, al sur de

Sitges, a una profundidad de 160

metros, en fondo fangoso. Una vez

lavado y tamizado, se obtuvieron unos

2,5 litros de material formado básica-

mente por restos de bivalvos. El sedi-

mento era pobre en especies vivas,

excepto de las familias Nuculidae y

Thyasiridae.

Entre los meses citados, se analizó el

contenido estomacal de unos 200 ejem-

plares de Astropecten irregularis (Linck),

dragados entre 150 y 250 metros de pro-

fundidad, frente a Vilanova i la Geltrú.

Gran cantidad de ejemplares perte-

necientes a la familia Janthinidae fueron

recogidos, la mayoría con restos de

partes blandas, durante los veranos de

1998 y 1999 en las playas de Cubelles

por los malacólogos Ramón Beneito y

Vicente Buñuel.

Se han fotografiado al microscopio

electrónico de barrido aquellas especies

nuevas para la zona sobre las que se

aporta información adicional.

El listado de especies ha sido confec-

cionado básicamente de acuerdo con la

nomenclatura de la CLEMAM (Check

List of European Marine Mollusca:

http: //www.mnhn.fr/cgi-

bin /mamlist).

Abreviaturas utilizadas:

MNCN: Museo Nacional de Ciencias

Naturales, Madrid.

c: significa concha

v: valva

H: altura total de la concha

h: altura de la última vuelta

D: diámetro

=: aproximadamente igual

RESULTADOS

En el nuevo material estudiado se

han identificado 255 especies, la

mayoría ya citadas por GIRIBET Y PEÑAS

(1997). Por tanto, en el presente trabajo

PEÑAS Y GIRIBET: Adiciones a la fauna malacologica del litoral del Garraf

sólo se relacionan las especies no citadas

previamente, que en total son 54 (47 gas-

terópodos y 7 bivalvos), 4 de ellas iden-

tificadas solamente a nivel genérico.

En la Tabla I se ofrece una relación

de estas 54 especies, indicando cuáles de

ellas se citan por primera vez para el

Mediterráneo español o para el Medite-

rráneo en general. La especie irá en

negrita cuando sea objeto de comenta-

rios. Se señala, asimismo, el lugar de

recolección, su abundancia relativa y las

especies ilustradas.

COMENTARIOS SOBRE ALGUNOS

TAXONES

De la mayoría de especies citadas en

este trabajo existen fotografías y descrip-

ciones actualizadas. Por tanto, en este

apartado nos hemos limitado a comentar

algunos de los taxones que nos han pare-

cido de mayor interés, ya sea por su

rareza O por que se aporta nueva infor-

mación. Además incluimos la descrip-

ción de una nueva especie para la ciencia

perteneciente a la familia Eulimidae.

Clase GASTROPODA

Tricolia deschampsi Gotas, 1993

Tricolia deschampsi Gofas, 1993. J. Moll. Stud., 59: 354-356, figs. 6-8, 19, 29, 30.

Material examinado: 3 c, procedentes del “Mar de Nacra”, Sitges, dragadas a 105 m de profundi-

dad.

Comentarios: Esta especie, cuya deter-

minación no nos ofrece dudas, es abun-

dante en el infralitoral y circalitoral del estre-

cho de Gibraltar y Mar de Alborán. La cita

en el Garraf representa la localización más

al norte del Mediterráneo español.

Tricolia tingitana Gofas, 1982

Tricolia tingitana Gofas, 1982. J. Moll. Stud., 48: 202-205, figs. 54-64.

Material examinado: 4 c, procedentes del “Mar de Nacra”, Sitges, dragadas a 105 m de profundi-

dad; 1 c, procedente de “El Vinyet”, al sur de Sitges, dragada a 160 m de profundidad.

Comentarios: Esta especie fue descrita

para la zona del estrecho de Gibraltar y

para Calvi, Cerdeña. Posteriormente,

GOFAS (1993) amplió su área de distribu-

ción al sur de Portugal y banco atlántico

de Gorringe. Además, revisó el material

de Córcega y llegó a la conclusión que

pertenecía a una especie diferente, que

describió como T. punctura. De esta

forma, la cita en el Garraf representa la

localización más oriental y septentrional

del Mediterráneo español.

Bathycrinicola nacraensis spec. nov. (Figs.4-7)

Material tipo: Holotipo (Fig. 4) y un paratipo, ambos con dimensiones de 1,2 x 0,5 mm y 4 vueltas

de espira, depositados en el MNCN (n? 15.05/46482) , ambos procedentes del “Mar de Nacra”,

Sitges, dragado a 105 metros de profundidad.

Localidad tipo: “Mar de Nacra”, Sitges, provincia de Barcelona (España), a 7,25 millas de la costa,

a 105 m de profundidad.

Etimología: El nombre específico proviene del caladero en el que ha sido encontrada por primera

vez, el “Mar de Nacra”. Nacra proviene del Catalán, y es el nombre común con el que se conocen

los bivalvos de la familia Pinnidae.

179

Iberus, 21 (1), 2003

Tabla I. Relación de las especies encontradas en el área de estudio, lugar donde se han encontrado,

figuras en las que se representan y abundancia relativa. Las especies en negrita se comentan en el

texto. Códigos. *: primera cita en el Mediterráneo español; **: primera cita en el Mediterráneo o

nueva cita para la ciencia; nm: Mar de Nacra; v: El Vinyet; e: estrellas; c: Cubelles; +: 1-2 ejemplares;

++: 3-10 ejemplares; +++: más de 10 ejemplares.

Table I. Checklist of species found in the area of study, location within the area, figures where the species

are represented, and abundante. Species in boldface are discussed in the text. Codes. *: first record for the

Spanish Mediterranean; **: firs record for the Mediterranean or new species; n: Mar de Nacra; v: El

Vinyet; e: in starfish gut contents; c: Cubelles; +: 1-2 specimens; ++: 3-10 specimens; +++: more than

10 specimens.

Close GASTROPODA

Familia SCISSURELLIDAE

Scissurella costata D'Orbigny, 1842: n +

Familia SKENEIDAE

Skenea serpuloides (Montagu, 1808): n +

* Lissomphalia bithynoides (Monterosato, 1880): n, v Figs. 1-2 ++

Dikoleps marianae Rubio, Dantart y Luque, 1998: n ++

Anekes turrita (Gaglini, 1987): n ++

* Polazzia ausoniae (Palazzi, 1988): n Fig. 3 +

Familia TRICOLIIDAE

Tricolia deschampsi Gofas, 1993: n +

Tricolia tingitana Gofos, 1982: n, v +

Familia TRIPHORIDAE

Monophorus thiriotae Bouchet, 1984: n | +

Familia CERITHIOPSIDAE

* Cerithiopsis fayalensis Watson, 1886: n, v +++

Familia JANTHINIDAE

Janthina nitens Menke, 1828: c ++

Jonthina pallida W. Thompson, 1840: c ++

Familia ACLIDIDAE

Aclis minor (Brown, 1827): n +

Fomilia EPITONIIDAE

Epitonium tiberii (de Boury, 1890): n, v +

Acirsa subdecussata (Cantraine, 1835): n +

Familia EULIMIDAE

** Bathycrinicola nacraensis spec. nov. n Figs. 4-7 +

* Crinophtheiros comatulicola (Graft, 1875): n +

Curveulima devians (Monterosato, 1884): n, v ++

Vitreolina antiflexa Monterosato, 1884: n +

Vitreolina curva (Monterosato, 1874 ex Jeffreys ms.): n +

Vitreolina incurva (Bucquoy, Dautzenberg y Dollfus, 1883): n, v He

Familia RISSOIDAE

Alvania hispidula (Monterosato, 1884): n +

Alvania lanciae (Calcara, 1841): n +

* Alvania sp. n Figs. 8-10 +

Pusillina parva (Da Costa, 1778): n Figs. 11-12 +

* Pusillina sp.: e Figs. 13-14 +

** Pseudosetia ficaratiensis (Brugnone, 1876): n, e Figs. 15-16 +H

180

PEÑAS Y GIRIBET: Adiciones a la fauna malacologica del litoral del Garraf

Tabla I. Continuación.

Table [. Continuation.

Familia CAECIDAE

Caecum subannulatum De Folin, 1870: n

Familia ATLANTIDAE

Atlanta fusca Souleyet, 1852: n

Atlanta inflata Souleyet, 1852: n. v

Familia FIROLIDAE

* Firoloida desmarestia Lesueur, 1817: n

Familia CONIDAE

Raphitoma erronea (Monterosato, 1884): n

Raphitoma histrix (Bellardi, 1847): n

Raphitoma pseudohystrix (Sykes, 1906): v

Familia CIMIDAE

* Cima sp.: n

Familia PYRAMIDELLIDAE

* Tiberia octaviana Di Geronimo, 1973: n

Chrysallida multicostata (Jeffreys, 1884): n, v

Miralda elegans (De Folin, 1870): n

Odostomella bicincta (Tiberi, 1868): n

Eulimella neoattenuata Gaglini, 1992: v

Ondina crystallina Locard, 1892: n, v

Ondina warreni (W. Thompson, 1845): n

Familia ACTEONIDAE

Acteon monterosatoi Dautzenberg, 1889: n

Familia DIAPHANIDAE

Colpodaspis pusilla M. Sars, 1870: n

Familia RETUSIDAE

Retusa mamillata (Philippi, 1836): n

Fomilia PHILINIDAE

Philine punctata (J. Adams, 1800): n

* Philine striatula Monteroscto, 1874: n

Close BIVALVIA

Familia YOLDIIDAE

* Microgloma pusilla (Jeffreys, 1879): n,

Fomilia LIMIDAE

Limatula subovata (Jeffreys, 1876): n

* Limatula sp.: n

Familia THYASIRIDAE

* Thyasira alleni Carrozza, 1981: n, v

Familia MONTACUTIDAE

* Montacuta semirubra Goglini, 1992: n

* Kelliopsis jozinae van Aartsen y Corrozza, 1997: n, v

Familia MESODESMATIDAE

* Monterosotus primus (Locard, 1899): n

Figs. 17-20

Fig. 21

181

Iberus, 21 (1), 2003

Descripción: Concha diminuta (Fig. 4),

cónico-piramidal, transparente, muy bri-

llante. Protoconcha (Figs. 5-7) lisa,

globosa, de casi 1,5 vueltas, con la sutura

de separación con la teloconcha casi ver-

tical, con un diámetro de unas 200 pum.

Teleoconcha de 2,5 vueltas convexas, la

última ancha, redondeada, h > 60% H.

Sutura estrecha pero profunda. Superficie

lisa, sin trazos de escultura, con excep-

ción de las líneas de crecimiento, muy

tenues, que son ortoclinas, salvo justo

bajo la sutura donde son prosoclinas.

Abertura circular. Columela arqueada,

delgada, cortante, tras la cual se encuen-

tra un ombligo pequeño pero profundo.

Discusión: Creemos que el hospeda-

dor de esta especie debe ser Leptometra

phalangium (Muller, 1841), dado el

hábitat en el que ha sido encontrada,

aunque no ha sido observada sobre este

crinoideo. Ambas conchas se encontra-

ron con restos de partes blandas.

BOUCHET Y WARÉN (1986) describen el

nuevo género Bathycrinicola, para eulími-

dos de profundidad parásitos de crinoi-

deos e incluyen en él cinco especies atlán-

ticas. Sólo una de ellas, B. curta (Warén,

1972), tiene una protoconcha parecida,

pero el resto de características morfológi-

cas es diferente: concha postlarval mayor,

más estrecha y alargada, con las vueltas

menos convexas, la sutura es muy

somera y carece de ombligo.

Hemos ubicado esta especie en el

género Bathycrinicola como el más ade-

cuado, al considerarla parásita de un

crinoideo y por el tipo de protoconcha.

Las conchas pertenecientes al género

Umbilibalcis también son umbilicadas,

pero tienen un perfil muy cónico, con

numerosas vueltas, y tienen una micro-

escultura conspicua formada por surcos

axiales.

Es la primera vez que este género se

cita para el Mediterráneo.

Alvania sp. (Figs. 8-10)

Material examinado: 1 c, en el “Mar de Nacra”, Sitges, dragada a 105 m. Dimensiones: 2,4 x 1,3 mm.

Comentarios: Se trata de una concha

(Fig. 8) probablemente rodada, con la

protoconcha (Figs. 9, 10) gastada, en la

que no se aprecia bien la escultura. A pe-

sar de ello, no se asemeja a ninguna de

las especies descritas en el Mediterráneo

ni Atlántico próximo. Solamente A. cimi-

coides (Forbes, 1844) tiene un perfil pare-

cido, pero presenta la escultura conspi-

cua, una sutura canaliculada y, sobre to-

do, su protoconcha es multiespiral, mien-

tras que en Alvanía sp. es pauciespiral.

Pusillina sp. (Figs. 13, 14)

Material examinado: 2 c, en Astropecten irregularis, a 150-250 m de profundidad. Dimensiones

máximas 2,7 x 1,3 mm.

Comentarios: Esta especie se caracte-

riza por carecer de la escultura, presente

en la mayoría de las pertenecientes a este

género, si bien su color blanquecino con

débiles flámulas verticales amarillentas

es típico de él. Los ejemplares hallados

recuerdan alguna forma de P. interrupta

(J. Adams, 1800), que podría haber sido

transportada a esas cotas batimétricas

pues, excepto P. inconspicua (Alder, 1844),

ninguna otra especie de este género vive

en las aguas profundas del Mediterráneo.

Pseudosetia ficaratiensis (Brugnone, 1876) (Figs. 15, 16)

Rissoa ficaratiensis Brugnone, 1876. Miscellanea Malacologica, pars 2: 21.

182

PEÑAS Y GIRIBET: Adiciones a la fauna malacologica del litoral del Garraf

100 um

Figuras 1-2. Lissomphalia bithynoides. 1: concha, 0,8 mm; 2: protoconcha. Figura 3. Palazzia auso-

niae. Concha, 0,57 mm. Figuras 4-7. Bathycrinicola nacraensis. 4: holotipo, 1,2 mm; 5, 6: proto-

concha; 7: detalle de la sutura.

Figures 1-2. Lissomphalia bithynoides. 1: shell, 0.8 mm; 2: protoconch. Figure 3. Palazzia ausoniae.

Shell, 0.57 mm. Figures 4-7. Bathycrinicola nacraensis. 4: holotype, 1.2 mm; 5, 6: protoconch; 7:

detail of the suture.

Iberus, 21 (1), 2003

Material examinado: 4 c, 2 en Astropecten irregularis (Linck), frente a Vilanova i la Geltrú, entre 150

y 250 my 2 en el “Mar de Nacra”, Sitges, a 105 m de profundidad.

Descripción: En BOUCHET Y WARÉN

(1993: 688-690, figs. 1595-1596, 1613,

1614). Se muestra aquí una concha

(Fig. 15), con unas dimensiones de 1,5

x 0,9 mm y su protoconcha (Fig. 16),

con un diámetro de unas 335 um.

Distribución: Especie solamente

conocida en aguas profundas entre el

Estrecho de Gibraltar y el suroeste de

Portugal, y como fósil Plio-Pleisto-

ceno en Sicilia. Esta es la primera vez

que se cita en el Mediterráneo,

creemos que como fauna actual, ya

que dos de las conchas estudiadas, si

bien carecían de partes blandas, eran

frescas.

Cima sp. (Figs. 17-20)

Material examinado: 1 c, procedente del “Mar de Nacra”, Sitges, dragada a 105 m de profundidad.

Descripción: Concha (Figs. 17, 18)

muy pequeña, conoidea alargada,

blanca vítrea. Dimensiones 1,3 x 0,5 mm.

Protoconcha (Figs. 19, 20) grande y

globosa, más bien heterostrofa, con un

diámetro de 165 um. Teleoconcha con

las vueltas convexas de rápido creci-

miento, la última grande, oval-redonde-

ada, h= 50% H. Sutura profunda. Aber-

tura oval. Columela arqueada. No umbi-

licada.

Comentarios: Esta rara especie parece

tener características cercanas a Aclis,

Eulimella y Cima. La protoconcha

recuerda algunas formas de Eulimella,

especialmente E. similebala Peñas y

Rolán, 1999, de los bancos atlánticos

submarinos al sur de las Azores, pero en

ella se observa claramente la sutura del

núcleo. A pesar de que parece más

cercana al género Aclis, hemos optado

por ubicar provisionalmente esta

especie en la familia Cimidae, por las

características de la protoconcha hete-

rostrofa, aunque distinta a las conocidas

de la familia Pyramidellidae.

Tiberia octaviana Di Geronimo, 1973 (Fig. 21)

Tiberia octaviana Di Geronimo, 1973. Conchiglie, 9(11-12): 217-222.

Material examinado: 1 c, procedente del “Mar de Nacra”, Sitges, dragada a 105 m. de profundi-

dad.

Descripción: En DI GERONIMO (1973).

Se muestra aquí la concha encontrada

(Fig. 21).

Distribución: Sólo conocida en aguas

de Catania, Italia. Esta es la primera cita

en el Mediterráneo español.

Se muestra aquí una concha con

unas dimensiones de 5,3 x 2,1 mm.

Comentarios: Esta especie, descrita

con dos únicos ejemplares, ha sido consi-

derada dudosa o una forma más esbelta

de T. minuscula Monterosato, 1880. El

único ejemplar recolectado ha sido com-

184

parado con varias conchas de T. minus-

cula de la misma comarca del Garraf, con

las cuales presenta diferencias. La

concha se ajusta a la descripción original

no sólo en su forma y relación H/D, sino

en las bandas de color marrón que son

más anchas que en T. minuscula, además

de tener una tercera banda periumbilical

más difuminada. Seguramente el estudio

de un mayor número de ejemplares

podría confirmar que T. octaviana y T.

minúscula son dos especies diferentes,

como creemos, sin formas intermedias.

PEÑAS Y GIRIBET: Adiciones a la fauna malacologica del litoral del Garraf

Figuras 8-10. Alvanía sp. 8: concha, 2,4 mm; 9-10: protoconcha. Figuras 11, 12. Pusillina parva.

11: concha, 3,4 mm, Vigo; 12: concha, 1,8 mm, Garraf. Figuras 13, 14. Pusillina sp. 13: concha,

2,7 mm; 14: protoconcha.

Figures 8-10. Alvania sp. 8: shell, 2.4 mm; 9-10: protoconch. Figures 11, 12. Pusillina parva. 11:

shell, 3.4 mm, Vigo; 12: shell, 1.8 mm, Garraf. Figures 13, 14. Pusillina sp. 13: shell, 2.7 mm; 14:

protoconch.

185

Iberus, 21 (1), 2003

Miralda elegans De Folin, 1870

Mathilda elegans De Folin, 1870. Les Fonds de la Mer, 1: 212, 213, pl. 28, fig. 15.

Pyrgulina sculptatissima Dautzenberg, 1913. Ann. Inst. Océanogr., 1:67, 68, lám. 3, figs. 15-16.

Chrysallida pulchra Gaglini, 1992. Argonauta, 7(1-7): 138, 139, fig. 146.

Material examinado: 1 c, procedente del “Mar de Nacra”, Sitges, dragada a 105 m.

Comentarios: Esta especie, típica del

África Occidental, fue citada en el Medi-

terráneo por HOENSELAAR Y MOOLEN-

BEEK (1990: 65-66, figs. 1-4) en Formen-

tera, Baleares, y por GAGLINI (1992: 138,

139, fig. 146) como Chrysallida pulchra en

Sicilia. Esta es la primera cita para el

Mediterráneo español peninsular.

Philine striatula Monterosato, 1874 ex Jeffreys MS

Philine striatula Monterosato, 1874. Journal de Conchyliologie, 22: 281.

Material examinado: 1 c, procedente del “Mar de Nacra”, Sitges, dragada a 105 m.

Descripción: En OLIVERIO Y T'RINGALI

(2001: 138, 139), quienes ilustran un

sintipo (figs. 73-75).

Comentarios: OLIVERIO Y 'TRINGALI

(2001) en su revisión de los tipos de los

Opistobranquios descritos por Montero-

sato dan validez a esta especie encon-

trada en aguas profundas de Sicilia.

Aquí se cita por primera vez en el Medi-

terráneo español.

Clase BIVALVIA

Microgloma pusilla (Jeffreys, 1879)

Leda pusilla Jeffreys, 1879. Proc. Zool. Soc. London, 581, pl. 46, fig. 6.

Material examinado: 2 v, procedentes del “Mar de Nacra”, Sitges, dragadas a 105 m.

Comentarios: Ver ilustración en

SALAS (1996: figs. 58-61), quien incluye

esta especie en la familia Pristiglo-

midae, y la cita para aguas profundas

frente al Cabo San Vicente, Portugal, y

frente a Rabat, Marruecos. Aquí se cita

por primera vez en el Mediterráneo

español.

Limatula sp.

Material examinado: 3 v, procedentes del “Mar de Nacra”, Sitges, dragadas a 105 m.

Comentarios: Las 3 valvas encon-

tradas, ninguna completa, se corres-

ponden con el ejemplar citado en

GIRIBET Y PEÑAS (1997) como Limatula

cf. gwyni (Sykes, 1903), sin escultura

radial aparente, pero con las líneas

concéntricas de crecimiento muy mar-

cadas.

Montacuta semirubra Gaglini, 1992

Montacuta semirubra Gaglini, 1992. Argonauta, 7(1-6): 178, 179, figs. 165-166.

Montacuta semirubra Monterosato, 1872 (nomen nudum)

186

PEÑAS Y GIRIBET: Adiciones a la. fauna malacologica del litoral del Garraf

100 um

Figuras 15, 16. Pseudosetia ficaratiensis. 15: concha, 1,5 mm; 16: protoconcha. Figuras 17-20. Cima

sp. 17, 18: concha, 1,3 mm; 19-20: protoconcha. Figura 21. Tiberia octaviana. Concha, 5,3 mm.

Figures 15, 16. Pseudosetia ficaratiensis. 15: shell, 1.5 mm; 16: protoconch. Figures 17-20. Cima sp.

17, 18: shell, 1.3 mm; 19-20: protoconch. Figure 21. Tiberia octaviana. Shell, 5.3 mm.

187

Iberus, 21 (1), 2003

Material examinado: 2 v, procedentes del “Mar de Nacra”, Sitges, dragadas a 105 m.

Comentarios: En el trabajo de

GAGLINI (1992: 178, 179, figs, 165-166)

puede verse la descripción e ilustración

CONCLUSIONES

Como se señaló anteriormente, se han

identificado con un asterisco las especies

que se citan por primera vez para el

Mediterráneo español, que son 18 (12

Gasterópodos y 6 Bivalvos), 4anivel gené-

rico. Deellas, Pseudosetia ficaratiensis se cita

por primera vez reciente en el Mediterráneo

en general y Bathycrinicola nacraensis sedes-

cribe como nueva especie para la ciencia.

Este nuevo estudio reafirma lo cons-

tatado en el primer trabajo de los autores:

la presencia en el Garraf de la mayoría de

la malacofauna típica del Mediterráeo

español (exceptuando la endémica del

Mar de Alborán y estrecho de Gibraltar).

También podemos confirmar que la

costa situada frente a El Garraf, a pesar

de su escasa extensión, es una zona muy

rica en moluscos marinos, debido a la

diversidad de sus fondos. Hasta el pre-

sente se han citado en la comarca 678 es-

pecies, 69 por primera vez para el Medi-

terráneo español, de las cuales 3 se han

citado por primera vez en el Mediterrá-

neo en general y se han descrito dos

nuevas especies para la ciencia.

BIBLIOGRAFÍA

BOUCHET, P. Y WARÉN, A., 1986. Revision of

the Northeast Atlantic bathyal and abyssal

Aclididae, Eulimidae, Epitoniidae (Mollusca,

Gastropoda). Bollettino Malacologico, Suppl. 2:

299-576.

BOUCHET, P. Y WARÉN, A., 1993. Revision of

the Northeast Atlantic bathyal and abyssal

Mesogastropoda. Bollettino Malacologico,

Suppl. 3: 580-839.

GAGLINI, A., 1992 (1991). Terze spigolature ...

monterosatiane. Argonauta, 7 (1-6): 125-180.

GERONIMO, 1. DI, 1973. Tiberia octaviana n. sp. di

Pyramidellidae (Gastropoda, Opistobran-

chia) del Mediterraneo. Conchiglie, 9 (11-12):

217-222.

188

de esta especie, cuya localidad tipo es

Palermo, Sicilia. Se cita por primera vez

para el Mediterráneo español.

AGRADECIMIENTOS

Nuestro agradecimiento a Manuel

Roca y sus hijos Jesús y Pavel, de Vila-

nova i la Geltrú, que nos facilitaron los

asteroideos y los sedimentos dragados

por su embarcación “Joven Mateo”; a

Ramón Beneito, de Tarragona y a

Vicente Buñuel, de Valls (Tarragona),

quienes recolectaron numerosos ejem-

plares de la familia Janthinidae; a

Carmen Salas, del Dpto. de Biología

Animal de la Universidad de Málaga,

por su ayuda en la determinación de

algunos ejemplares de bivalvos; a

Anders Warén (Swedish Museum of

Natural History, Stockholm) por sus

comentarios sobre el género Bathycrini-

cola; a Jesús Méndez, del CACTI (Centro

de Apoyo Científico y Tecnológico a la

Investigación), de la Universidad de

Vigo por la realización de las fotografías

al MEB; y especialmente a Emilio Rolán,

por sus comentarios, permanente cola-

boración y ayuda en la digitalización de

las láminas.

GIRIBET, G. Y PEÑAS, A., 1997. Fauna malacoló-

gica del litoral del Garraf (NE de la Península

Ibérica). Iberus, 15 (1): 41-93.

GIRIBET, G. Y PEÑAS, A., 1998. A new Epilepton

species (Bivalvia, Montacutidae) from the

Western Mediterranean. lberus, 16(2): 117-

127.

GOFAS, S., 1982. The genus Tricolia in the Eas-

tern Atlantic and the Mediterranean. The

Journal of Molluscan Studies, 48: 182-213.

GOEAS, S., 1993. Notes on some Ibero-Maroccan

and Mediterranean Tricolia (Gastropoda, Tri-

coliidae), with descriptions of new species.

The Journal of Molluscan studies, 59: 351-361.

PEÑAS Y GIRIBET: Adiciones a la fauna malacologica del litoral del Garraf

HOENSELAAR, H. J. Y MOOLENBEEK, R. G., 1990.

First record of Miralda elegans (De Folin, 1870)

nov. comb. from the Mediterranean Sea (Gas-

tropoda, Pyramidellidae). Bollettino Malaco-

logico, 26 (1-4): 65-66.

OLIVERIO, M. Y TRINGALI, L. P., 2001. The types

of marine molluscan species described by

Monterosato, in the Museo Civico di Zoolo-

gia, Roma. General scope of the work, and

part 1: the opisthobranch gastropods. Bollet-

tino Malacologico, 37 (5-8): 121-142.

PEÑAS, A. Y ROLÁN, E., 1999. Pyramidellidae

(Gastropoda, Heterostropha) de la Misión

Oceanográfica “Seamount 2”. Iberus, Suple-

mento 5: 151-199.

PEÑAS, A. Y ROLÁN, E., 2000. The family Pyra-

midellidae Gray, 1840 (Mollusca, Gastro-

poda, Heterostropha) in West Africa. 7. Ad-

denda to the genera Eulimella and Turbonilla,

with a list of the east Atlantic species and sy-

nonyms. Argonauta, 13 (2): 59-80.

SALAS, C., 1996. Marine Bivalves from off the

Southern Iberian Peninsula collected by the

Balgim and Fauna 1 expeditions. Haliotis, 25:

33-1.

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Iberus, 21 (1);191-206, 2003

Nuevas especies de la familia Hydrobiidae (Mollusca,

Orthogastropoda) de la Comunidad Valenciana (España)

New species of the family Hydrobiidae (Mollusca, Orthogastropoda)

from “Comunidad Valenciana” (Spain)

Emilio ROLÁN* y Alberto MARTÍNEZ-ORTÍ**

Recibido el 27-X1-2002. Aceptado el 10-111-2003

RESUMEN

Se describen cinco nuevas especies de Hydrobiidae encontradas en dos surgencias del

municipio de Sueras y en otra más en el de Cabanes, ambos de la provincia de Caste-

llón, Comunidad Valenciana; estas especies se asignan tentativamente a los géneros By-

thiospeum (1) y Sardopaladilhia (4), y se discuten sus asignaciones genéricas.

ABSTRACT

Five new species of the Hydrobiidae found in two springs in the locality of Sueras and one

more in Cabanas (both from Castellón province, Comunidad Valenciana, Spain) are des-

cribed and their relationships are discussed. They are tentatively assigned to the genera

Bythiospeum (1) and Sardopaladilhia (4).

PALABRAS CLAVE: Hydrobiidae, taxonomía, nuevas especies, Bythiospeum, Sardopaladilhia, Comunidad

Valenciana, España.

KEYWORDS: Hydrobiidae, taxonomy, new species, Bythiospeum, Sardopaladilhia, Comunidad Valenciana,

Spain.

INTRODUCCIÓN

Los moluscos de aguas freáticas son

poco conocidos debido a sus pequeñas

áreas de dispersión, y a las dificultades

de obtener muestras de animales vivos

y poder estudiar así la anatomía de las

partes blandas. Antiguos trabajos como

el de BOURGUIGNAT (1863) inician nues-

tros conocimientos sobre estos grupos.

Mucho más escasa es la información

existente sobre estos moluscos en la

Península Ibérica, que aparece en traba-

jos generales como los de Haas (1929) y

BECH (1990), en los que se recogen citas

* Cánovas del Castillo, 22, 36202 Vigo, España.

de cuatro especies del género Moitessie-

ria Bourguignat, 1863 que, según

BOETERS (1988), son dos especies sola-

mente. A éstas, habría que añadir más

recientemente, la primera especie de

Paladilhiopsis para la Península Ibérica

descrita por ROLÁN Y RAMOS (1996). En

zonas próximas de Francia BOETERS Y

GITTENBERGER .(1980) describe una

nueva Moitessieria.

Las relaciones sistemáticas entre

estos géneros, todavía insuficientemente

estudiadas, han sido comentadas por

** Museu Valencia d' Historia Natural, Passeig de la Petxina, 15, 46008 Valencia, España.

191

Iberus, 21 (1), 2003

B El Quito

En m

Corra de Amón

Mi ara

HA]

EME Emil

le Miravet

(Cabanes)

Figura 1. A. Localización geográfica de las surgencias estudiadas en la Provincia de Castellón

(España). B. Mapa detallado del área de Sueras (Castellón).

Figure 1. A. Geograpbical location of the springs studied in the Castellón province (Spain). B. Detailed

map of the Sueras area.

BERNASCONI (1984) y BODON Y GIUSTI

(1991). Las sinonimias y trabajos que

han estudiado la taxonomía y diversas

aportaciones en la familia Hidrobiidae

han sido recogidas en KABAT y HERSH-

LER (1993).

Los descubrimientos de nuevas

especies e incluso nuevos géneros para

moluscos de aguas subterráneas (MAN-

GANELLI, BODON, CIANFANELLI, TALENTI

Y GiusTI 1998; RAMOS, ARCONADA,

ROLÁN Y MORENO, 2000; ARCONADA Y

RAMOS, 2001; BERTRAND, 2001; GIRARDI,

2001; ARCONADA Y RAMOS, 2002) se

siguen produciendo todavía hoy en día,

192

pese a la dificultad para conseguir la

recolección de estos moluscos.

En la Comunidad Valenciana los

estudios sobre las fuentes realizados por

MORELL (1992) en la provincia de Caste-

llón y TAPIA (1996), en las de Castellón y

Valencia, han sido el punto de partida

para el estudio de estos diminutos

moluscos. Posteriormente los trabajos

sobre anatomía y de descripción de

nuevas especies de RAMOS ET AL. (2000),

ARCONADA Y RAMOS (2001) y de ARCO-

NADA Y RAMOS (2002) han contribuido

notablemente al conocimiento de la

familia Hydrobiidae en esta región.

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

MATERIAL Y MÉTODOS

Dos surgencias muestreadas se

encuentran situadas en el parque

natural de la Sierra de Espadán, en la

provincia de Castellón, concretamente

en el término de Sueras: Font de Castro

(UTM: 305YK249245, 420 m) y Piscina

de Sueras (UTM: 305YK277262, 300 m).

Este último nombre hace referencia a

que se encuentra a unos 100 m, aguas

arriba, de la piscina municipal, a la que

suministra agua en verano. La geología

de estas localidades corresponde a dolo-

mías, margas, calizas y arcillas con

yesos en la primera y sin ellos en la

segunda, presentando las dos facies

bicarbonatada cálcica (TAPIA, 1996).

Otra surgencia está en la localidad de

Cabanes: Ullal de Miravet (Castellón)

(UTM: 31TBE495446, 144 m) y su geolo-

gía corresponde a calizas con recubri-

mientos de arcillas. Para la situación de

estas surgencias ver la Figura 1.

El material fue inicialmente recolec-

tado por Gloria Tapia en 1994, y cons-

taba de varias conchas de distintas espe-

cies. Con posterioridad, este material se

incrementó con nuevas prospecciones

realizadas por el segundo autor en 2002.

Para la recolección del material se ha

seguido la metodología aplicada por

VIAL (1999). Hay que resaltar la necesi-

dad de sumergirse para recoger sedi-

mentos en el interior de la surgencia de

RESULTADOS

la Piscina de Sueras. Los ejemplares

fueron separados del sedimento

mediante columna de tamices y poste-

rior selección con el esteromicroscopio

Leica Wild-M8.

Las vueltas de espira del material

estudiado son contadas a partir de un

núcleo inicial, según la definición reali-

zada por VERDUIN (1977).

El estudio al MEB fue realizado

montando el material en soportes de

aluminio con cinta adhesiva Scothch de

doble cara y cemento de carbón conduc-

tivo, haciendo la metalización con oro.

De cada especie fueron estudiadas entre

12 y 35 conchas.

Abreviaturas utilizadas:

AMNH American Museum of Natural

History, Nueva York

BMNH The Natural History Museum,

Londres

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Museum National d'Histoire

Naturelle, Paris

MVHN Museu Valencia d'Historia

Natural, Valencia

MZB Museu de Zoologia, Barcelona

NNMN Nationaal Natuurhistorich

Museum Naturalis, Leiden

CER colección de E. Rolán

c concha

f fragmento

“Bythiospeum” gloriae spec. nov. (Figs. 2-8)

Material tipo: Holotipo (Fig. 2) depositado en el MNCN, n* 15.05/46586. Paratipos en las siguien-

tes colecciones: MNCN (2 c), MNHN (1 c), MVHN (8 c, 3 f, n2 805), MZB (1 c, n£ 2002-691), NNMN

(1 c, n* 94818) y CER (8 c, 11 f).

Localidad típica: Font de Castro, Sueras (Castellón).

Etimología: El nombre específico se dedica a Gloria Tapia, quién estudió los macroinvertebrados

de las fuentes de la provincia de Castellón y encontró por primera vez las conchas de las especies

estudiadas en este trabajo.

Descripción: Concha (Figs. 2, 3) alar-

gada, ligeramente cónica, hialina, con una

altura máxima entre 2,0 y 2,4 mm, ápice

romo y un número de vueltas de espira

comprendido entre 4 y 5, siendo estas

vueltas convexas, aparentemente lisas, con

superficie Opaca y una sutura profunda.

Las líneas de crecimiento son ortoclinas,

poco visibles en algunas zonas, apare-

ciendo de vez en cuando alguna línea o

193

Iberus, 21 (1), 2003

varias muy marcadas, como si fuesen

pequeñas costillas. La abertura es redon-

deada, muy ligeramente oval, con el borde

sencillo y fino, pero algo incurvado hacia

afuera. Peristoma simple y libre, excepto

en su parte interna donde tiene una

pequeña zona de contacto con la pared de

la última vuelta. Visto lateralmente (Fig.

4) se aprecia que no todo el borde del

peristoma se encuentra en el mismo plano,

sino que se prolonga ligeramente en la

parte en contacto con la última vuelta,

mientras por arriba y afuera se ondula,

retrasándose ligeramente, con la forma de

una mínima escotadura. Hay un ombligo

pequeño que queda semioculto por la

eversión del peristoma.

La protoconcha (Fig. 5) no presenta

un límite claro y apreciable con la telo-

concha. El núcleo mide 110-130 um.

Microescultura: en la protoconcha (Fig.

5), prácticamente no se observa microes-

cultura en la primera media vuelta, pero

con posterioridad aparecen pequeñísimas

elevaciones en forma de líneas más o

menos evidentes que se colocan en posi-

ción espiral (Fig. 6), excepto en la parte

subsutural, donde se hacen un poco obli-

cuas. Después de las primeras vueltas (Fig.

7) las líneas se hacen más finas y apare-

cen en toda la superficie de la vuelta (Fig.

8) que está formada por pequeñísimas ele-

vaciones muy numerosas y discontinuas

que se sitúan en sentido espiral.

Dimensiones: El holotipo mide 2,1 x

0,9 mm. Alguna concha alcanza 2,3 mm.

El animal es desconocido.

Discusión: Asignación genérica: La

carencia de ejemplares con partes blandas

dificulta la asignación genérica. Por su

forma corta, ligeramente cónica y el escaso

número de vueltas de espira, parece que

no debería asignarse al género Moitessie-

ria Bourguignat, 1863, que además suele

presentar una escultura espiral bastante

prominente y frecuentemente formada por

cavidades (BODON Y GIUSTI, 1991).

Por su forma, podría pertenecer al

género Paladilhiopsis Pavlovic, 1913,

(especie tipo P. robiciana Clessin, 1882),

género que ha sufrido diferentes conside-

raciones: según SCHUTT (1970) sería un

subgénero de Paladilhia y para GIUSTI y

PEZZOLI (1982) sería un sinónimo de By-

194

thiospeum, aunque para BERNASCONI (1985)

sería un subgénero; sus conchas tienen

una espira de crecimiento uniforme y

lento; el peristoma está bastante expan-

dido pero el contacto con la vuelta ante-

rior es mayor; el ombligo está casi oculto

por el peristoma y, además, la escultura

puede ser muy ligera o inexistente (GIUSTI

y PEZZOLI, 1980). GrusTI Y PEZZOLI (1982)

Opinan que el género está mal definido

porque su especie típica ha sido extinguida

y que es preferible no sea usado. En cual-

quier caso, las especies de Paladilhiopsis

pueden o no tener microescultura, pero la

última vuelta es más adherente en la parte

superior de la abertura a la vuelta ante-

rior, por lo que decidimos descartar su

pertenencia a este género.

Algo parecido a ésto también ocurre

en Paladilha Bourguignat, 1865 (especie

tipo P. pleurotoma Bourguignat, 1865) ya

que las especies de este género, según

BOETERS Y GITTENBERGER (1990), suelen

tener una separación entre el borde de la

abertura y la vuelta anterior, y son casi

lisas O tienen líneas espirales muy

ténues. Y lo mismo se podría decir de

Palaospeum Boeters, 1999, que también

proviene del Sur de Francia y Pirineos.

También la nueva especie presenta

cierto parecido en su concha con las del

género Clameia Boeters y Gittenberger

(1990) (especie tipo C. brooki Boeters y Git-

tenberger, 1990) pero ésta también tiene la

abertura separada de la vuelta anterior y

la estriación espiral suele ser más marcada.

Dadas las diferencias de esta especie

con los géneros conocidos de morfología

similar, los antecedentes de que en la

Península Ibérica nuevos géneros han

sido recientemente establecidos, como

Tarraconia, Boetersiella, y otros (ver

RAMOS ET AL., 2000, ARCONADA Y

RAMOS, 2001, 2002), no sería sorpren-

dente que esta nueva especie, junto con

“Paladilhiopsis” septentrionalis y otras de

las descritas a continuación pudiesen

pertenecer a nuevos géneros.

Sin embargo, por la carencia de partes

blandas en el material estudiado, hemos

decidido no crear nuevos taxones supra-

específicos y mantener tentativamente su

inclusión en el género Bythiospeum Bour-

guignat, 1882, que se extiende por Suiza,

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

200 um

20 um

100 um

50 um

Figuras 2-8. “Bythiospeum” gloriae spec. nov. 2: holotipo, 2,1 mm (MNCN); 3: concha, 1,9 mm

(CER); 4: detalle de la abertura, paratipo (CER); 5: protoconcha, paratipo (CER); 6: microescul-

tura de la protoconcha; 7: microescultura de las primeras vueltas de la teloconcha; 8: detalle de la

microescultura de la teloconcha.

Figures 2-8. “Bythiospeum” gloriae spec. nov. 2: holotype, 2.1 mm (MNCN); 3: shell, 1.9 mm (CER);

4: detail of the aperture, paratype (CER); 5: protoconch, paratype (CER); 6: microsculpture of the pro-

toconch; 7: microsculpture of the first whorls of' the teleoconch; 8: detail of the microsculpture of the tele-

oconch.

195

Iberus, 21 (1), 2003

Alemania y Austria (ZILCH, 1970 y BER-

NASCONI, 1990), y cuyas conchas parecen

presentar una silueta similar, aunque no

coincidan en la microescultura.

Diferencias con otras especies: En

cuanto a las diferencias con especies ya

conocidas de áreas más o menos próxi-

mas, y cuya asignación genérica tam-

poco esté bien definida, hay que estable-

cerlas con “Paladilhiopsis” septentrionalis

Rolán y Ramos, 1996, conocida sólo por

conchas vacías y descrita del norte de la

Península Ibérica. Esta especie tiene una

microescultura con pequeños hundi-

mientos en la protoconcha y surcos en

zigzag en la teloconcha; además, el

borde de la abertura está en un plano, se

mantiene un poco separado de la vuelta

anterior y apenas sufre eversión.

Todas las especies españolas asigna-

das a los géneros Alzoniella Giusti y Bodon,

1984, Belgrandiella Wagner, 1927 y Bythi-

nella Moquin-Tandon, 1855, presentes en

el norte y oeste de la Península Ibérica,

carecen de microescultura en la superfi-

cie externa de la teloconcha y en cambio

tienen depresiones en la protoconcha; su

abertura está en un plano, contacta amplia-

mente con la vuelta anterior, carecen de

ombligo y el peristoma no está evertido.

“Mottessieria” juvenisanguis Boeters y

Gittenberger, 1980, se encuentra en el oeste

del Pirineo francés y se diferencia por tener

vueltas de espira más convexas, que crecen

más rápidamente, la abertura con ondu-

laciones más marcadas y, además, presenta

una microescultura espiral de la telocon-

cha formada por cordoncillos.

Sardopaladilhia plagigeyerica Manga-

nelli, Bodon, Cianfanelli, Talenti y

Giusti, 1998 tiene el borde externo de la

abertura más sobresaliente de la silueta

de la concha y prácticamente carece de

escultura espiral en la teloconcha mien-

tras en la protoconcha está formada por

pequeñas depresiones.

Género Sardopaladilhia Manganelli, Bodon, Cianfanelli, Talenti y Giusti, 1998

Especie tipo: S. plagigeyerica Manganelli, Bodon, Cianfanelli, Talenti y Giusti, 1998 (por original

designación y monotipia).

Comentarios: Las características mor-

fológicas de este género son: espira algo

elevada; abertura con peristoma sencillo

y continuo, algo evertido como en una

trompeta; la abertura se adhiere en una

zona pequeña a la vuelta anterior y, en

este punto, se hace un poco más promi-

nente con respecto al resto del borde; en

la parte superior de la abertura, un poco

hacia su parte externa, hay una escota-

dura ligera, que desvía hacia atrás el

borde de la misma; el peristoma sobre-

sale de forma clara de la silueta de la

teloconcha; escultura espiral de puntos

o líneas (poco marcada en la especie

tipo); protoconcha con su final mal deli-

mitado con la teloconcha y con escultura

poco pronunciada.

Probablemente, “Moitessieria” juveni-

sanguis Boeters y Gittenberger, 1980 per-

tenece a este género, ya que ni el peris-

toma, sobresaliente y onduloso, ni la

existencia de ombligo, ni la escultura

carente de depresiones, parecen ser

caracteres que se encuentren en la

especie tipo de Mottessieria.

Sardopaladilhia marianae spec. nov. (Figs. 9-21)

Material tipo: Holotipo (Fig. 9) depositado en el MNCN, n* 15.05/46587. Paratipos en las siguien-

tes colecciones: MNCN (1 c, Fig. 10), AMNH ( c, Fig. 11), BMNH (1 c), MNAHN (1 c, Fig. 12), MVHN

(44 c, n* 806), MZB (2 c, n* 2002-692), NNMN (2 c, n* 94819) y CER (90 c, 56 f, Figs. 16-19).

Otro material estudiado: 16 c, de Cabanes (J. Albesa leg.) (MVHN n*%804).

Localidad típica: Piscina de Sueras (Castellón).

Etimología: El nombre específico se dedica a Mariangeles Ramos “Marian”, quién inició con noso-

tros hace años este trabajo pero sus múltiples ocupaciones le impidieron concluirlo.

196

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

50 um

Figuras 9-21. Sardopaladilhia marianae spec. nov. 9: holotipo, 3,4 mm (MNCN); 10: paratipo,

3,4 mm (MNCN); 11: paratipo, 3,4 mm (AMNH); 12: paratipo, 3,2 mm (MNHN); 13: concha,

3,3 mm (CER); 14, 15: detalle de la abertura; 16: protoconcha, paratipo (CER); 17: detalle de la

protoconcha; 18, 19: microescultura de la protoconcha; 20, 21: microescultura de la teloconcha.

Figures 9-21. Sardopaladilhia marianae spec. nov. 9: holotype, 3.4 mm (MNCN); 10: paratype, 3.4

mm (MNCN); 11: paratype, 3.4 mm (AMNH); 12: paratype, 3.2 mm (MNHN); 13: shell, 3.3 mm

(CER); 14, 15: detail of the aperture; 16: protoconch, paratype (CER); 17: detail of the protoconch; 18,

19: microsculpture of the protoconch; 20, 21: microsculpture of the teleoconch.

Iberus, 21 (1), 2003

Descripción: Concha (Figs. 9-13)

cónica, hialina, con una máxima dimen-

sión entre 3,0 y 3,7 mm, con ápice romo,

y un número de vueltas de espira com-

prendido entre 6 y 7, siendo estas

vueltas convexas, aparentemente lisas,

con superficie no brillante y sutura pro-

funda. El crecimiento de la espira es

lento pero continuo. Ocasionalmente

aparecen vueltas que tienen un abomba-

miento más acusado que las demás.

Líneas de crecimiento ligeramente opis-

toclinas, finas y numerosas. Ombligo

bien definido (Fig. 15). Abertura de

forma ligeramente oval, sobresaliendo

de la silueta de la concha. El peristoma

es sencillo y fino, y está algo evertido,

como en una trompeta. La parte supe-

rior del peristoma está adherida a la

vuelta anterior en un corto sector. En

una visión lateral (Figs. 11, 14), este

borde no está en un mismo plano en su

totalidad y se encuentra algo más pro-

longado en la zona de contacto con la

pared de la última vuelta, mientras, algo

más arriba y afuera, el borde está

curvado como en una escotadura (Figs.

11, 14), existiendo otra más suave en su

parte más interna (Fig. 15).

La protoconcha (Figs. 16, 17) tiene

un número de vueltas difícil de precisar

porque no existe un límite claro con la

teloconcha. El núcleo mide 135-140 um,

y tiene una microescultura (Fig. 18)

formada por concavidades irregulares

con bordes elevados; esta escultura se

cambia a continuación (Fig. 19) por otra

que tiene pequeñas depresiones rodea-

das por zonas elevadas irregulares. Poco

después de algo más de una vuelta

desde el núcleo, esta escultura se cambia

por otra en la que aparecen finísimos

cordoncillos espirales formados por la

fusión de tubérculos (Figs. 18, 20). Una

vuelta después, aproximadamente, la

escultura ha cambiado ligeramente, per-

sistiendo una estriación espiral formada

por tubérculos fusionados, que forman

cortos trazos, pero con una densidad

inferior a la anterior (Fig. 21).

Dimensiones: el holotipo mide 3,4 x

1,7 mm. Una buena parte de las conchas

estudiadas alcanzan estas dimensiones, e

incluso alguna llegó a medir 3,76 mm de

altura siendo otras algo más pequeñas.

El animal es desconocido.

Discusión: La asignación genérica de

esta especie se basa en la gran similari-

dad de la concha con la de S. plagigeye-

rica (especie tipo del género) por lo que,

incluso con la carencia de datos anató-

micos, la relación con el género Sardopa-

ladilhia nos parece extremadamente pro-

bable, y creemos justificada la asigna-

ción categórica a este género.

Las diferencias específicas con S. pla-

gigeyerica se basan en que esta especie

tiene una concha un poco menos alar-

gada, la microescultura de la protocon-

cha tiene depresiones mucho menos

marcadas y la microescultura de la telo-

concha es mucho menos evidente

(BODON Y GIUSTI, 1991).

“Moitessieria” juvenisanguis tiene una

forma similar, pero su abertura es

menos evertida mientras la escotadura

superior de la abertura es mayor; tiene

una diferente microescultura de la pro-

toconcha (líneas de tubérculos en vez de

depresiones) y de la teloconcha (filetes

espirales bien diferenciados).

Por su mayor tamaño y su forma

más alargada y cónica se diferencia

fácilmente de “Paladilhiopsis” septentrio-

nalis Rolán y Ramos, 1996, y de “Bythios-

peum” gloriae spec. nov.

“Sardopaladilhia” buccina spec. nov. (Figs. 22-32)

Material tipo: Holotipo (Fig. 22) depositado en el MNCN, n? 15.05 /46588. Paratipos en las siguien-

tes colecciones: MNCN (10 c), AMNH (1 c, Fig. 23), BMNH (1 c), MNAHN (1 c, Fig. 24), MVHN (156

c, n* 807), MZB (5 c, 2002-693), NNMN (5 c, n* 94820) y CER (84 c, 10 f, Fig. 25).

Otro material estudiado: Más de 500 conchas y otros tantos fragmentos de la localidad tipo.

Localidad típica: Font de Castro, Sueras (Castellón).

Etimología: El nombre específico proviene de la palabra latina buccina que significa trompeta,

haciendo referencia a la forma de la concha y a la abertura con el peristoma evertido.

198

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

Figuras 22-32. “Sardopaladilhia” buccina spec. nov. 22: holotipo, 2,4 mm (MNCN); 23: paratipo, 2,4

mm (AMNH); 24: paratipo, 2,2 mm (MNHN); 25: paratipo, 2,5 mm (CER), 26: visión inferior

(MNCN); 27: visión superior (UNCN); 28: protoconcha, paratipo (CER); 29: detalle de la proto-

concha (CER); 30, 31: microescultura de la protoconcha; 32: microescultura de la teloconcha.

Figures 22-32. “Sardopaladilhia” buccina spec. nov. 22: holotype, 2.4 mm (MNCN); 23: paratype,

2.4 mm (AMNAH); 24: paratype, 2.2 mm (MNHN); 25: paratype, 2.5 mm (CER), 26: visión inferior

(MNCN); 27: apical vision (MNCN); 28: protoconch, paratype (CER); 29: detail of the protoconch

(CER); 30, 31: microsculpture of the protoconch; 32: microsculpture of the teleoconch.

199

Iberus, 21 (1), 2003

Descripción: Concha (Figs. 22-25)

cónica, hialina, con una máxima dimen-

sión entre 2,1 y 2,6 mm de altura y 1,53

mm de diámetro, con ápice romo, y un

número de vueltas de espira de 4 a 4

3/4, siendo estas vueltas muy convexas,

aparentemente lisas, con superficie no

brillante y sutura profunda. En ocasio-

nes se observan pequeñas irregularida-

des o deformaciones de las vueltas, que

modifican la regularidad del perfil de la

concha. Líneas de crecimiento finas,

poco manifiestas y ligeramente opisto-

clinas. La abertura es redondeada, lige-

ramente oval, con el borde sencillo y

fino, pero algo girado hacia afuera

como en una trompeta, y estando su

parte superior e interna en contacto con

la vuelta anterior, únicamente en un

corto sector. En una visión lateral (Figs.

24, 26) se aprecia que no todo el borde

labial se encuentra en el mismo plano,

sino que se prolonga ligeramente en la

parte en contacto con la vuelta anterior

(Fig. 26), mientras por arriba hacia la

parte externa de la abertura (Figs. 24,

26, 27) se ondula, formando una

pequeña escotadura, que se retrasa lige-

ramente. Hay un ombligo apreciable

que, en una visión anterior, queda par-

cialmente oculto por la eversión de la

abertura.

La protoconcha (Fig. 28) tiene un

límite no bien apreciable con la telo-

concha. El núcleo mide entre 140-150

um.

-Microescultura: La protoconcha (Figs.

29, 31) presenta una escultura formada,

en su comienzo, por líneas que se

colocan espiralmente en forma de

espigas, transformándose a continua-

ción en tubérculos que se agrupan for-

mando líneas espirales, y que, en la

parte media e inferior de la vuelta, se

fusionan (Fig. 30) dando lugar a estruc-

turas espiralmente alargadas. En la telo-

concha (Fig. 32), la microescultura está

formada por tubérculos que adoptan

una cierta alineación espiral y que, en

muchos lugares, se encuentran fusiona-

dos con otros próximos formando líneas

irregulares y produciendo dibujos ara-

bescos.

200

El holotipo mide 2,4 x 1,1 mm.

Alguna concha puede ser ligeramente

más grande.

El animal es desconocido.

Discusión: La asignación genérica

está basada en la semejanza de algunos

caracteres de de la concha de esta

especie con la de S. plagigeyerica, la

especie tipo del género que, sin

embargo, tiene la abertura menos ever-

tida en su borde y, en cambio, sobre-

sale más del perfil de la concha.

Además tiene microescultura con

depresiones en la protoconcha y, en la

teloconcha, la escultura es menos

manifiesta. Por estas diferencias, esta

asignación no puede ser más que

tentativa.

De Sardopaladilhia marianae spec. nov.

se diferencia porque esta especie es más

grande, más cónica, con el peristoma

más sobresaliente del perfil de la concha

y la microescultura de la protoconcha

está formada depresiones mientras que

la de la teloconcha tiene tubérculos en

vez de pequeños trazos.

Las diferencias con otras especies

conocidas de la Península Ibérica hay

que establecerlas con “Paladilhiopsis”

septentrionalis, que tiene menos escul-

tura, formada por líneas en zigzag, y el

peristoma se separa de la vuelta anterior

manteniéndose todo el borde en un

plano.

“Moitessieria” juvenisanguis Boeters

y Gittenberger, 1980, tiene una silueta

más alargada, la escotadura de la aber-

tura es más pronunciada, la abertura

sobresale más de la silueta de la

concha y tiene una microescultura de

la protoconcha formada por líneas de

tubérculos mientras la de la teloconcha

está formada por cordoncillos bien

definidos.

“Bythiospeum” gloriae spec. nov., que

vive en simpatría en la localidad típica,

se diferencia porque en su espira las

vueltas tienen menor velocidad de creci-

miento, y por lo tanto su forma es más

esbelta por ser menos ancha en la última

vuelta; además la abertura es menos

sobresaliente, y su escultura es más ate-

nuada.

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

100 um 100 um

Figuras. 33-45. “Sardopaladilhia” distorta spec. nov. 33: holotipo, 2,0 mm (MNCN); 34: paratipo,

2,3 mm (AMNH); 35: paratipo, 1,9 mm (MNHN); 36: paratipo, 2,0 mm (CER); 37-40: parati-

pos, 2,1, 2,1, 1,8 y 2,0 mm (MNCN); 41: paratipo, 1,9 mm (CER); 42: protoconcha, paratipo

(CER); 43: microescultura de la protoconcha; 44, 45: microescultura de la teloconcha.

Figures. 33-45. “Sardopaladilhia” distorta spec. nov. 33: holotype, 2.0 mm (MNCN); 34: paratype,

2.3 mm (AMNH); 35: paratype, 1.9 mm (MNHN); 36: paratype, 2.0 mm (CER); 37-40: paratypes,

2.1, 2.1, 1.8 y 2.0 mm (MNCN); 41: paratype, 1.9 mm (CER); 42: protoconch of a paratype (CER);

43: microsculpture of the protoconch; 44, 45: microsculpture of the teleoconch.

201

Iberus, 21 (1), 2003

“Sardopaladilhia” distorta spec. nov. (Figs. 33-45)

Material tipo: Holotipo (Fig. 33) depositado en el MNCN, n? 15.05/46589. Paratipos en las siguien-

tes colecciones: MNCN (4 c, Figs. 37-40), AMNH (1 c, Fig. 34), BMNH (1 c), MNHN (1 c, Fig. 35),

MVHN (7 c, n* 808), MZB (1 c, n* 2002-694), NNHM (1 c, n* 94821) y CER (44 c, Figs. 36, 41).

Otro material examinado: 17 f, de la localidad típica.

Localidad típica: Piscina de Sueras (Castellón).

Etimología: El nombre específico hace alusión a la forma de la concha con desviaciones del eje de

la espira que le da un aspecto contrahecho.

Descripción: Concha (Figs. 33-41)

oval-cónica, hialina, con una máxima

dimensión entre 1,8 y 2,4 mm, con ápice

romo, y un número de vueltas de espira

de 4-4 !/2, siendo estas vueltas muy con-

vexas, con superficie no brillante y

sutura profunda. Existen frecuentes irre-

gularidades o deformaciones de las

vueltas, que dan la impresión de no

adaptarse exactamente al eje de la

concha, O variar su eje en momentos dis-

tintos. Sin escultura aparente, excepto

líneas de crecimiento que son finas y

numerosas, algo sinuosas, especial-

mente hacia la base; parecen ortoclinas o

ligeramente prosoclinas, en especial al

final de la espira cuando la vuelta está

distorsionada. La última vuelta de

espira se proyecta hacia afuera y un

poco hacia arriba, sobresaliendo nota-

blemente de la silueta de la concha (Fig.

39, 41). La abertura es redondeada, lige-

ramente piriforme, con el peristoma

sencillo y fino, pero algo evertido hacia

fuera, como en una trompeta, y mante-

niendo un espacio muy corto de su

parte interna y superior en contacto con

la vuelta anterior. En una visión lateral

(Figs. 34, 35) el borde del peristoma se

muestra ondulado, con una depresión

en la parte superior y otra en la más

interna. Ombligo abierto y profundo.

La protoconcha (Figs. 41, 42) no pre-

senta una clara separación con la telo-

concha, aunque se aprecia, en algunas

ocasiones, un cambio en la microescul-

tura después de algo más de una vuelta.

El núcleo mide unas 130 um.

Microescultura: en la protoconcha, la

escultura está formada por un dibujo

irregular en forma de pequeñas excava-

ciones y lineas espirales, apenas insi-

nuadas, dispuestas ligeramente en

sentido espiral (Fig. 43), apareciendo al

202

comienzo de la teloconcha cordoncillos

espirales finos que recorren toda la

concha haciéndose más evidentes (Figs.

44, 45) y que están cruzados por nume-

rosas estrías de crecimiento.

Dimensiones: el holotipo mide 2,0 x

1,7 mm. Algunas conchas pueden alcan-

zar hasta 2,4 mm y 1,8 de diámetro.

El animal es desconocido.

Discusión: La posición genérica de

esta especie en Sardopaladilha, es tenta-

tiva, y está basada en la abertura con el

borde del peristoma evertido y con

ondulaciones semejantes a las que apa-

recen en las especies antes descritas, así

como una microescultura más o menos

similar en protoconcha y teloconcha.

Otros géneros que podrían haber

sido utilizados pero fueron descartados

para esta especie fueron los siguientes:

e Bythiospeum Bourguignat, 1882

(ver anteriormente) tiene conchas con

espira regular y la abertura carece de

una gran eversión.

e Paladilhiopsis Pavlovic, 1913. (ver

anteriormente) ocurre lo mismo que con

el género anterior.

e Iglica Wagner, 1927 (especie tipo 1.

gratulabunda Wagner, 1927) no parece el

adecuado porque (según GIUSTI Y

PEZZOLI, 1980) tiene forma alargada,

sutura no profunda, ombligo apenas

visible, peristoma no expandido y apenas

sinuoso, sin microescultura externa.

e Moitessieria Bourguignat, 1863

(especie tipo: M. simoniana Saint-Simon,

1848) tiene la concha muy uniforme-

mente alargada, la abertura está adhe-

rida en parte a la vuelta anterior, no hay

ombligo evidente y el peristoma no está

o está muy ligeramente evertido; su

microescultura está formada por perfo-

raciones espirales aunque, a veces, tiene

cordoncillos. Según BERNASCONI (1984)

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

100 um

200 um

Figs. 46-57. “Sardopaladilhia” subdistorta spec. nov. 46: holotipo, 1,8 mm (MNCN); 47: paratipo,

2,1 mm (AMNH); 48: paratipo, 2,3 mm (BMNH); 49: paratipo, 2,05 mm (CER); 50: paratipo, 2,0

(CER); 51, 52: paratipos, 2,0 y 2,2 mm (MNCN); 53: paratipo, 1,9 mm (CER); 54: protoconcha

(CER); 55: detalle de la microescultura de la protoconcha; 56, 57: microescultura de la teloconcha.

Figs. 46-57. “Sardopaladilhia” subdistorta spec. nov. 46: holotype, 1.8 mm (MNCN); 47: paratype,

2.1 mm (AMNA); 48: paratype, 2.3 mm (BMNA); 49: paratype, 2.05 mm (CER); 50: paratype, 2.0

(CER); 51, 52: paratypes, 2.0 y 2.2 mm (MNCN); 53: paratype, 1.9 mm (CER); 54: protoconch

(CER); 55: detail of the microsculpture of the protoconch; 56, 57: microsculpture of'the teleoconch.

203

Iberus, 21 (1), 2003

y BODON Y GIUSTI (1991) Spiralix Boeters,

1972 sería un sinónimo.

e Clameia Boeters y Gittenberger,

1990 (especie tipo C. brooki Boeters y Git-

tenberger, 1990) tiene una concha uni-

formemente alargada, el peristoma está

evertido como en una trompeta, pero

está separado de la vuelta anterior. La

microescultura está formada por líneas

espirales formadas por segmentos obli-

cuos conectados.

e Paladilhia Bourguignat, 1865

(especie tipo P. pleurotoma Bourguignat,

1865) tiene la abertura piriforme con el

borde interno del peristoma rectilíneo y

en posición oblicua al eje de la concha,

el ombligo está casi cerrado por el peris-

toma y la microescultura no existe o es

casi inexistente (LOCARD, 1893, BODON Y

Giusti, 1991).

e Lartetia Bourguignat, 1869 (especie

tipo L. belgrandi (Bourguignat, 1869)

tiene la abertura bastante adherida a la

vuelta anterior y el ombligo aparece

aplastado.

Por las razones explicadas y no con-

siderando su posible pertenencia a los

géneros mencionados, decidimos su

inclusión provisionalmente en Sardopa-

ladilhia que parece reunir más caracteres

similares, aún teniendo en cuenta ciertas

diferencias que presenta “S”. distorta,

como son las desviaciones del eje de la

espira, la dirección hacia arriba del final

de la última vuelta y la microescultura

que es un poco diferente de la de la

especie típica.

La diferenciación de “S”. distorta de

la especie tipo del género, S. plagigeye-

rica, se basa en que esta última especie

tiene una espira con crecimiento más

lento, con menor tendencia a la distor-

sión, la última vuelta no tiende a ele-

varse y la microescultura de la telocon-

cha es muy poco aparente.

Con las especies consideradas con-

genéricas y que se han descrito anterior-

mente, se diferencia por su desviación

del eje de la concha y elevación final de

la última vuelta.

“Sardopaladilhia” subdistorta spec. nov. (Figs. 46-57)

Material tipo: Holotipo (Fig. 46) depositado en el MNCN, n? 15.05 /46590. Paratipos en las siguien-

tes colecciones: MNCN (4 c, Figs. 51, 52), AMNH (1 c, Fig. 47), BMNH (1 c, Fig. 48), MNHN (1 c),

MVHN (64 c, n* 809), MZB (5 c, n* 2002-695), NNMN (5 c, n* 94822) y CER (133 c, 6 f, Figs. 49, 50,

53).

Otro material examinado: más de 100 c, y de 250 fragmentos, de la localidad típica.

Localidad típica: Font de Castro, Sueras (Castellón).

Etimología: El nombre específico hace alusión a la forma de la concha, similar a S. distorta, pero

con desviaciones menores y menor aspecto de concha contrahecha.

Descripción: Concha (Figs. 46-53)

oval cónica, hialina, con una máxima

dimensión entre 1,8 y 2,3 mm, con ápice

romo, y un número de vueltas de espira

comprendido entre 3 */2 y 4 */2, siendo

estas vueltas muy convexas, con super-

ficie no brillante y sutura profunda. Las

vueltas siguen el mismo eje de la

concha, aunque no son extremadamente

regulares, y a veces parece que se

separan ligeramente de este eje. No hay

una escultura aparente, excepto líneas

de crecimiento que son finas y numero-

sas, prosoclinas y algo sinuosas, espe-

cialmente hacia la base. La última vuelta

de espira es prominente, sobresaliendo

204

un poco de la silueta de la concha. La

abertura es redondeada, ligeramente

oval, con el peristoma sencillo y fino, y

ligeramente evertido hacia afuera; hay

un espacio muy corto de su parte

interna y superior en contacto con la

vuelta anterior. En una visión lateral

(Fig. 51) el borde del peristoma se

muestra un poco ondulado, con una

depresión en la parte superior y otra en

la más interna (Fig. 52). Ombligo abierto

y profundo.

La protoconcha (Fig. 54) tiene poco

preciso el punto de separación con la

teloconcha aunque, en algunas ocasio-

nes, se puede apreciar un cambio en la

ROLÁN Y MARTÍNEZ-ORTÍ: Nuevos Hydrobiidae de la Comunidad Valenciana

microescultura un poco después de la

primera vuelta de espira. El núcleo mide

alrededor de 157 um.

Microescultura: en la protoconcha, la

escultura está formada por un dibujo

irregular en forma de pequeñas escava-

ciones, apenas insinuadas, y diminutos

trazos espirales (Fig. 55); un poco más

adelante aparecen cordoncillos espirales

finos que recorren toda la concha

haciéndose más evidentes (Figs. 56, 57)

aunque a veces se interrumpen al ser

cruzados por las estrías de crecimiento.

Dimensiones: el holotipo mide 1,8 x

1,3 mm. Otras conchas llegan a medir

2,3 mm de altura y 1,61 de diámetro.

El animal es desconocido.

Discusión: La especie más próxima a

“5”. subdistorta spec. nov. es “S”. distorta

spec. nov., pero se diferencia porque

esta última especie es proporcional-

mente más ancha, sus primeras vueltas

AGRADECIMIENTOS

A Gloria Tapia quién remitió el

primer material para su estudio. A Jesús

Méndez de los Servicios Generales de

Apoyo a la Investigación de la Universi-

dad de Vigo por las fotografías realiza-

das al MEB. También a Joaquín Albesa,

quién nos proporcionó material de

Cabanes, y Sergio Montagut, por su

colaboriación en los distintos muestreos

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O Sociedad Española de Malacología —_—_—_——— Iberus, 21 (1): 207-240, 2003

The deep-water Indo-Pacific radiation of Fusinus

(Chryseofusus subgen. nov.) (Gastropoda: Fasciolariidae)

La radiación Indo-Pacífica de aguas profundas del género Fusinus

(Chryseofusus subgen. nov.) (Gastropoda: Fasciolariidae)

Roland HADORN* and Koen FRAUSSEN**

Recibido el 2-XII-2002. Aceptado el 11-111-2003

ABSTRACT

A number of fusinids from the Indo-Pacific deep-water fauna are studied to get more

insight in the distribution and variability. The subgenus Chryseofusus (Gastropoda: Fascio-

lariidae: Fusinus Rafinesque, 1815) is described as new to accommodate a number of

species sharing conchological characteristics different from typical Fusinus. Their separa-

tion from Fusinus s.s. is based on differences in axial sculpture (usually absent on body

whorl), spiral sculpture (weak, close-set, regular, crossed by distinct growth lines), shape

(shorter spire, shorter siphonal canal, less convex whorls with subsutural concavity, less

constricted suture) and parietal callus [inner lip smooth, parietal wall covered with an

extended, adherent thin layer of callus).

Fusinus (Chryseofusus) bradneri (Drivas and Jay, 1990), F. (C.) chrysodomoides (Schep-

man, 1911), F. (C.) graciliformis (Sowerby, 1880), FE. (C.) hyphalus M. Smith, 1940, F.

(C.) jurgeni Hadorn and Fraussen, 2002, F. (C.) kazdailisi Fraussen and Hadorn, 2000

and F. (C.) subangulatus (von Martens, 1901) are briefly described and their taxonomic

placement in the new subgenus is discussed.

To avoid further taxonomic complications, a lectotype is designated for the correct F. (C.)

chrysodomoides.

F. (C.) acherusius (west Madagascar, Mozambique Channel, 1475-1530 m), F. (C.) alisae

(north New Caledonia, 444-452 m), F. (C.) artutus (Philippines, Bohol, deep water), F. (C.)

cadus [south New Caledonia, 460-470 m], F. (C.) dapsilis (Vietnam, deep water), F. (C.)

riscus (New Caledonia, Norfolk Ridge, 394-401 mj), F. (C.) scissus [south New Caledonia,

535 m), F. (C.) wareni (New Caledonia, 480 m), and F. (C.) westralis (northwest Aus-

tralia, off Port Hedland, 450 m) are described as new to science.

RESUMEN

Se estudia un grupo de fusínidos de la fauna de aguas profundas del Indo-Pacífico, con el

fin de profundizar en su distribución y variabilidad. Se describe el subgénero Chryseofu-

sus (Gastropoda: Fasciolariidae: Fusinus Rafinesque, 1815) para incluir a un número de

especies que comparten caracteres conquiológicos distintos de los típicos Fusinus. Su

separación de Fusinus s.s. se basa en diferencias en la escultura axial (normalmente

ausente en la espira del cuerpo), escultura espiral (debil, compacta, regular, atravesada

por claras líneas de crecimiento), forma (espira más corta, canal sifonal más corto, vueltas

menos convexas con concavidad subsutural, sutura menos constreñida) y callo parietal

(labio interno liso, pared parietal cubierta con una fina capa de callo adherente).

* Schuetzenweg 1, CH-3373 Roethenbach, Switzerland.

** Leuvensestraat 25, B-3200 Aarschot, Belgium.

207

Iberus, 21 (1), 2003

- Fusinus (Chryseofusus) bradneri (Drivas y Jay, 1990), F. (C.) chrysodomoides [Schepman,

1911), F (C.) graciliformis (Sowerby, 1880), F. (C.) hyphalus M. Smith, 1940, F. (C.) jurgeni

Hadorn y Fraussen, 2002, F. (C.) kazdailisi Fraussen y Hadorn, 2000 y F. (C.) subangulatus (von

Martens, 1901) se describen brevemente, y se discute su adscripción al nuevo subgénero.

Para evitar futuras complicaciones taxonómicas, se designa un lectotipo para la forma

correcta de F. (C.) chrysodomoides.

F. (C.) acherusius (Madagascar oeste, Mozambique Channel, 1475-1530 m), F. (C.) ali-

sae [N de Nueva Caledonia, 444-452 m), F. (C.) artutus (Filipinas, Bohol, aguas profun-

das), F. (C.) cadus [(S de Nueva Caledonia, 460-470 m), F. (C.) dapsilis (Vietnam, aguas

profundas), F. (C.) riscus (Nueva Caledonia, Norfolk Ridge, 394-401 m), F. (C.) scissus [S

de Nueva Caledonia, 535 m), F. (C.) wareni (Nueva Caledonia, 480 m), y F. (C.) westra-

lis (NO de Australia, frente a Port Hedland, 450 m) se describen como especies nuevas.

KEY WORDS: Mollusca, Gastropoda, Fasciolariidae, Fusinus, Chryseofusus, MUSORSTOM, Indo-Pacific, new

subgenus, new species.

PALABRAS CLAVE: Mollusca, Gastropoda, Fasciolariidae, Fusinus, Chryseofusus, MUSORSTOM, Indo-

Pacífico, nuevo subgénero, nuevas especies.

INTRODUCTION

During French expeditions, called

CAMPAGNES MUSORSTOM, conducted

by ORSTOM (New Caledonia) and

MNHN (Paris) in the Indo-West Pacific

and in particular in the seas around New

Caledonia, a huge number of interesting

species, many of them new to science, are

collected. The Fasciolariidae are well rep-

resented in this rich material.

The purpose of the present paper is

to report on this material, starting with

the study of a number of peculiar

fusinids from the upper bathyal zone of

the Indo-Pacific, which we assign to the

new subgenus Chryseofusus. Thanks to

the large quantity of available material, a

profound study was possible, comparing

material from all over the Indo-West

Pacific. A number of species formerly

considered to be endemic, or restricted

to a certain area, are now found to have

a much more extensive range, a situation

also observed by both authors in some

lower bathyal Buccinidae. We must con-

clude the bathyal fauna of some areas is

still poorly known and information

about distribution is tentative for many

bathyal Neogastropoda. In the present

paper an effort is made to produce

correct information on the distribution of

Chryseofusus species. As a result F. val-

208

diviae Hadorn and Fraussen, 1999 (type

locality Somalia) is synonymized with EF.

graciliformis (Sowerby, 1880) (type local-

ity Japan). The west Australian species

commonly assigned to “Siphonofusus

chrysodomoides” is found to be different

from the Indonesian type material of

Fusus chrysodomoides Schepman, 1911,

and herein described as new.

MATERIAL AND METHODS

The present study is essentially

based on the material collected by

French research vessels and expeditions

in the tropical Indo-West Pacific during

the last 30 years:

(a) Material from several expeditions

and surveys in the South-West Indian

Ocean: BENTHEDI (1977) on board R.V.

Suroit in the northern part of the Mozam-

bique Channel; MD32 (1982) on board

R.V. Marion-Dufresne around Réunion ls.;

shrimp surveys conducted in the

Mozambique Channel by A. Crosnier in

1972-74 on board R.V. Vauban, and by R.

von Cosel in 1986 on board commercial

trawler Mascareignes III; as well as mater-

ial recently obtained as a by-product of

commercial shrimp fisheries.

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

(b) Material from numerous expedi-

tions in the New Caledonia region since

1984 (BIOCAL, MUSORSTOM 4-6,

CHALCAL 2, CALSUB, SMIB 1-8,

BATHUS 1-4, BERYX 11); we refer to

RICHER DE FORGES (1990, 1993b), RICHER

DE FORGES AND CHEVILLON (1996) and

Roux (1994) for a narrative of these

cruises and station lists.

other mainly West Pacific regions: In-

donesia (CORINDON 2; KARUBAR, see

CROSNIER, RICHER DE FORGES AND

BOUCHET, 1997); Vanuatu (MU-

SORSTOM 8, see RICHER DE FORGES,

FALIEX AND MENOU, 1996); Wallis and

Futuna (MUSORSTOM 7, see RICHER DE

FORGES, 1993a); Fiji (MUSORSTOM 10,

BORDAU 1, see RICHER DE FORGES,

NEWELL, SCLACHER-HOENLINGER,

SCHLACHTER, NATING, CÉSA AND

BOUCHET, 2000; RICHER DE FORGES,

BOUCHET, DAYRAT, WARÉN AND

PHILIPPE, 2000) and Tonga (BORDAU 2).

Material from these expeditions is,

unless otherwise stated, deposited in

MNHN. No individual catalogue

number is allocated, but material is

unambiguously designated (and retriev-

able) by the combination of expedition

acronym and station number.

The method of VERDUIN (1977) was

employed to count the number of proto-

conch whorls.

Abbreviations used in this paper:

AMS Australian Museum, Sydney, Aus-

tralia

BMNH The Natural History Museum,

London, Great Britain

IMT Institute of Malacology, Tokyo,

Japan

IRD Institut de recherche pour le devel-

oppement, Nouméa, New Caledonia

SYSTEMATICS

KMMA Klaipeda Maritime Museum

and Aquarium, Klaipeda, Lithuania

MNHN Muséum national d'Histoire

naturelle, Paris, France

NM Natal Museum, Pietermaritzburg,

South Africa

NMBE Naturhistorisches Museum Bern,

Switzerland

NMNZ Museum of New Zealand Te

Papa Tongarewa, Wellington, New

Zealand

NSMT National Science Museum,

Tokyo, Japan

RMNH National Museum of Natural

History — Naturalis, Leiden, The

Netherlands

SMNH Swedish Museum of Natural

History, Stockholm, Sweden

USNM National Museum of Natural

History, Smithsonian Institution,

Washington D.C., USA

WAM Western Australian Museum,

Perth, Australia

ZMA Zoologisch Museum, University

of Amsterdam, Amsterdam, the

Netherlands

ZMB Museum fur Naturkunde (Zool.

Museum), Berlin, Germany

KF Collection Koen Fraussen, Aarschot,

Belgium

RH Collection Roland Hadorn, Róthen-

bach, Switzerland

CA (casier) trap

CC (chalut a crevettes) shrimp trawl

CH (chalut) trawl

CP (chalut á perche) beam trawl

DC (drague Charcot) Charcot dredge

DE (drague épibenthique) epibenthic

dredge

DR (drague a roche) rock dredge

DW (drague Warén) Warén dredge

dd dead collected specimen

juv juvenile specimen

lv live collected specimen

subad subadult specimen

Family FASCIOLARIIDAE Gray, 1853

Genus Fusinus Rafinesque, 1815

Fusinus Rafinesque, 1815. Anal. nat. tabl. univ. corps org.: 145. Substitute name for “Fusus Lamarck'

[=Fusus Bruguiere, 1789], non Fusus Helbling, 1779.

209

Iberus, 21 (1), 2003

Type species: Murex cólus Linnaeus, 1758, by typification of replaced name.

Description: Shells fusiform, small to

large in size, elongate, with tall spire

and long siphonal canal. Usually well-

ribbed and with spiral threads. Col-

umellar folds absent. Operculum cor-

neous, shape and size corresponding to

aperture, nucleus terminal.

Radula: with small tricuspid

central tooth, lateral teeth curved,

with 4 to about 15 strong, long point-

ed cusps.

Remarks: Colus Humphrey, 1797 is

invalid (Humphrey has been ruled as

non-binominal by ICZN).

Chryseofusus subgen. nov.

Type species: Fusus chrysodomoides Schepman, 1911 (Figs. 8-9).

Etymology: Derived from the Latin “chryseus” (adjective: made of gold), meaning “a Fusus made

of gold”, after the yellowish coloured periostracum of some of the species. Hereby also evocating

the name of the type species: Fusus chrysodomoides Schepman, 1911.

Diagnosis: Shell small to medium,

usually thin, “light in weight, shape

fusiform. Whorls slightly convex, shoul-

der slope on latter whorls concave, some-

times slightly keeled, suture indistinct.

Rather strong axial ribs on upper whorls,

axial sculpture on latter whorls usually

weak or absent. Spiral sculpture indistinct,

close-set, regular, consisting of weak

primary cords and intercalated fine

threads, crossed by strong, close-set,

curved axial growth lines, giving the

surface the texture of linen. Protoconch

typical of genus, -1 to -1 */4 whorls,

smooth, glossy, last part (—* /4 whorl) often

sculptured with some axial riblets. Aper-

ture ovate, white, yellowish or brownish;

outer lip simple, with or without internal

lirae; inner lip completely smooth, pari-

etal wall covered with an extended, adher-

ent thin layer of callus, never forming a

detached inner lip like in F. colus. Col-

lumellar folds absent. Siphonal canal as

long as aperture or slightly shorter, slightly

curved or straight, open. Shell uniformly

whitish, dirty greyish or brownish. Some

species with pale reddish coloured spiral

sculpture. Operculum typical of genus,

corneous, shape and size corresponding

to aperture, nucleus terminal. Perios-

tracum usually yellowish or brownish,

often well adherent, thick and velvety.

Radula typical of genus, central tooth tri-

cuspid, lateral teeth curved, with 4-8 strong

pointed cusps.

210

Range and habitat: Chryseofusus is eco-

logically a deep water group, accommo-

dating species from the Indo-Pacific

upper bathyal zone, between 100 and

1900 m deep. Until present date we have

not recognized any Atlantic species as

belonging to Chryseofusus.

Species in Chryseofusus (in alphabetical

order): F. acherusius sp. nov., Mozambique.

Channel and New Caledonia in 1530-

1900 m; F. alisae sp: nov., New Caledonia

and Coral Sea in 300-545 m; F. artutus sp.

nov., Philippines, Indonesia and New

Caledonia in 271-435 m; E bradneri (Dri-

vas and Jay, 1990), Reunion, north Mada-

gascar, Glorieuses and Comoro Islands in

300-750 m;, F. cadus sp. nov., New Caledo-

nia in 460-675 m; E. chrysodomoides

(Schepman, 1911), Indonesia, southwest

Pacific and north Madagascar in 179-797

my EF. dapsilis sp. nov., Vietnam, deep wa-

ter; F. graciliformis (Sowerby, 1880), east

Asia and east Africa in 50-600 m;, F. hy-

phalus M. Smith, 1940, Japan, Philippines

and east China Sea in 100-300 m;, F. jur-

geni Hadorn and Fraussen, 2002, south-

west Madagascar in 530 m, F. kazdailisi

Fraussen and Hadorn, 2000, Chile in 150-

1200 m; F. riscus sp. nov., south New

Caledonia in 401-430 m;, F. scissus sp. nov.,

south New Caledonia in 410-580 m; E.

subangulatus (von Martens, 1901), east

Africa in 400-1134 m, F. wareni sp. nov.,

New Caledonia in 480-755 m; F. westralis

sp. nov., Western Australia in 300-500 m.

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Comparison: Chryseofusus differs from

all other subgenera in Fusinus by the

slightly convex, usually unkeeled

whorls with subsutural concavity, the

presence of axial ribs usually only on

upper whorls, the weak, close-set,

regular spiral sculpture crossed by

strong, close-set, curved axial growth

lines, giving the surface the texture of

linen or a sometimes pearled appear-

ance. Outer lip always simple; inner lip

completely smooth, the never detached

parietal callus consists of an extended,

adherent thin layer of callus.

F. bradneri and F. chrysodomoides, now

placed in Chryseofusus on the base of

conchological characteristics and the

fusinid radula, have generally been

referred to the buccinid genus Siphono-

fusus Kuroda and Habe, 1952.

F. graciliformis and E. hyphalus have

been referred to the conchologically

similar genus Simplicifusus Kuroda and

Habe in Kuroda, Habe and Oyama,

1971, but can be distinguished by oper-

culum morphology.

Simplicifusus noguchii (Fig. 1) was

named by Habe and Masuda for all the

. Japanese material previously referred to

as S. simplex (E.A. Smith, 1879). Kuroda

et al. referred to that material when they

designated “Fusus simplex (SmithY as the

type species of Simplicifusus. BIELER AND

PerrT (1996: 33), HIGO, CALLOMON AND

GOTO (1999: 263) and SNYDER (2000: 175)

already reported on this problem. This

matter will be dealt in a future paper by

Snyder (pers. comm. M. A. Snyder).

For comparison we figure the holo-

type of Fusus simplex E.A. Smith, 1879

[BMNH 1878.11.7.27, a juvenile specimen

of 18.0 x 5.5 mm, type locality: south of

Korea] (Figs. 5, 6). It belongs to Fusinus

s.s. and has a typical fusinid operculum

with terminal nucleus (Fig. 7).

For comparison we also figure S.

noguchii Habe and Masuda, 1990 (Fig.

1), and its operculum (Fig. 2) which is

completely different from all other

treated species belonging to the new

subgenus Chryseofusus and all other sub-

genera in Fusinus Rafinesque, 1815,

which all have a typically fusinid oper-

culum with terminal nucleus (Fig. 3).

The operculum of S. noguchii (Fig. 2) is

small, ovate, thin, much smaller than

the aperture, and the nucleus is situated

at lower outer side. It is identical to the

operculae of the genus Granulifusus

Kuroda and Habe, 1954 [Type species:

Fusus niponicus E.A. Smith, 1879] (Fig.

4). Future study may show that Simplici-

fusus may be a junior synonym of Gra-

nulifusus.

Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911) (Figs. 8-13, 76, 86)

Fusus chrysodomoides Schepman, 1911. Rés. Siboga Exp., Mon 49 (1), part 4: 293, pl. 19, fig. 4; pl. 23,

fig. 10.

SUBSEQUENT USE

Siphonofusus chrysodomoides (Schepman, 1911). non KOsuGE (1985: 59, pl. 23, fig. 7), WILSON (1994:

66, pl. 12, figs. 7a-b), HADORN AND FRAUSSEN (1999: pl. 3, figs. 17-18).

Fusinus chrysodomoides (Schepman, 1911). HADORN AND FRAUSSEN (1999: 117, 120).

Type material: Lectotype ZMA, SIBOGA (70.7 x 24.9 mm, lv, preserved with animal in alcohol),

designated herein (Figs. 8-9). — 1 paralectotype ZMA 3.11.021, Indonesia, near Kai Islands, SIBOGA

stn 262, 5” 53' 8” S, 132” 48' 8” E, 560 m (66.9 x 25.6 mm, lv).

Because of the previous confusion with regard to this species and to ensure a taxonomic correct

interpretation in the future, the selection of a lectotype for this species is advisable. We hereby select

the specimen figured by Schepman as lectotype.

Type locality: Indonesia, Molucca-Passage, SIBOGA stn 139, 0? 11' S, 127” 25' E, 397 m.

Material examined: Lectotype and paralectotype of Fusus chrysodomoides in ZMA.

North Madagascar, 12” 50' S, 48” 09 E, 580-585 m, 1 dd/1 dd juv; 12” 43' S, 48” 15' E, 300-340 m, 1

dd. -— North Madagascar, 350-400 m, 1 dd juv/1 dd, collected by commercial boats, RH.

Indonesia, north Makassar Strait, CORINDON Il stn CH276, 01* 55' S, 119* 13' E, 395-450 m, 1

dd. — Tanimbar Islands, KARUBAR stn DW44, 07* 52 S, 132? 48' E, 291-295 m, 1 dd juv. — Kai

Iberus, 21 (1), 2003

Figures 1, 2. Simplicifusus noguchii Habe and Masuda, 1990. 1: RH, Japan, Mikawa, Aichi Pref., 54.8

mm; 2: Operculum. Figure 3. Operculum of Fusinus colus (Linnaeus, 1758). Figure 4: Operculum of

Granulifusus niponicus (E.A. Smith, 1879). Figures 5-7. Fusinus simplex (E.A. Smith, 1879), Holotype

BMNH 1878.11.7.27, south of Korea, 18.0 mm. 7: Operculum. Figures 8-13. Fusinus (Chryseofusus)

chrysodomoides (Schepman, 1911). 8, 9: Lectotype ZMA, Indonesia, Molucca-Passage, 70.7 mm; 10, 11:

Paralectotype ZMA 3.11.021, Indonesia, near Kai Islands, 66.9 mm; 12, 13: MNHN, Fiji, 70.9 mm.

Figuras 1, 2. Simplicifusus noguchii Habe y Masuda, 1990. 1: RH, Japón, Mikawa, Aichi Pref, 54,8

mm; 2: Opérculo. Figura 3. Opérculo de Fusinus colus (Linnaeus, 1758). Figura 4. Opérculo de Granuli-

fusus niponicus (E.A. Smith, 1879). Figuras 5-7. Fusinus simplex (E.A. Smith, 1879), Holotipo

BMNH 1878.11.7.27, S de Corea, 18,0 mm. 7: Opérculo. Figuras 8-13. Fusinus (Chryseofusus) chry-

sodomoides (Schepman, 1911). 8, 9: Lectotipo ZMA, Indonesia, Molucca-Passage, 70,7 mm; 10, 11:

Paralectotipo ZMA 3.11.021, Indonesia, cerca de Kai Islands, 66,9 mm; 12, 13: MNHN, Fiji, 70,9 mm.

212

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Figures 14-21. Fusinus (Chryseofusus) graciliformis (Sowerby, 1880). 14, 15: Holotype BMNH

1880.10.15.2, Japan, 52.5 mm; 16, 17: Holotype of Fusus sieboldí Schepman, 1891 (junior

synonym), RMNH 86858, Japan, 40.0 mm; 18, 19: KF 3200, Philippines, 81.9 mm; 20, 21:

Paratype 1 of Fusinus valdiviae Hadorn and Fraussen, 1999 (junior synonym), KF 2811, Somalia,

47.4 mm. Figures 22, 23. Fusinus (Chryseofusus) hyphalus M. Smith, 1940, RH, Taiwan, 50.3 mm.

Figuras 14-21. Fusinus (Chryseofusus) graciliformis (Sowerby, 1880). 14, 15: Holotipo BMNH

1880.10.15.2, Japón, 52,5 mm; 16, 17: Holotipo de Fusus sieboldi Schepman, 1891 (sinónimo

junior), RMNH 86858, Japón, 40,0 mm; 18, 19: KF 3200, Filipinas, 81,9 mm; 20, 21: Paratipo 1

de Fusinus valdiviae Hadorn y Fraussen, 1999 (sinónimo junior), KF 2811, Somalia, 47,4 mm.

Figuras 22, 23. Fusinus (Chryseofusus) hyphalus M. Smith, 1940, RH, Taiwan, 50,3 mm.

213

Iberus, 21 (1), 2003

Islands, KARUBAR stn CP16, 05? 17' S, 132? 50' E, 315-349 m, 1 dd juv; stn CP27, 05* 33' S, 132*

51 E, 304-314 m, 2 dd juv.

Philippines, northwest of Mindanao, Aliguay Island, 150 m, 1 dd, Snyder coll.

Vanuatu, MUSORSTOM 8 stn DW958, 20? 21' S, 169* 47' E, 497-570 m, 1 lv; stn DW1105, 15* 03

S, 167? 07' E, 154-179 m, 1 dd; stn CP1124, 15* 02' S, 166" 57' E, 532-599 m, 1 dd.

North New Caledonia, BATHUS 4 stn CP948, 20* 33' S, 164" 57' E, 533-610 m, 1 dd; stn CP950,

20* 32 S, 164" 56' E, 705-750 m, 1 dd.

South New Caledonia, SMIB 1 stn DW, 22* 52! S, 167* 13 E, 415 m, 1 dd.

New Caledonia, Loyality Ridge, CALSUB pl. 11, 20? 52” 5” S, 167” 03' E, 577 m, 1 dd; MUSORSTOM

6 stan DW483, 21” 20' S, 167” 48' E, 600 m, 1 dd juv.

Southwest Pacific, Tuscarora Bank, MUSORSTOM 7 stn DW555, 11* 47' S, 178* 1% W, 540-542 m, 1 dd.

Fiji, Bligh Water, MUSORSTOM 10 stn DW1314, 17” 16.1' S, 178” 14.8' E, 656-660 m, 1 lv juv; stn

CP1327, 17 13.3' S, 177” 51.6' E, 370-389 m, 1 fragment; stn CP1330, 17* 09.5' S, 177* 56.3' E, 567-

699 m, 2 1v/2 lv juv; stn CP1341, 16? 52.5' S, 177 43.7" E, 500-614 m, 1 lv/1 dd.

Fiji, BORDAU 1 stn CP1401, 16* 35' S, 179” 41” W, 600-648 m, 4 lv juv/2 dd juv; stn CP1407, 16" 40"

S, 179” 39 E, 499-527 m, 1 lv juv; stn CP1448, 16* 45' S, 179? 59 E, 410-500 m, 1 lv juv; stn DW1451,

16" 45' S, 179? 59.5' E, 400-460 m, 2 lv juv; stn CP1502, 18* 21' S, 178” 27” W, 640-660 m, 1 dd.

Tonga Islands, BORDAU 2 stn DW1538, 21” 39 S, 175” 19 W, 471-508 m, 1 dd juv; south Nomuka

group, sta DW1554, 20” 38' S, 174” 58” W, 482-498 m, 1 dd juv; Chenal north Nomuka, stn DW1555,

20” 11'S, 174? 45 W, 591 m, 1 dd juv; “Seamount”, sta DW1605, 22* 17' S, 175” 16' W, 441 m, 2 frag-

ments; stn CP1620, 24” 18' S, 176” 20” W, 572 m, 1 dd; northwest Tongatapu, stn CP1640, 21” 09 5,

175? 24” W, 564-569 m, 1 lv juv; stn CP1642, 21* 05' S, 175” 23" W, 532 m, 1 dd juv.

Description: Shell medium-sized

(60.0-85.0 mm), uniformly off-white to

brownish, fusiform, with rather long

conical spire, solid, about 10 convex

unkeeled whorls, slightly concave

below suture, appressed to the preced-

ing whorl. Suture not deep, wavy on

upper whorls following the axial sculp-

ture, and straight on latter whorls.

Upper teleoconch whorls with rela-

tively strong axial ribs, separated by

interspaces of same width. 10-14 axial

ribs on uppermost whorl, 9-11 on the

following 3-5 whorls, disappearing on

latter whorls.

Spiral sculpture fine. 4 or 5 primary

spiral cords on uppermost teleoconch

whorls; 1 fine intercalated thread on

fourth whorl becoming as strong as the

primary ones on following whorls. Fine

tertiary threads appear from sixth or

seventh whorl on; the number increases

to 3 or 4 of unequal strength on body

whorl. Spiral sculpture crossed by

curved, strong, well-visible and close-

set growth lines, giving surface the

texture of linen.

Aperture ovate, pinched at both

ends, white. Parietal callus thin, smooth

and glossy, extended and adherent to

parietal wall. Columellar folds absent.

Outer lip thin, simple, smooth inside.

214

Siphonal canal relatively short,

about as long as aperture, curved. Outer

side sculptured with fine spirals,

crossed by growth lines.

Operculum typical of genus, cor-

neous, brown, shape and size corre-

sponding to aperture, nucleus terminal.

Periostracum brown.

Radula (Figs. 76, 86) typically fusinid.

Central tooth with tricuspid base, the

median cusp longest, base broader than

the notched anterior margin. Lateral

teeth broad, strongly curved, with 6-9

cusps of subequal length, at both ends

with a small denticle.

Range and habitat: Indonesia, Philip-

pines, northern Madagascar, and south-

western Pacific (New Caledonia,

Vanuatu, Fiji and Tonga) between 179

and 797 m deep.

Remarks: F. chrysodomoides was

placed in the genus Siphonofusus (Buc-

cinidae) by authors, but the radula (Figs.

76, 86) and conchological characteristics

are typical of the genus Fusinus. The

Australian shells figured as Siphonofusus

chrysodomoides by WILSON (1994: 66, pl.

12, figs. 7a-b) or as Fusinus

chrysodomoides by HADORN AND

FRAUSSEN (1999: pl. 3, figs. 17-18) are

clearly different from the Indonesian

type material. The Australian species is

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

described as F. westralis sp. nov. later on

in this paper. F. westralis differs from F.

chrysodomoides by the clearly larger shell

size, the more slender and more ex-

tended ire, the larger number of

whorls, by the more concave shoulder

slope especially on the lower whorls,

the narrower interspaces between the

axial ribs on upper whorls, and finally

by the finer spiral sculpture and the

larger number of very fine intercalated

spiral threads.

F. chrysodomoides from New Caledo-

nia usually have more prominent and

broader axial ribs on upper whorls. All

specimens from Fiji and Tonga and most

specimens from New Caledonia are

brownish coloured as opposed to the

white coloured type material of F.

chrysodomoides from Indonesia, the white

coloured specimens from the East

African coast and one specimen from

New Caledonia (CALSUB 1989 pl. 11).

However, we found no evidence to sep-

arate the white and the brownish

coloured specimens. Moreover, the

radulae of the New Caledonian speci-

mens are nearly identical to the figured

radula of the paralectotype of HF

chrysodomoides (SCHEPMAN, 1911: Fig. 10).

BOUCHET AND WARÉN (1986: 475)

noted, that Manaria insularis Okutani,

1968 “resembles closely Fusus chryso-

domoides Schepman, 1911”. After exami-

nation of the holotype of Manaria insu-

laris [NSMT Mo. 60220; Type locality:

Japan, off Hachijo Island, R/V Soyo-

Maru stn B3, 33” 08.3” N, 140” 01.8” E,

460 m] and direct comparison with the

type material of F. chrysodomoides we

found out, that both species are clearly

distinct. F. chrysodomoides clearly has a

larger adult size, a smaller spire angle

(-35* -38? /M. insularis: -42* ), a propor-

tionally longer spire and a narrower

siphonal canal. The axial ribs are usually

restricted to the spire tip, are more

prominent and separated by wider in-

terspaces. The spiral sculpture on upper

whorls clearly consists of less distinct

spiral cords and is also present in the in-

terspaces between the axial ribs. The

growth lines are prominent and well-

visible on all whorls, a characteristic of

all Chryseofusus. M. insularis only has in-

conspicuous growth lines. F. chryso-

domoides has a smooth columella as op-

posed to M. insularis, which has one

weak columellar fold.

No genus nor family (Buccinidae or

Fasciolariidae) can be ascertained to

accomodate M. insularis, until more

material with animal and radula, is

found.

Fusinus (Chryseofusus) graciliformis (Sowerby, 1880) (Figs. 14-21, 87)

Fusus graciliformis Sowerby, 1880. Thes. Conch., Monogr. Fusus, 4.: 80, pl. VIL fig. 62.

SYNONYMS

Fusus sieboldi Schepman, 1891. Notes Leyden Mus., 13 (2): 62; 1891: Notes Leyden Mus., 13 (17): 157,

pl. 9, figs. 3a, b.

Fusinus valdiviae Hadorn and Fraussen, 1999. Vita Marina, 46 (3-4): 117-120, figs. 11-16, text fig. 2.

SUBSEQUENT USE

Fusinus graciliformis (Sowerby, 1880). Fujira (1929: 90, pl. 3, fig. 11).

Simplicifusus graciliformis (Sowerby, 1880). KURODA AND HABE in Kuroda, Habe and Oyama

(1971: 282-283, pl. 51, fig. 2).

Type material: F. graciliformis: Holotype BMNH 1880.10.15.2 (52.5 x 18.0 mm, dd) - F. sieboldi:

Holotype RMNH 86858 (40.0 x 15.2 mm, dd). — F. valdiviae: Holotype MNHN (50.2 x 18.9 mm, lv

subad); 11 paratypes, Somalia, deep water, KF, RH and B. Briano (42.5-62.9 mm, dd subad); 1

paratype BMNH 1998187, Somalia, deep water (48.0 x 17.1 mm, dd subad); 1 paratype NMBE

1727.99, Somalia, deep water (45.7 x 17.0 mm, dd subad); 1 paratype IMT-99-2, Somalia, deep

water (44.9 x 18.3 mm, dd subad); 3 paratypes, Gulf of Aden, Somalia, deep water, H. Dekker

(49.0-60.5 mm, dd subad); 1 paratype ZMB, VALDIVIA stn 256, 01? 49 N, 45? 29 E, 1134 m (47.2

x 18.3 mm, dd subad).

Iberus, 21 (1), 2003

Figures 24-27. Fusinus (Chryseofusus) subangulatus (von Martens, 1901). 24, 25: Lectotype ZMB 59931,

south Somalia, near Kismaayo, 71.1 mm; 26, 27: RH, Somalia, 55.3 mm. Figures 28-31. Fusinus

(Chryseofusus) bradnerí (Drivas and Jay, 1990). 28, 29: MNHN, Reunion, off Saint-Gilles, 53.5 mm;

30, 31: MNAHN, Glorieuses Islands, dwarf form?, 48.4 mm. Figures 32, 33. Fusinus (Chryseofusus)

kazdailisí Eraussen and Hadorn, 2000, Holotype KMMA LJM925, Chile, Nazca Ridge, 40.0 mm.

Figuras 24-27. Fusinus (Chryseofusus) subangulatus (von Martens, 1901). 24, 25: Lectotipo ZMB 59931,

S de Somalia, cerca de Kismaayo, 71,1 mm; 26, 27: RH, Somalia, 55,3 mm. Figuras 28-31. Fusinus

(Chryseofusus) bradneri (Drivas y Jay 1990). 28, 29: MNHN, Reunion, frente a Saint-Gilles, 53,5

mm; 30, 31: MNAN, Glorieuses Islands, ¿forma enana?, 48,4 mm. Figuras 32, 33. Fusinus (Chryseo-

fusus) kazdailisi Fraussen y Hadorn, 2000, Holotipo KMMA LJM925, Chile, Nazca Ridge, 40.0 mm.

216

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Type locality: F. graciliformis: Japan. F. sieboldi: Japan. F. valdiviae: Somalia, between Ras Hafun and

Djibouti, approximately 400 m deep.

Material examined: The holotypes of F. graciliformis and F. sieboldi, and all the type material of F.

valdiviae.

Madagascar, southwest Madagascar, Tulear, 500-800 m, collected by commercial boats, 4 dd, B.

Briano; 1 dd, RH. — West Madagascar, off Morondava, 600 m, collected by commercial boats, 3 dd,

KF; 2 dd, RH. — North Madagascar, 12” 52' 0” S, 48* 10' 3” E, 420-428 m, 1 dd juv; 12” 42 9” S, 48?

12' 1” E, 445-455 m, 1 dd juv.

Philippines, south Bohol, Balicasag Island, off Panglao, 380 m, 1 lv, KF 2819. — Bohol, Balicasag, 275

m, 1 lv, C. Takahashi; 1 lv, B. Rogers. — North Mindanao, north of Dipolog, Aliguay Island, deep

water, 3 lv, KF 3200; 2 lv, RH.

Description: Medium-sized shell

(42.5-90.0 mm) with about 9-11 convex

whorls, surrounded by a weak subsu-

tural concavity. Suture appressed to pre-

ceding whorl, not incised.

Protoconch broken or eroded in all

examined specimens and therefore

unknown.

Eight or nine rather strong and

narrow axial ribs, reaching from suture to

suture and separated by narrow and not

very deep impressed interspaces on the 2

uppermost teleoconch whorls. On the fol-

lowing 2 or 3 whorls the number of ribs

increases up to 11-13 per whorl, becom-

ing somewhat broader and lower and

usually stopping below the upper suture;

interspaces between ribs become shal-

lower and less distinct. On antepenulti-

mate or penultimate whorl the axial ribs

become very low and less distinct and

the spacing becomes irregular. Axial ribs

always obsolete on body whorl and

usually also on penultimate whorl.

Teleoconch begins with 5 or 6 strong

but low and rounded spiral cords. Start-

ing with the second or third whorl an

additional clearly finer spiral cord

appears just below suture, and a fine

intercalated spiral thread appears

between each pair of cords. On the fol-

lowing whorls the number of fine spiral

threads increases by intercalation; up to

5 or 6 between each pair of primary

cords on body whorl. In the subsutural

concavity only some fine threads. The

spiral sculpture is crossed by rather

strong and well-visible curved growth

lines, giving the surface of the shell the

texture of linen.

Aperture ovate, whitish to yellowish

in colour, acute at posterior end. Parietal

callus thin, smooth and glossy, appressed

to lower part of body whorl. Columellar

folds absent. Outer lip simple and thin,

sculptured with numerous fine internal

lirae; outline of lip strikingly curved like

an elongated reversed “S'. Siphonal canal

shorter than aperture length, conspicu-

ously curved. Outer side ornamented

with numerous fine, low and regularly

spaced spiral cords which are interca-

lated by numerous fine spiral threads.

Operculum corneous, colour pale

reddish brown, thin, shape and size cor-

responding to aperture. Outer side with

fine concentric growth lines. Typical of

genus, with terminal nucleus.

Periostracum thin, olive-green, not

hairy.

Radula (Fig. 87) typical of genus.

Central tooth nearly rectangular, tricus-

pid, median cusp somewhat stronger.

Lateral teeth strongly curved, with 6 or

7 long, strong and pointed cusps; outer-

most one much larger than all others. At

both ends of lateral tooth with a small

denticle.

Range and. habitat: Japan (B0só penin-

sula and southwards), East China Sea,

Philippines, South China Sea (HIGO ET

AL., 1999: 263), and along the East

African coast, from the Gulf of Aden,

Djibouti, Somalia (HADORN AND

FRAUSSEN, 1999: 120) to southwestern

Madagascar. Between 50 and 1134 m

deep, on sandy bottom.

Remarks: Comparing both the holo-

types of F. graciliformis (Figs. 14-15) and

F. sieboldi (Figs. 16-17) shows, that the

latter is doubtless a junior synonym.

Both holotypes are nearly identical in

size, shape, sculpture and both were

found in Japan.

Iberus, 21 (1), 2003

F. valdiviae (Figs. 20-21) appears to be

conspecific, the description based on

subadult East African specimens. HADORN

AND FRAUSSEN (1999) compared F. valdiviae

with F. westralis (described in this paper)

misidentified in the 1999 paper as F.

chrysodomoides. After examination of the

holotype of EF. graciliformis and other mate-

rial mainly from the Philippines we treat

the conchologically not distinguishable F.

valdiviae from East Africa provisionally as

a junior synonym of EF. graciliformis.

Further study can reveal a subspecific

status for F. valdiviae. The radula of F. gra-

ciliformis is still unknown, the radula of F.

valdiviae was figured by HADORN AND

FRAUSSEN (1999, Fig. 87).

F. hyphalus differs from EF. gracili-

formis by having a more slender spire

tip, a smaller adult size, a thinner and

lighter shell, and by the less convex

whorls.

Fusinus (Chryseofusus) hyphalus M. Smith, 1940 (Figs. 22, 23)

Fusinus hyphalus M. Smith, 1940. The Nautilus, 54 (2): 43, pl. 2, fig. 9.

SUBSEQUENT USE.

Fusinus (Simplicifusus) hyphalus M. Smith, 1940. KIRA (1962: 85).

Simplicifusus hyphalus (M. Smith, 1940). HIGO ET AL. (1999: 263).

Type material: Holotype in M. Smith” s collection (36.5 x 12.5 mm) (not seen).

Type locality: Japan, off Tosa, 183 m deep.

Material examined: Taiwan, deep water, 1 lv juv/1 lv, RH. — Keelung, 1 lv/1 dd subad, RH.

Description: Shell entirely white or

pale, medium-sized (36.0-75.0 mm),

thin, lightweight, fusiform, elongate,

consisting of about 8 or 9 slightly

convex whorls with only slight subsu-

tural concavity. Body whorl often

inflated and ventricose in adult speci-

mens. Suture indistinct, straight.

Protoconch typical of genus, 1 !/2

whorls, bulbous, white, glossy, final

part (*/4 whorl) sculptured with 4 or 5

strong axial riblets, diameter 1.0-1.1

mm.

Weak, narrow and inconspicuous

axial ribs on upper 5 or 6 teleoconch

whorls, separated by narrow inter-

spaces, disappearing on latter whorls.

10 or 11 axial ribs on the 4 uppermost

teleoconch whorls extending from

suture to suture, 11-13 on fifth or sixth

whorl, becoming weaker and disappear-

ing on the following whorl.

Teleoconch beginning with 3 rela-

tively weak primary spiral cords, the 2

lower ones slightly stronger. From

second whorl on, a fine secondary spiral

thread appears between primary cords

and below suture, becoming as strong as

the primary cords on following whorls.

From fourth postnuclear whorl on, 2-4

218

fine intercalated tertiary threads appear

between primary and secondary cords.

Axial growth lines fine but distinct on

all whorls, crossing spiral sculpture and

giving the surface a pearled appearance.

Aperture ovate, whitish, relatively

large. Edge of outer lip simple, slightly

crenulated, internal side smooth or with

some weak internal lirae. Inner lip

smooth, glossy, parietal callus appressed

to parietal wall, columellar folds absent.

Siphonal canal thin, slightly curved, as

long as aperture, open.

Periostracum thin, persistent, straw-

brown in colour.

Operculum typical of genus, cor-

neous, brown, shape and size corre-

sponding to aperture, nucleus terminal.

Range and habitat: Japan, Enshu-nada

and westwards, East China Sea at 100-

300 m on sandy bottom (HIGO ET AL.,

1999: 263); Philippines, Cebu and Bohol

(SPRINGSTEEN AND LEOBRERA, 1986: 177).

Remarks: F. hyphalus resembles Sim-

plicifusus noguchii, but belongs to

Fusinus because of the typical fusinid

operculum. Simplicifusus noguchii differs

from E. hyphalus by having a smaller

adult size, a more slender white-

coloured shell, and by the presence of

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

axial ribs usually on all whorls, stronger

on upper whorls, becoming weaker on

body whorl, and by the small, nearly

round operculum, which is typical of

Simplicifusus (Fig. 2).

F. graciliformis can be distinguished

by the larger adult size, the longer spire,

the more convex whorls, and by often

having rather strong, broad axial ribs on

upper whorls.

Fusinus (Chryseofusus) subangulatus (von Martens, 1901) (Figs. 24-27, 77)

Fusus? subangulatus von Martens, 1901. Sitz.-Ber. Ges. Nat. Freunde Berlin, Jahrg. 1901: 21.

SUBSEQUENT USE

Fusus subangulatus von Martens. VON MARTENS (1904: 102-103, pl. 2, fig. 11).

Fusinus subangulatus (von Martens, 1903). HADORN AND FRAUSSEN (1999: 112-117, pl. 1, figs. 1-8,

text fig. 1, non pl. 2, figs. 9-10 =Fusinus jurgeni Hadorn and Fraussen, 2002).

Type material: Lectotype ZMB 59931, VALDIVIA (71.1 x 23.7 mm, dd), designated by HADORN

AND FRAUSSEN (1999). — 3 paralectotypes ZMB, south Somalia, near Kismaayo, VALDIVIA stn

253, 0? 27' S, 42? 47' E, 638 m (71.3 x 25.4 mm, lv); south Somalia, near Mogadisho, VALDIVIA

stn 256, 1” 49 N, 45” 29 E, 1134 m (37.1 mm, spire fragment, dd); Tanzania, Pemba Channel,

VALDIVIA stn 246, 5” 24” S, 39” 19 E, 818 m (39.9 x 14.5 mm, incomplete specimen with heavy

incrustations, dd).

Type locality: South Somalia, near Kismaayo, VALDIVIA stn 254, 0? 29 S, 42? 47' E, 977 m.

Material examined: Lectotype and the 3 paralectotypes in ZMB.

Somalia, deep water, 3 dd, KE; 1 dd, RH. — Between Ras Hafun and Djibouti, 400 m, 1 dd, KF; 1

dd, RH.

Description: Shell rather large (up to

77.2 mm), relatively heavy in weight.

Background colour whitish to yellowish

with pale reddish coloured spiral sculp-

ture. 9 or 10 convex whorls, latter

whorls distinctly keeled, shoulder slope

clearly concave. Axial and spiral sculp-

ture rather weak. Aperture relatively

large, siphonal canal short and distinctly

curved. Protoconch broken in all known

specimens. Suture appressed to the pre-

ceding whorl, wavy on upper whorls,

straight on penultimate and body

whorl.

Upper postnuclear whorls slightly

convex, sculptured with 9 or 10 narrow

and rather weak axial ribs, reaching

from suture to suture; interspaces

between them narrow and only weakly

impressed. Beginning at the appearance

of the peripheral keel (usually on fourth

or fifth teleoconch whorl) the inter-

spaces between the axial ribs become

wider; the ribs terminate below the

upper suture and are reduced to broad,

rounded knobs which are most promi-

nent at the periphery. On the penulti-

mate whorl they become weaker and

irregular, fading out suddenly. About 9-

13 axial knobs on latter whorls, but

usually obsolete on body whorl and

sometimes also on penultimate whorl.

Four strong spiral cords on the first

teleoconch whorl, 5 or 6 on the follow-

ing whorls. On latter whorls, beginning

with the peripheral keel, several rather

fine additional spiral threads appear on

shoulder. Starting with third or fourth

postnuclear whorl a fine spiral thread

appears between each pair of stronger

cords. The number of fine threads

increases by intercalation; up to 8 fine

intercalated threads of unequal strength

on body whorl. Central spiral cord and

the 2 cords below it somewhat stronger

than all others. Spiral sculpture crossed

by conspicuously strong growth lines,

well-visible on all whorls.

Aperture rather large, narrowly

ovate, acute at posterior end, yellowish

or white in colour. Outer lip simple and

sharp, without or with only weak inter-

nal lirae. Inner lip appressed to parietal

wall, smooth. Columellar folds absent.

Siphonal canal rather short for genus,

shorter than aperture length, slightly

Iberus, 21 (1), 2003

curved, widely open. Outer side orna-

mented with fine spiral cords and

numerous intercalated threads.

Operculum rather thin, colour pale

reddish brown, corneous. Shape and

size corresponding to aperture, typical

of genus, with terminal nucleus.

Radula (Fig. 77) typical of the genus,

consisting of an elongated central tooth

with a tricuspid base. Median cusp

clearly longest. Rounded base broader

than notched top. Lateral teeth conspic-

uously broad, strongly curved with 7-8

rather short, strong and pointed cusps; 2

outermost cusps broader and stronger.

Range and habitat: East Africa, from

Djibouti, Gulf of Aden to the Pemba

Channel, Tanzania, 400-1134 m deep on

blue silt and pteropod ooze.

Remarks: Rediscovered by HADORN

AND FRAUSSEN (1999), the typically

fusinid radula was figured for the first

time (Fig. 77). A specimen from off the

Somalian coast figured on pl. 2, figs. 9-

10 as F. subangulatus “broad form” is

now referred to F. jurgeni Hadorn and

Fraussen, 2002.

F. jurgeni has a larger adult size and

has a broader shell, a longer siphonal

canal, a larger number of spiral cords

which are clearly finer and denser.

F. chrysodomotides is most similar to F.

subangulatus and can be distinguished

by having a somewhat larger adult size,

uncoloured spiral cords, a larger

number of weaker spiral cords, often

broader and somewhat stronger axial

ribs, a more ventricose body whorl,

stronger axial growth lines, usually a

more curved siphonal canal and finally

some minor differences in radula mor-

phology.

Fusinus (Chryseofusus) bradneri (Drivas and Jay, 1990) (Figs. 28-31, 78)

Siphonofusus bradneri Drivas and Jay, 1990. Venus, 49 (4): 272-273, pl. 1, fig. 6.

SUBSEQUENT USE

Peristernia caledonica (Petit, 1851). DRIVAS AND JAY (1998: 39, sp. 27).

Fusinus bradneri (Drivas and Jay, 1990). FRAUSSEN (1999: 81); HADORN AND FRAUSSEN (1999: 117, 120).

Type material: Holotype in MNHN (54.0 x 18.0 mm, dd). — 4 paratypes (46.6-77.5 mm, dd), from

the type locality, in the collections of Jean Drivas and Maurice Jay.

Type locality: West coast of Reunion Island, St. Paul's Bay, 750 m.

Material examined: Holotype in MNHN.

Comoro Islands, Mayotte, east Passe Longogori, BENTHEDI 1977 stn 33, 12? 53' 5” S, 45” 16' 3” E,

275-400 m, 1 dd.

Glorieuses, west Grande Glorieuse, BENTHEDI 1977 stn 98, 11* 35' 5” S, 47" 16' 4” E, 280-460 m, 1

dd (dwarf form).

North Madagascar, 12? 42' 4” S, 48” 14 1” E, 375-380 m, 1 lv.

Reunion, Baie de la Possession, 600 m, 1 lv/2 dd. — Off Saint-Gilles, 700 m, 3 dd. — Reunion, Cam-

pagne MD32 stn CA7O, 21” 23' 01” S, 55* 29 03” E, 700-730 m, 1 dd; stn CP129, 20? 51' S, 55* 36' E,

290-300 m, 1 dd juv; stn DR62, 21? 09 S, 55* 12” E, 630-710 m, 1 dd juv; stn DC134, 20? 51' S, 55” 39

E, 650-750 m, 1 dd juv; stn CP144, 20? 50' S, 55? 35' E, 605-620 m, 1 lv juv. — Reunion, west coast, St.

Paul, 750-800 m, 1 dd, KF 2746; 5 dd, RH. — Reunion, off St. Paul's Bay, 750-800 m, 1 lv, KF.

Description: Shell fusiform, medium

sized (46.6-77.5 mm), solid, consisting of

7 or 8 convex teleoconch whorls with

subsutural concavity. Suture indistinct,

appressed to the preceding whorl, wavy

on axially ribbed part of spire, straight

on latter whorls. Reddish-brown

coloured, the stronger spirals and below

the suture darker.

Protoconch pale, typically fusinid.

220

About 7 strong, conspicuously broad

axial ribs separated by wide interspaces

on upper whorls, disappearing on

penultimate whorl.

Four strong primary spiral cords on

upper teleoconch whorls; from fourth

whorl on an intercalated secondary fine

spiral thread between primary cords,

soon becoming as strong as primary ones

on antepenultimate whorl. At the same

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

time 1-3 clearly weaker tertiary interca-

lated threads appear on body whorl.

Fine, but well-visible growth lines

intersect the spiral sculpture giving the

surface a finely granulate appearance.

Aperture whitish, relatively small,

ovate, pinched at both ends. Outer lip

relatively thick in adult specimens,

weakly denticulate, sculptured with

numerous close-set and distinct internal

lirae. Columellar callus smooth, adher-

ent. Columellar folds absent.

Siphonal canal long, rather narrow,

slightly curved.

Operculum typical of genus, cor-

neous, reddish-brown, shape and size

corresponding to aperture, nucleus ter-

minal.

Radula (Fig. 78) typical of genus. Tri-

cuspid elongate central tooth. Cusps

strong, long and pointed, projecting

below the base, central one somewhat

stronger. The rounded top is narrower

than the base. Lateral teeth have 5

strong pointed cusps with incurved tips.

Outermost one clearly the strongest.

With a small denticle at both ends.

Range and habitat: Reunion (300-750

m deep), north Madagascar (375-380 m),

Glorieuses (280-460 m) and Comoro

Islands (275-400 m). Previously only

known from Reunion. Live collected

specimens between 380 and 750 m deep,

dead shells between 300 and 750 m.

Remarks: Described as a Siphonofusus

(Buccinidae), but referred to Fusinus by

HADORN AND FRAUSSEN (1999) on the

basis of conchological resemblance to F.

valdiviae (a junior synonym of F. gracili-

formis), F. westralis sp. nov. (misidenti-

fied as “F. chrysodomoides”) and F. suban-

gulatus. The generic position was still

not confirmed because of the unknown

radula. A juvenile specimen of EF. brad-

neri, stored in MNHN, had the dried

animal preserved inside the shell. The

radula (Fig. 78) is found to be typical of

the genus Fusinus and the placement in

Fasciolariidae is therefore correct.

F. chrysodomoides most closely resem-

bles F. bradneri, but differs in having a

slightly larger number of axial ribs on

upper teleoconch whorls, in the nar-

rower axial ribs and the narrower inter-

spaces, the weak or absent lirae inside

the aperture, in the uniformly whitish or

greyish colour of the shell, in the usually

shorter and less twisted siphonal canal,

and finally in different radula morph-

ology (more elongated and narrower

central tooth, smaller number of cusps

on lateral teeth).

F. graciliformis differs by having a

somewhat larger and thinner shell, a

larger number of narrower axial ribs on

upper whorls with narrower inter-

spaces, an aperture with weaker and

indistinct internal lirae, and a uniformly

white or rarely flesh coloured shell.

One probably dwarf specimen (Figs.

30-31) from the Glorieuses Islands

(BENTHEDI stn 98) clearly has a shorter

shell, a shorter spire and a shorter

siphonal canal, a less constricted suture

and less convex whorls. More material

and study is necessary to conclude if

this is a dwarf form or a distinct species.

DRIVAS AND JAY (1998: 39, sp. 27)

figured a specimen of Chryseofusus

misidentified as Peristernia caledonica

(Petit, 1851). We examined two syntypes

of Turbinella caledonica Petit, 1851 in

MNHN [Type locality: New Caledonia]

and found out, that the identification

made by Drivas and Jay could defini-

tively not be correct. We consider the

figured shell a juvenile F. bradneri,

because of the close resemblance to the

juvenile material collected during the

Campagne MD32 1982 near Reunion.

Fusinus (Chryseofusus) jurgeni Hadorn and Fraussen, 2002 (Figs. 34-37, 81)

Fusinus subangulatus (von Martens, 1903) “broad form”. HADORN AND FRAUSSEN (1999: 115, pl. 2,

figs. 9-10; 116).

Fusinus cf. bradneri Drivas and Jay, 1990. MALLARD (2001: 12, fig. 12).

Fusinus jurgeni Hadorn and Fraussen, 2002. Iberus, 20 (1): 67-76, figs. 9-14, 20 (radula).

221

Iberus, 21 (1), 2003

Type material: Holotype (94.2 x 31.7 mm, lv) and one paratype (83.4 x 30.1 mm, dd) in MNHN, 5

paratypes in NM L5693/T1875 (89.7 x 32.6 mm, dd), KF (84.0 x 30.4 mm, dd), RH (89.1 x 31.0 mm,

76.5 x 26.6 mm, both dd), B. Rogers (90.5 x 32.2 mm, dd).

Type locality: Southwest Madagascar, Mozambique channel, 22? 22” S, 43? 03' E, 530 m.

Material examined: The live-taken holotype, collected by R. von Cosel during a shrimp stock sur-

vey by the commercial trawler Mascareignes III in 1986.

Madagascar, southwest Madagascar, Tulear, 500-800 m, collected by commercial boats, 5 paratypes

MNHN, NM L5693/T1875, KF, RH; 4 dd, B. Briano; 1 dd, RH. — Southwest Madagascar, off Tulear,

deep water, 1 dd, paratype B. Rogers. — West Madagascar, off Morondava, 600 m, 3 dd, KF 3208.

Somalia, trawled between Ras Hafun and Djibouti, 400 m, 1 dd, KF 1631.

Description: Shell large (up to 100.1

mm), fusiform, light in weight, spire

elongate, siphonal canal relatively long

for subgenus. 11 or 12 convex whorls,

latter whorls often slightly keeled,

shoulder slope concave. Shell uniformly

white, sometimes with reddish-brown

tinged spiral cords and with some weak

reddish brown axial strikes.

Protoconch typically fusinid, consist-

ing of 1 to 1 */4 smooth, glossy whorls.

Transition to teleoconch eroded and details

not recognizable. Diameter 0.9-1.0 mm.

Eight or nine rather narrow and

weak axial ribs on 3 uppermost teleo-

conch whorls, extending from suture to

suture. 8-10 axial ribs on following

whorls, withdrawing from upper su-

ture. Axial sculpture weak or absent on

penultimate whorl. Body whorl without

axial sculpture. Interspaces narrow on

upper whorls, slightly broader and less

pronounced on latter whorls.

Spiral sculpture crossed by strong,

curved axial growth lines. 3 strong

spiral cords on first teleoconch whorl,

the uppermost weaker. From second

whorl on, a fine intercalated secondary

spiral thread appears. From fifth or sixth

whorl on, secondary spirals becoming

as strong as primary ones and addi-

tional fine tertiary spiral threads appear.

On latter whorls 3-5 intercalated fine

threads visible between stronger spirals.

Aperture rather large, ovate, upper

end pointed, white coloured. Outer lip

simple and smooth, without internal

lirae. Inner lip completely smooth,

curved. Parietal callus thin, parietal wall

covered with an extended adherent

layer of callus. Columellar folds absent.

Siphonal canal almost straight or

slightly curved, as long as aperture.

222

Operculum typically fusinid, cor-

neous, Ovate, pointed below, dark

brown, shape and size corresponding to

aperture, with terminal nucleus. Outer

side ornamented with strong concentric

growth lines; inner side with a strongly

callused edge along the outer side.

Periostracum unknown.

Radula (Fig. 81) typical of genus.

Central tooth almost rectangular in shape;

base slightly convex, top straight or slightly

concave, both sides concave, tricuspid,

with strong, short, pointed cusps project-

ing below base. Lateral teeth elongate,

curved, with 7 strong, long, pointed cusps.

At both ends with a small denticle.

Range and habitat: Known from

southwestern Madagascar, between 500

and 800 m deep. One live collected

specimen 530 m deep. One record from

Somalia, collected between Ras Hafun

and Djibouti, 400 m deep (KF 1631).

Remarks: E. subangulatus has a some-

what smaller and heavier shell, a

smaller number of whorls, a more

slender shape, a shorter siphonal canal

and a stronger spiral sculpture with a

smaller number of spiral cords.

F. chrysodomoides differs in being

usually smaller, in having a heavier and

stouter shell, a less constricted suture, a

sometimes brownish coloured aperture

and a uniformly coloured shell, and a

spiral sculpture with a more beaded

appearance.

F. bradneri differs by its smaller size,

the more solid and reddish brown

coloured shell, the less constricted suture,

the broad and strong axial ribs on upper

whorls, the strong close-set lirae inside

the aperture, and by having a smaller

number of intercalated fine threads

between the stronger spiral cords.

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Fusinus (Chryseofusus) kazdailisi Fraussen and Hadorn, 2000 (Figs. 32-33, 79)

Fusinus kazdailisi Fraussen and Hadorn, 2000. Novapex, 1 (1): 15-19, figs. 1-12.

Type material: Holotype in KMMA LJM925 (40.0 x 17.0 mm, lv). - Chile, Nazca Ridge, Ecliptic

Bank, 110-150 m (56.0 x 23.0 mm, dd, paratype 1, H. Danila; 57.0 x 23.0 mm, dd, paratype 2, KF; 48.0

x 20.0 mm, dd, paratype 3, RH; 43.0 x 18.0 mm, lv, paratype 4, MNHN). — Chile, Nazca Ridge,

Mesyatsev Bank, 260-280 m (45.0 x 19.0 mm, lv, paratype 5, KF). — Chile, off Valparaiso, 1200 m

(52.0 x 22.0 mm, lv, paratype 6, A. Nora; 51.0 x 20.0 mm, lv, paratype 7, KF). — unknown locality

(47.0 x 19.0 mm, lv, paratype 8, C. and J. Hemmen; 57.0 x 22.0 mm, lv, paratype 9, Haus der Natur,

Cismar, Germany; 48.0 x 20.0 mm, lv, paratype 10, Haus der Natur, Cismar, Germany).

Type locality: Off Chile, Nazca Ridge, Ecliptic Bank, in deep water.

Material examined: Holotype in KMMA and all paratypes.

Chile, Nazca Ridge, Ecliptic Bank, 110-150 m, 1 dd, KF; 1 lv, RH.

Unknown locality, 1 dd, RH; 1 dd, KF 2944; 1 dd, R. Kelly.

Description: Shell medium sized

(40.0-57.0 mm), solid, shape fusiform,

semi-slender. Whorls about 6 in number,

showing a weak subsutural concavity.

Suture appressed to preceding whorl.

Shell dirty greyish to brown or pale

reddish-brown. Protoconch missing in

all known specimens.

Eight to fourteen rather strong and

narrow axial ribs, traversing from suture

to suture, on upper whorls. Interspaces

rather narrow. Axial ribs become weaker

on following whorls and finally disappear.

All whorls densely covered with fine,

sometimes slightly curved growth lines,

often not visible on their own but recog-

nizable by low papillae formed on spiral

cords, giving the surface of latter whorls

a pearled appearance.

Seven to nine strong and rounded

primary spiral cords on first remaining

teleoconch whorl, separated by deep and

narrow interspaces. On second whorl,

interspaces become broader, tending to be

similar in size to spiral cords. 9 or 10

primary spiral cords on following whorl,

12-14 on penultimate whorl. On third or

fourth whorl 1 fine intercalated secondary

spiral thread appears between each pair

of primary cords. Furthermore, 1 or 2 addi-

tional intercalated tertiary spiral threads

between secondary and primary cords on

lower part of penultimate whorl.

Aperture narrowly ovate to lens-

shaped. Parietal callus thin, smooth and

glossy, appressed to body whorl. Col-

umellar folds absent. Outer lip thin.

Siphonal canal broad, shorter than aper-

ture, oblique at left-hand side and

straight.

Operculum corneous, thin and pale

reddish-brown, shape and size corre-

sponding to aperture, with terminal

nucleus.

Periostracum thin, olive-green in

colour.

Radula (Fig. 79) typical of genus.

Central tooth tricuspid, median cusp

strongest, occasionally with 1 or 2 small

additional denticles situated near

middle of cusp. Lateral teeth strongly

curved, with 6 or 7 long and pointed

cusps mostly of equal size. A small den-

ticle occasionally appears at both ends

of lateral tooth.

Range and habitat: Described from

Nazca Ridge and from off Valparaiso,

Chile. On mud and muddy sand in deep

water between 150 and 1200 m.

Remarks: Aeneator castilla: McLean

and Andrade, 1982, a buccinid species,

is similar in sculpture and colour, but

can be distinguished by the larger size,

the broader shape, the more ventricose

body whorl, the clearly larger aperture,

and finally the buccinid radula.

Fusinus (Chryseofusus) acherusius sp. nov. (Figs. 38-43, 80)

Type material: Holotype (58.6 x 22.6 mm, lv) and one paratype (48.0 x 20.9 mm, lv) in MNHN, 1

paratype RH (52.3 x 21.4 mm, dd).

Iberus, 21 (1), 2003

Type locality: West Madagascar, Mozambique Channel, stn CH126, 17” 50' S, 43? 07' E, 1475-1530 m.

Material examined: The live taken holotype and the paratype in MNHN from the type locality.

Madagascar, west Madagascar, Mozambique Channel, 18? 00' S, 43? 00' E, 1715-1750 m, 1 dd,

paratype RH. — Northwest Madagascar, Mozambique Channel, stn CH131, 13? 46/ S, 47? 33' E, 1490-

1600 m, 1 lv.

South New Caledonia, BIOCAL stn CP27, 23" 06' S, 166" 26' E, 1850-1900 m, 1 dd.

Etymology: “acherusius” (Latin, adjective), meaning “from the underworld” and “sombre, dreary”.

Description: Shell medium sized (up

to 58.6 mm), dirty grey, rather heavy,

body whorl short, spire high, consisting

of about 8 carinated whorls which are

appressed to preceding one. Shoulder

slope straight or slightly concave.

Protoconch decollated in all avail-

able specimens.

Upper teleoconch whorls with about

12 rather weak axial ribs, which are

most prominent at periphery. Inter-

spaces shallow, as broad as ribs. Ribs

slightly more numerous (up to 17) on

latter whorls, withdrawing from upper

suture and becoming simultaneously

weaker, gradually fading away towards

body whorl. Usually without axial ribs

on body whorl.

Four or five strong spiral cords on

upper teleoconch whorls. Central one

somewhat stronger, forming a distinct

carina on all whorls. From early whorls

on, 1 fine intercalated secondary thread

appears between each pair of stronger

primary cords. On body whorl often

becoming as strong as primary ones.

Spiral sculpture crossed by numerous

unequal close-set axial growth lines

giving surface the texture of linen.

Aperture elongate, narrow, white to

yellowish. Outer lip sharp, simple,

finely crenulated. Posterior canal well

developed, without internal lirae. Inner

lip smooth, without callus and columel-

lar folds. Siphonal canal short, slightly

longer than half of aperture length,

widely open, strongly curved to the left

and slightly curved backwards.

Periostracum thin, light brownish,

well-adherent.

Operculum typical of genus, brown

coloured, corneous, size and shape cor-

responding to aperture, with terminal

nucleus.

Radula (Fig. 80) fusinid, central

tooth tricuspid, somewhat atypical for

Fusinus, extremely narrow, elongated,

with a strong central cusp and 1 rudi-

mentary, reduced, hardly visible small

cusp at both sides of the central one.

Lateral teeth typical of genus, slightly

curved, base broad, with 7 rather short

pointed cusps.

Range and habitat: West Madagascar,

Mozambique Channel, 1530-1715 m

deep. One single record (Figs. 42, 43)

from south New Caledonia, 1850-1900

m deep.

Comparison: F. kazdailisi from off the

Chilean coast most closely resembles F.

acherusius, but can be separated by

having weaker and more numerous

axial ribs on upper teleoconch whorls,

no axial ribs on penultimate whorl, non-

carinated whorls, a narrower and

straight siphonal canal, and finally a dif-

ferent radula (central tooth nearly round

with 3 well-developed cusps, the central

one occasionally with 1 or 2 small addi-

tional denticles).

Fusinus (Chryseofusus) artutus sp. nov. (Figs. 44-47, 82)

Type material: Holotype NM L2083 (72.2 x 26.0 mm, dd) in NM, paratype 1 (59.5 x 23.2 mm, lv),

B. Rogers.

Type locality: Philippines, Bohol, Panglao, 200 m deep.

Material examined: Holotype and paratype both from the type locality, collected by local fishermen.

Indonesia, Tanimbar Islands, KARUBAR stn CP46, 08* 01' S, 132? 51' E, 271-273 m, 1 lv subad.

New Caledonia, BIOCAL stn DWO08, 20* 34' S, 166" 54' E, 435 m, 1 lv/2 dd juv.

Etymology: Named after the Latin expression “artutus” (adjective), meaning “strongly built” (or

“sturdy”, “solid”, “firm”), remembering the broad and solid shell.

224

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Figures 34-37. Fusinus (Chryseofusus) jurgeni Hadorn and Fraussen, 2002. 34, 35: Holotype MNHN,

southwest Madagascar, 94.2 mm; 36, 37: Paratype MNHN, southwest Madagascar, Tulear, 83.4 mm.

Figures 38-43. Fusinus (Chryseofusus) acherusius sp. nov. 38, 39: Holotype MNHN, west Madagas-

car, Mozambique Channel, 58.6 mm; 40, 41: Paratype MNHN, west Madagascar, Mozambique

Channel, 48.0 mm; 42, 43: MNHN, south New Caledonia, 35.5 mm.

Figuras 34-37. Fusinus (Chryseofusus) jurgeni Hadorn y Fraussen, 2002. 34, 35: Holotipo MNHN, SE

de Madagascar, 94,2 mm; 36, 37: Paratipo MNHN, SO de Madagascar, Tulear, 83,4 mm. Figuras 38-

43. Fusinus (Chryseofusus) acherusius sp. nov. 38, 39: Holotipo MNHN, O de Madagascar, Mozambi-

que Channel, 58,6 mm; 40, 41: Paratipo MNHN, O de Madagascar, Mozambique Channel, 48,0 mm);

42, 43: MNHN, S de Nueva Caledonia, 35,5 mm.

Iberus, 21 (1), 2003

Description: Shell up to 72.2 mm,

fusiform, stout, with broad spire angle, 9

or 10 convex, shouldered whorls with

slight subsutural concavity. Suture wavy,

following axial sculpture of preceding

whorl on ribbed upper part of the spire,

straight on lower whorls. Spire tip white,

latter whorls flesh coloured, inside aper-

ture light brownish or whitish.

Protoconch rather large, white, 1 to 1

1/4 whorls, smooth and glossy, some-

times with axial growth lines on final

part, ending in a varix. 1.0 mm in diam-

eter.

Strong, close-set axial ribs on upper

teleoconch whorls, traversing from

suture to suture. Interspaces narrow and

deep. About 9-11 narrow and distinct

ribs on upper whorls, suddenly becom-

ing lower and broader, and fading away

on penultimate whorl.

Teleoconch begins with 4 strong

spiral cords. 2 additional cords appear

on second whorl: one immediately

below suture, the other just above lower

suture. From third whorl on, a fine sec-

ondary spiral thread appears between

each pair of primary cords, becoming

soon as strong as primary ones. In addi-

tion very fine intercalated tertiary

spirals appear. This fine spiral sculpture

is crossed by strong close-set axial

growth lines, giving the surface the

texture of linen.

Aperture ovate, white or light

brownish coloured. Outer lip thin, edge

slightly crenulated, inside smooth or

inconspicuously lirate. Parietal callus

thin, smooth, columellar folds absent.

Siphonal canal rather short, about as

long as aperture, curved, slightly

bending backwards.

Periostracum thin, greenish, well-

adherent.

Operculum typically fusinid,

reddish-brown, corneous, shape and

size corresponding to aperture, with ter-

minal nucleus.

Radula typically fusinid (Fig. 82).

Central tooth small, broad, top and both

sides concave, base convex, tricuspid,

cusps of about equal size, projecting

below base. Lateral teeth curved, having

6 or 7 rather small, short, pointed cusps

with incurved tips. Small denticle at

inner end.

Range and habitat: Philippines, Bohol,

200-540 m deep. One record from

Indonesia, Tanimbar Islands, 271-273 m

deep, and one record from New Caledo-

nia, 435 m deep.

Comparison: F. chrysodomoides has a

less constricted suture, a white to

brownish shell, coarser and less numer-

ous axial ribs on upper whorls, and a

finer, more close-set spiral sculpture.

F. graciliformis -has a more elegant

shell with elongate spire, a larger adult

size, less ventricose, less convex and

longer whorls, and has a weaker spiral

sculpture with a larger number of inter-

calated fine threads on body whorl.

Fusinus (Chryseofusus) dapsilis sp. nov. (Figs. 48, 49)

Type material: Holotype (74.1 x 29.2 mm, dd) in MNHN.

Type locality: Vietnam, deep water, not accompanied by detailed data.

Material examined: The holotype is the only specimen known, and originates from material

traded by Russian collectors in the late 1980s — early 1990s.

Etymology: Named after the Latin expression “dapsilis” (adjective), meaning “precious”

“expensive”, remembering the golden colour.

Description: Shell 74.1 mm, reddish-

brown, fusiform, thin, light in weight

but solid. Whorls convex and ventri-

cose, surrounded by distinct subsutural

concavity.

Protoconch and spire tip broken

leaving 6 */4 remaining whorls. Original

226

number of teleoconch whorls 9 or 10 by

estimation.

Upper part of spire covered with 10 or

11 rounded, dense, relatively strong axial

ribs, extending from suture to suture, sud-

denly fading away on antepenultimate

whorl. Last 3 whorls smooth, except for

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

strong, curved axial growth lines crossed

by weak close-set spiral threads, giving

surface the texture of linen.

Four rather strong spiral cords on

uppermost remaining whorl; subsutural

cord somewhat weaker. On next whorl,

a fine intercalated secondary thread

appears between each pair of primary

cords, becoming as strong as primary

ones on antepenultimate whorl. Up to 3

tertiary, clearly finer intercalated threads

appear between the strong primary and

secondary cords on penultimate and

body whorl. Spiral sculpture crossed by

strong growth lines, giving the spirals a

pearled appearance.

Aperture rather large, light purplish

coloured. Outer lip simple, thin, edge

minutely crenulated; interior side

smooth. Parietal wall covered with a

thin, glossy and smooth callus, some-

what extending outside aperture. Col-

umellar folds absent. Siphonal canal

conspicuously short, broad, open and

strongly curved.

Periostracum,

radula unknown.

Range and habitat: Only known from

the South China Sea, off Vietnam. No

precise information about locality and

habitat available.

Comparison: E. chrysodomoides differs

from F. dapsilis by the heavier shell, the

broader and less close-set axial ribs and

the longer siphonal canal.

FE bradneri has a smaller adult size, is

more elegant and has a more elongate

spire, a longer and narrower siphonal

canal, and the inside of the aperture is

strongly lirate. Moreover, the axial

sculpture on upper teleoconch whorls

consists of strong, broad axial ribs, and

the stronger spiral cords are reddish-

brown in colour.

protoconch and

Fusinus (Chryseofusus) riscus sp. nov. (Figs. 50-53, 83)

Type material: Holotype (19.1 x 8.3 mm, lv subad) and one paratype (18.9 x 8.9 mm, lv) in MNHN,

4 paratypes in AMS C.205150 (21.5 x 9.7 mm, dd), NMNZ M.273192 (19.4 x 8.4 mm, dd), RH (21.4

x 9.3 mm, lv subad), KF (19.3 x 8.3 mm, lv subad).

Type locality: South New Caledonia, Norfolk Ridge, BATHUS 3 stn DW818, 23” 44' S, 168” 16' E,

394-401 m. |

Material examined: The live collected holotype, the 5 paratypes, and 8 specimens (2 dd /3 lv subad /3

dd juv) from the type locality.

South New Caledonia, Norfolk Ridge, BATHUS 3 stn DW817, 23* 42 S, 168” 16' E, 405-410 m, 11

dd/5 dd juv/3 lv/2 lv juv. — South New Caledonia, BERYX 11 stn CP21, 24? 44' S, 168” 07' E, 430-

450 m, 2 dd.

Etymology: The name is derived from the Latin “riscus” (noun, masculine), meaning a rotan box.

F. riscus resembles a rotan box in surface sculpture (fine spiral sculpture crossed by strong growth

lines giving surface the texture of linen).

Description: Shell small for genus (up

to 21.5 mm), fusiform, thick, whitish to

yellowish, consisting of 7 or 8 convex,

slightly carinated whorls, appressed to

preceding whorl, surrounded by a dis-

tinct subsutural concavity. Suture wavy,

inconspicuous.

Protoconch consisting of 1 whorl,

white, transparent, glossy, smooth, final

part (about */4 whorl) with some very

fine axial growth lines and 2 or 3 weak

spiral threads; transition to teleoconch

indistinct. 0.6-0.8 mm in diameter.

Axial sculpture consists of broad, low

knobs on all whorls, separated by wide

and low interspaces. On upper teleo-

conch whorls 7 or 8 ribs traversing from

suture to suture, on latter whorls 5 or 6

shorter ribs abapically prominent, often

somewhat irregular on body whorl.

Teleoconch begins with 4 spiral cords,

abapical ones clearly stronger. On fourth

or fifth teleoconch whorl, a fifth strong

spiral cord appears just above lower

suture, stronger when crossing axial

knobs and weak in interspaces. Number

of spirals increasing by intercalation on

latter whorls, secondary ones becoming

as strong as primary ones on penultimate

and body whorl. On body whorl fine ter-

227

Iberus, 21 (1), 2003

Figures 44-47. Fusinus (Chryseofusus) artutus sp. nov. 44, 45: Holotype NM L2083, Philippines, Bohol,

Panglao, 72.2 mm; 46, 47: Paratype B. Rogers, Philippines, Bohol, Panglao, 59.5 mm. Figures 48,

49. Fusinus (Chryseofusus) dapsilis sp. nov., Holotype MNHN, Vietnam, 74.1 mm. Figures 50-53.

Fusinus (Chryseofusus) riscus sp. nov. 50, 51: Holotype MNHN, New Caledonia, Norfolk Ridge, 19.1

mm; 52, 53: MNHN, New Caledonia, Norfolk Ridge, 24.8 mm.

Figuras 44-47. Fusinus (Chryseofusus) artutus sp. nov. 44, 45: Holotipo NM L2083, Filipinas, Bohol,

Panglao, 72,2 mm; 46, 47: Paratipo B. Rogers, Filipinas, Bohol, Panglao, 59,5 mm. Figuras 48, 49. Fusinus

(Chryseofusus) dapsilis sp. n0v., Holotipo MNHN, Vietnam, 74,1 mm. Figuras 50-53. FEusinus (Chry-

seofusus) riscus sp. nov. 50, 51: Holotipo MNHN, Nueva Caledonia, Norfolk Ridge, 19,1 mm; 52, 53:

MNHN, Nueva Caledoniz, Norfolk Ridge, 24,8 mm.

228

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Figures 54-57. Fusinus (Chryseofusus) cadus sp. nov. 54, 55: Holotype MNHN, south New Caledonia,

35.5 mm; 56, 57: MNHN, south New Caledonia, 30.5 mm. Figures 58-61. Fusinus (Chryseofusus)

alisae sp. nov. 58, 59: Holotype MNHN, north New Caledonia, 29.6 mm; 60, 61: Paratype MNHN,

New Caledonia, 27.5 mm. Figures 62-65. Fusinus (Chryseofusus) scissus sp. nov. 62, 63: Holotype

MNHN, south New Caledonia, 33.0 mm; 64, 65: Paratype MNHN, south New Caledonia, 31.1 mm.

Figuras 54-57. Fusinus (Chryseofusus) cadus sp. nov. 54, 55: Holotipo MNHN, S de Nueva Caledo-

nia, 35,5 mm; 56, 57: MNHAN, S de Nueva Caledonia, 30,5 mm. Figuras 58-61. Fusinus (Chryseo-

fusus) alisae sp. nov. 58, 59: Holotipo MNHN, N de Nueva Caledonia, 29,6 mm; 60, 61: Paratipo MNHN,

Nueva Caledonia, 27,5 mm. Figuras 62-65. Fusinus (Chryseofusus) scissus sp. nov. 62, 63: Holotipo

MNHAN, S de Nueva Caledonia, 33,0 mm; 64, 65: Paratipo MNHN, S de Nueva Caledonia, 31,1 mm.

229

Iberus, 21 (1), 2003

tiary threads appear at both sides of sec-

ondary spirals. Spiral sculpture crossed

by conspicuously strong growth lines

giving surface the texture of linen.

Aperture small, ovate, white

coloured. Outer lip slightly crenulated,

simple. Conspicuously strong, broad

internal lirae. Parietal callus appressed,

smooth, columellar folds absent.

Siphonal canal shorter than aperture

length, straight, widely opened. Perios-

tracum light brownish, rather thick.

Operculum brownish, typical of

genus, shape and size corresponding to

aperture, with terminal nucleus.

Radula typical of Fusinus (Fig. 83).

Central tooth large, elongate, almost

oblong with slightly concave sides, tri-

cuspid. All 3 cusps conspicuously

strong and pointed, clearly projecting

below base. Central cusp the strongest.

Lateral teeth curved, 4-6 strong, long

and pointed cusps with incurved tips.

At both ends with a small pointed denti-

cle.

Range and habitat: Southern New

Caledonia, between 401 and 430 m

deep.

Comparison: F. cadus and F. alisae,

both described later in this paper, have a

larger adult size, clearly narrower and

more axial ribs on upper whorls, and

usually a ribless penultimate and body

whorl.

Fusinus (Chryseofusus) cadus sp. nov. (Figs. 54-57, 84)

Type material: Holotype (35.5 x 13.4 mm, dd) and 1 paratype (22.3 x 8.9 mm, lv juv) in MNHN, 4

paratypes in AMS C.205151 (26.9 x 10.3 mm, dd subad), NMNZ M.273193 (25.7 x 11.0 mm, dd), RH

(23.9 x 9.8 mm, lv subad), KF (19.8 x 8.0 mm, dd juv).

Type locality: South New Caledonia, BERYX 11 stn DW27, 23* 37' S, 167” 41' E, 460-470 m.

Material examined: The holotype, 1 paratype MNHN and 6 specimens (dd) from the type local-

ity, all in MNHN. |

North New Caledonia, MUSORSTOM 4 stn DW162, 18? 35' S, 163” 10' E, 525 m, 1 dd juv. - BATHUS

4 stn DW908, 18* 58' S, 163 11' E, 502-527 m, 1 dd.

South New Caledonia, BERYX 11 stn DW38, 23* 38' S, 167* 39 E, 550-690 m, 5 dd/1 lv subad/1 dd

juv; stn DW39, 23" 37' S, 167” 40' E, 490-500 m, 2 dd juv. - SMIB 3 stn DW 12, 23” 38' S, 167" 42 E,

470 m, 2 dd. - BIOCAL stn DW33, 23” 10' S, 167" 10' E, 675-680 m, 1 lv juv/2 dd juv; stn CP52, 23

06' S, 167” 47" E, 540-600 m, 1 dd subad, paratype AMS C.205151. - CHALCAL 2 stn DW76, 23? 41"

S, 167” 45' E, 470 m, 2 dd juv/1 lv subad, paratypes RH and KF. - MUSORSTOM 4 stn DW221, 22?

59 S, 167” 37' E, 535-560 m, 1 dd juv; stn DW229, 22* 51' S, 167” 13' E, 445-460 m, 1 dd, paratype

NMNZ M.273193.

Etymology: The name is derived from the Latin “cadus” (noun, masculine), meaning a small ter-

racotta bottle. F. cadus, small and with subsutural concavity, also resembles pottery in colour.

Description: Shell small (up to 35.5 mm),

whitish to light brownish, consisting of 8

whorls. Whorls convex, with slight sub-

sutural concavity. Suture indistinct.

Protoconch porcellaneous white,

swollen, smooth and glossy, consisting

of 1 whorl. Final part (about */4 whorl)

sometimes with some fine axial riblets.

0.7-0.9 mm in diameter.

On upper teleoconch whorls, 8 or 9

strong, broad, low axial ribs, traversing

from suture to suture, somewhat nar-

rower at upper end. Interspaces half as

wide as the axial ribs. Axial sculpture

suddenly fading out on antepenultimate

or penultimate whorl.

230

Four strong spiral cords on upper teleo-

conch whorls. From third whorl on, a fine

spiral thread appears between primary

ones. On fifth whorl, up to 3 fine interca-

lated tertiary spirals appear and secondary

spirals become as broad as the primary

ones. Spiral sculpture weak, crossed by

numerous well-visible close-set growth

lines giving surface the texture of linen.

Aperture narrowly ovate, white,

pointed at both ends, with numerous

weak internal lirae. Outer lip thin, finely

crenulated. Parietal callus thin, smooth,

appressed, columellar folds absent.

Siphonal canal slightly curved, some-

what shorter than aperture.

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Periostracum light brownish, thick.

Operculum typical of genus, cor-

neous, brown, shape and size corre-

sponding to aperture, with terminal

nucleus.

Radula typically fusinid (Fig. 84),

central tooth rather large, almost ovate

(top and base convex, both sides straight

or slightly concave), with 3 prominent

cusps projecting below base. Central

cusp somewhat stronger than the others.

Lateral teeth curved, with 6 prominent

long cusps with incurved tips. Outer-

most one the broadest and strongest.

With a small denticle at both ends.

Range and habitat: From north New

Caledonia to south New Caledonia (18?

aa 163” 10E to 23-41" 5, 167” 47" E),

between 460 and 675 m deep.

Comparison: F. riscus has a smaller

adult size, less numerous and much

broader axial ribs with broad inter-

spaces on upper whorls, and has the

penultimate and body whorl axially

ribbed.

F. alisae is similar in size and sculp-

ture, but has a larger number of nar-

rower axial ribs, more convex whorls,

the upper whorls increase their diame-

ter much more rapidly, and different

radula morphology (lateral teeth with

more and clearly shorter cusps, and the

central tooth has a narrower, concave

top and more concave sides and oblique

cusps at both sides of the central one).

Juvenile specimens of F. wareni are

also similar in shape and sculpture, but

the whorl diameter increases more

rapidly and the spire is less extended. F.

wareni has more axial ribs per whorl,

adult shells are clearly larger in size and

have more whorls.

Fusinus (Chryseofusus) alisae sp. nov. (Figs. 58-61, 85)

Type material: Holotype (29.6 x 12.1 mm, dd) and 1 paratype (27.5 x 11.8 mm, dd) in MNHN, 4

paratypes in AMS C.205152 (26.4 x 11.0 mm, dd), NMNZ M.273194 (27.4 x 12.5 mm, dd), RH (35.5

x 15.5 mm, dd), KF (20.8 x 9.6 mm, dd). |

Type locality: North New Caledonia, BATHUS 4 stn DW927, 18* 56' S, 163” 22 E, 444-452 m.

Material examined: The holotype, 1 paratype MNHN and 14 specimens (9 dd juv/5 lv juv) from

the type locality.

Coral Sea, Chesterfield Islands, MUSORSTOM 5 stn 361, 19* 53' S, 158? 38' E, 400 m, 3 dd juv; stn

378, 19” 54” S, 158” 38' E, 355 m, 2 dd juv; stn 379, 19* 53' S, 158” 40' E, 370-400 m, 1 dd juv.

North New Caledonia, SMIB 6 stn DW118, 18* 58' S, 163” 26' E, 290-300 m, 1 dd, paratype RH; stn

Dw119, 18* 59 S, 163" 26' E, 295-305 m, 1 dd, paratype NMNZ M.273194; stn DW121, 18? 58' S,

163" 26' E, 315 m, 1 lv/1 dd/1 dd juv; stn DW122, 18* 58' S, 163” 25' E, 325-330 m, 2 lv subad; stn

DWw123, 18” 57' S, 163” 25' E, 330-360 m, 1 dd, paratype AMS C.205152. —- BATHUS 4 stn DW923,

18” 52 S, 163” 24” E, 470-502 m, 3 dd juv; stan DW924, 18* 55' S, 163” 24” E, 344-360 m, 3 dd; stn

DW925, 18* 55 S, 163” 24” E, 370-405 m, 4 dd juv; stan DW926, 18? 57' S, 163” 25' E, 325-330 m, 1

dd /2 dd juv; stn CP928, 18* 55' S, 1637 24” E, 420-452 m, 1 dd/1 dd juv; stn DW931, 18* 55' S, 163”

24” E, 360-377 m, 1 lv juv/3 dd; stn DW940, 19* 00' S, 163* 26' E, 305 m, 1 dd, paratype KF. —

MUSORSTOM 4 stn CP194, 18* 53' S, 163” 22” E, 545 m, 3 dd juv; stan DW181, 18? 57' S, 163” 22 E,

350 m, 4 dd juv; sta DW196, 18* 55' S, 163” 24' E, 450 m, 2 dd juv.

Etymology: Named after the French research vessel “Alis”, which collected most of the known

specimens during the cruises BATHUS 4 and SMIB 6.

Description: Shell small (up to 35.5

mm), whitish to light brownish,

fusiform, 7 or 8 convex whorls with sub-

sutural concavity. Upper whorls slightly

bicarinate. Diameter of upper whorls

increasing more rapidly than lower

ones.

Protoconch whitish, consisting of 1

1/4 whorls. Upper part smooth and

glossy, last part (about */4 to */2 whorl)

sculptured with some narrow but strong

axial riblets. 0.7-1.0 mm in diameter.

Axial ribs traversing from suture to

suture on upper whorls, withdrawing

from upper suture on latter whorls. 9-12

rather weak, narrow ribs on 2 upper-

most teleoconch whorls, separated by

conspicuously narrow interspaces. 10-14

231

Iberus, 21 (1), 2003

Figures 66, 67. Fusinus (Chryseofusus) scissus sp. nov., MNHN, south New Caledonia, 32.1 mm. Figures

68-71. Fusinus (Chryseofusus) wareni sp. nov. 68, 69: Holotype MNHN, New Caledonia, 59.5 mm;

70,71: Paratype MNHN, New Caledonia, 50.0 mm. Figures 72-75. Fusinus (Chryseofusus) westralis

sp. nov., northwest Australia, Rottnest Island. 72, 73: Holotype WAM S10876, 114.4 mm; 74, 75:

Paratype MNHN, 113.3 mm.

Figuras 66, 67. Fusinus (Chryseofusus) scissus sp. 20%., MNHN, S de Nueva Caledonia, 32,1 mm. Figuras

68-71. Fusinus (Chryseofusus) wareni sp. nov. 68, 69: Holotipo MNHN, Nueva Caledonia, 59,5 mm;

70, 71: Paratipo MNHN, Nueva Caledonia, 50,0 mm. Figuras 72-75. Fusinus (Chryseofusus) wes-

tralis sp. nov., NO de Australia, Rottnest Island. 72, 73: Holotipo WAM S10876, 114,4 mm; 74, 75:

Paratipo MNHN, 113.3 mm.

232

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

ribs on following 2 or 3 whorls, latter

whorls usually without ribs. Large adult

specimens sometimes with conspicu-

ously strong axial knobs on body whorl.

Teleoconch begins with 4 spiral

cords. 2 adapical ones clearly stronger

from beginning and forming a weak,

inconspicuous double keel on upper

whorls, fading away on latter whorls.

From second or third postnuclear whorl

on, a fine intercalated thread appears

between each pair of primary cords.

Number of fine threads increasing up to

2 by intercalation on latter whorls.

Rather weak spiral sculpture crossed by

prominent axial growth lines giving

surface the texture of linen.

Aperture ovate, whitish, pointed at

both ends. Outer lip slightly crenulated,

simple. About 18 rather fine internal lirae.

Inner lip smooth, parietal callus thin, col-

umellar folds absent. Siphonal canal

shorter than aperture, slightly curved.

Periostracum light brown, rather

thin.

Operculum corneous, brownish,

typical of genus, shape and size corre-

sponding to aperture, with terminal

nucleus.

Radula typical of genus (Fig. 85).

Central tooth small, short. Base convex,

broad, with 3 prominent cusps of about

equal size projecting below base; top

narrow, concave; both sides strongly

convex. Lateral teeth curved, having 6 or

7 short pointed cusps of about equal size

and a small pointed denticle at both ends.

Range and habitat: North of New

Caledonia and Coral Sea, Chesterfield

Islands, between 300 and 545 m deep.

Comparison: F. riscus differs in being

smaller, having less convex whorls, a

clearly smaller number of much broader

axial ribs with broader interspaces on

upper whorls, an axially ribbed penulti-

mate and body whorl and differences in

radula morphology (clearly larger, elon-

gate central tooth, a smaller number of

much longer cusps on lateral teeth) and

a different protoconch (only one whorl,

sculptured with fine axial growth lines

and 2 or 3 weak spiral threads on final

part).

F. cadus differs in the less numerous

and broader axial ribs, less convex

whorls, the upper whorls increasing in

diameter slower than in F. alisae, the last

part of the protoconch is smooth or has

some fine axial riblets, and has a differ-

ent radula morphology (lateral teeth

with less numerous but much longer

cusps).

Fusinus (Chryseofusus) scissus sp. nov. (Figs. 62-67, 88)

Type material: Holotype (33.0 x 14.5 mm, lv) and 2 paratypes (31.1 x 13.6 mm, lv; 33.4 x 15.9 mm,

dd, siphonal canal broken) in MNHN, and 4 paratypes in AMS C.205153 (31.9 x 14.7 mm, lv), NMNZ

M.273195 (29.9 x 13.8 mm, lv), RH (33.4 x 14.1 mm, lv), KF (27.2 x 11.4 mm, lv).

Type locality: South New Caledonia, SMIB 3 stn DW24, 22* 59 S, 167? 21' E, 535 m.

Material examined: The live taken holotype and all the paratypes from the type locality.

South New Caledonia, SMIB 1 stn DWZ, 22* 52 S, 167” 13' E, 415 m, 1 lv/1 lv juv; stn DW7, 22* 56'

S, 167” 16' E, 500 m, 1 dd juv. —- SMIB 2 stn DW3, 22* 56' S, 167" 15' E, 412-428 m, 1 dd; stn DWA4,

22” 53' S, 167” 13' E, 410-417 m, 1 dd juv; stn DW5, 22* 56' S, 167” 14” E, 398-410 m, 2 dd juv; stn

DW6, 22? 56' S, 167” 16' E, 442-460 m, 1 dd juv; stn DW9, 22* 54' S, 167" 15' E, 475-500 m, 2 lv juv;

stn DW 10, 22* 55 S, 167” 16' E, 490-495 m, 3 lv/7 dd; stn DW17, 22* 55' S, 167" 15' E, 428-448 m, 1

lv; sn DW18b, 22* 58' S, 167” 20' E, 530-535 m, 3 lv/6 dd; stn DC26, 22* 59 S, 167” 23' E, 500-535

m, 1 lv/1 dd. —- SMIB 3 stn DW21, 22* 59 S, 167* 19 E, 525 m, 1 lv/1 dd; stn DW22, 23* 03 $, 167"

19 E, 503 m, 3 lv juv/1 dd; stn DW23, 22* 58' S, 167* 20' E, 530 m, 1 lv/1 dd; stn DW26, 22* 55' S,

167” 16' E, 450 m, 1 dd juv. — SMIB 4 stn DW58, 23" 00' S, 167" 24' E, 480-560 m, 2 1v/1 dd; stn DWél,

23” 00' S, 167” 22” E, 520-550 m, 1 lv/1 dd; stn DWé2, 23* 00' S, 167” 22' E, 490-540 m, 1 lv; stn DW63,

22” 59 S, 167” 21' E, 580 m, 1 dd; stn DW65, 22* 55' S, 167" 15' E, 400-420 m, 1 dd. - MUSORSTOM

4 stn CP214, 22” 54' S, 167” 14 E, 425-440 m, 1 dd; stn CP216, 22* 59 S, 167* 22' E, 490-515 m, 1 dd

juv; stn DW222, 22* 58' S, 167” 33' E, 410-440 m, 1 dd. - BERYX 11 stn CP22, 24” 44' S, 168* 07' E,

490-510 m, 1 dd. - BIOCAL stn DW44, 22* 47' S, 167" 14” E, 440-450 m, 2 dd /22 dd juv; stn DW46,

22” 53' S, 167” 17' E, 570-610 m, 20 dd juv/4 lv juv. - BATHUS 2 stn DW7109, 22” 48' S, 167" 16' E,

233

Iberus, 21 (1), 2003

444-445 m, 2 1v/1 dd/15 dd juv; stn DW720, 22* 52' S, 167” 16' E, 530-541 m, 3 lv/24 dd /9 dd juv;

stn DW721, 22? 54' S, 167” 17' E, 525-547 m, 3 lv/12 dd/26 dd juv?/6 lv juv.

Ile des Pins, southeast Récif sud, SMIB 8 stns DW193-196, 22” 59 5-23" 00' S, 168* 21' E-168* 23' E,

491-558 m, 16 dd/18 dd juv; stan DW201, 22” 59 S, 168” 21' E, 500-504 m, 2 dd/1 dd juv.

Norfolk Ridge, SMIB 8 stn DW200, 24” 00' S, 168” 21” E, 514-525 m, 2 dd.

New Caledonia, 500 m, by local fisherman, 2 lv/3 dd, RH.

Etymology: “scissus” (Latin, adjective) means “wrinkled”, describing the weak and close-set axial

folds on all whorls.

Description: Shell small (up to 33.4

mm), fusiform, body whorl inflated, 8

whorls. Whorls convex, unkeeled or

with an inconspicuous double keel,

body whorl conspicuously ventricose.

Colour uniformly white.

Protoconch large, 1.0 mm in diame-

ter, smooth, whitish, consisting of 1*/4

whorls; final part (about */4 whorl)

sculptured with some fine axial riblets

traversing from suture to suture. Ending

in an indistinct varix.

Eleven to thirteen inconspicuous,

weak, close-set axial folds on all whorls,

traversing from suture to suture on up-

per whorls. Ribs withdrawing from up-

per suture on body whorl, becoming

somewhat stronger, forming more or less

prominent axial knobs at periphery.

Teleoconch beginning with 4 or 5

rather weak spiral cords. From second or

third whorl on, fine intercalated secondary

spiral threads appear between primary

cords. Number increasing to 6 on latter

whorls. 2 primary cords at periphery of-

ten somewhat stronger than others, form-

ing sometimes an indistinct double keel.

Aperture large, ovate, pinched at upper

end, white or cream coloured, often light

brownish or purplish along edge on inner

side of outer lip. Outer lip simple, slightly

crenulated; inner side ornamented with

close-set white lirae. Parietal callus thin,

appressed to parietal wall, smooth and

glossy. Columellar folds absent. Siphonal

canal conspicuously short, broad, curved

and widely open. Outer side sculptured

with some rather weak spiral cords and

up to 3 intercalated fine threads.

Periostracum straw coloured, thin.

Operculum typical of genus, cor-

neo0us, light brown, shape and size cor-

responding to aperture, outer side

sculptured with fine concentric growth

lines, with terminal nucleus.

Radula typical of genus (Fig. 88).

Central tooth tricuspid, middle cusp

strongly developed, long and straight;

cusps at both sides shorter with

incurved tips. All cusps clearly project

below base. Lateral teeth broad, curved,

consisting of 5 or 6 strong, long, pointed

cusps with incurved tips and with small

denticle at both ends. Outermost cusp

strongest and broadest.

Range and habitat: Only known from

southern New Caledonia, 410-580 m

deep.

Comparison: F. scissus can be distin-

guished from the other small southwest

Pacific species F. cadus, F. alisae and EF.

riscus by the somewhat larger adult size,

the large, ventricose body whorl, the

larger aperture and the axially ribbed

body whorl.

Manaria insularis Okutani, 1968, a

species of uncertain generic position

(see discussion under F. chrysodomotides),

most closely resembles F. scissus in

shape and size, but differs in having a

light brown shell, a smaller number of

whorls, indistinct axial ribs on body

whorl, and by the clearly different spiral

sculpture (close-set, strong, raised spiral

cords, strong on the top of ribs, obscure

in the interspaces, and by the presence

of only one intercalated fine thread on

penultimate and body whorl).

Fusinus (Chryseofusus) wareni sp. nov. (Figs. 68-71)

Type material: Holotype (59.5 x 22.3 mm, dd) and 1 paratype in MNHN (50.0 x 20.1 mm, dad).

Type locality: New Caledonia, MUSORSTOM 4 stn CP170, 18* 57' S, 163" 13' E, 480 m.

234

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Material examined: The holotype in MNHN.

New Caledonia, MUSORSTOM 4 stn CC201, 18* 56' S, 163” 14” E, 490 m, 1 dd, paratype MNHN.

Loyality Ridge, BATHUS 3 stn DW794, 23” 48' S, 169” 49 E, 751-755 m, 1 fragment.

Tonga Islands, seamount south of Eua, BORDAU 2 stn DW1617, 23” 03' S, 175” 53” W, 483-531 m,

1 dd/1 dd juv.

Etymology: This species is named to honour Anders Warén (SMNH) for his contributions to mala-

cology. The preparation of the radulae and SEM illustrations in this paper are his work.

Description: Shell medium sized (up

to 59.5 mm), fusiform, uniformly whitish

or brownish, consisting of about 9

rounded teleoconch whorls with slight

subsutural concavity. Suture indistinct,

whorls appressed to preceding one.

Protoconch white, bulbous, glossy,

smooth, consisting of 1 */4whorls, ending

in a varix. Diameter about 0.9-1.0 mm.

Uppermost 4 or 5 teleoconch whorls

rather weakly axially ribbed. 10-12 low,

broad ribs with narrow interspaces,

reaching from suture to suture on first 3

whorls, withdrawing from upper suture

on fourth or fifth postnuclear whorl.

Lower whorls without axial ribs. All

whorls ornamented with well-visible,

close-set, curved growth lines, crossing

the weak spiral cords and giving surface

the texture of linen.

Teleoconch beginning with 4 or 5

spiral cords, adapically more prominent

compared to upper ones. From third

whorl on, a fine secondary spiral thread

appears between each pair of primary

cords. From fifth whorl on, 1 tertiary

fine thread appears at both sides of sec-

ondary ones. From sixth whorl on, the

secondary ones become as strong as

primary ones. 2 or 3 fine intercalated

spiral threads between stronger cords

on body whorl.

Aperture ovate, pointed at both

ends, white or flesh coloured, inner lip

smooth. Parietal wall covered with a

thin callus, becoming well-developed on

columella. Columellar folds absent.

Outer lip simple. Internal side sculp-

tured with about 18-20 elongate, irregu-

lar and rather strong denticles. Siphonal

canal about as long as aperture, strongly

curved, rather narrow, open, tapering

anteriorly. Outer side sculptured with

close-set spiral cords and up to 3 fine

intercalated spiral threads.

Periostracum, operculum and radula

unknown.

Range and habitat: New Caledonia

and Tonga Islands, between 480 and 751

m deep.

Comparison: FEF. kazdailisi is most

similar to F. wareni in shape and sculp-

ture but differs in having a somewhat

smaller adult size, a dirty greyish to

brown or pale reddish-brown colour,

less numerous whorls, stronger and

more pronounced axial ribs with more

distinct interspaces, a smaller number

of somewhat stronger spiral cords

which are intercalated by 3-6 fine

spiral threads on body whorl, and

finally in having a broad and straight

siphonal canal which is oblique to the

left side.

FE acherustus has a heavier shell, cari-

nated upper whorls, numerous pro-

nounced axial ribs on most whorls, a

rough spiral sculpture, rough growth

lines of unequal strength, and a broader

and shorter siphonal canal.

F. alisae has a smaller adult size, a

whitish to light brownish colour,

broader upper postnuclear whorls, a

usually bicarinate profile of the upper

whorls, a smaller protoconch, a subsu-

tural concavity which is often more

prominent, and an often larger number

of axially ribbed whorls.

F. cadus also differs in having a

clearly smaller adult size, a whitish to

light brownish colour, a smaller proto-

conch with a smaller diameter, in

having a less prominent subsutural con-

cavity, a more slender shell, and a larger

number of axially ribbed whorls.

F. artutus is similar in sculpture, but

has a larger adult size, more convex

whorls, a longer spire, a more con-

stricted suture, a deeper subsutural con-

cavity, and a longer siphonal canal.

Iberus, 21 (1), 2003

Fusinus (Chryseofusus) westralis sp. nov. (Figs. 72-75)

Siphonofusus chrysodomoides (Schepman, 1911). — KosuGE (1985: 59, pl. 23, fig. 7); WILSON (1994: 66,

pl. 12, figs. 7a-b).

Fusinus chrysodomoides (Schepman, 1911). - HADORN AND FRAUSSEN (1999: pl. 3, figs. 17-18).

Type material: Holotype (114.4 x 36.2 mm, lv) in WAM 510876. 8 paratypes in MNHN (113.3 x 40.6

mm, lv), AMS C.205154 (99.3 x 34.0 mm, lv), NMNZ M.2731% (86.6 x 31.6 mm, lv), USNM (81.2 x

29.3 mm, lv), ZMA Moll. 4.03.005 (93.6 x 32.1 mm, lv), KF 3201 (123.5 x 40.2 mm, lv), RH (123.6 x

42.6 mm, lv), B. Rogers (114.1 x 38.4 mm, lv).

Type locality: Northwest Australia, Rottnest Island, 400-500 m deep.

Material examined: The live collected holotype and 8 paratypes from the type locality.

Northwest Australia, off Rottnest Island, 400-500 m deep, collected by commercial fishing boats,

33 lv, KF 3203; 1 lv, RH. — Off Port Hedland, deep water, 1 lv, KF3201; 1 lv, KF3202; 5 lv, RH.

Etymology: Named after Western Australia, which is the species present range. Also to remember

Perotrochus westralis (Whitehead, 1987), a sympatric species.

Description: Shell large (up to 140

mm), light, thin, uniformly white, shape

fusiform. 11-13 ventricose, unkeeled

whorls, with wide subsutural concavity.

Protoconch white, glossy, consisting

of 1 whorl, ending in a weak varix.

Surface of protoconch slightly eroded

and therefore no sculpture visible. 0.9

mm in diameter.

Nine to twelve rather weak, narrow

axial ribs on 4 uppermost teleoconch

whorls, reaching from suture to suture on

first postnuclear whorl. Withdrawing from

upper suture on second whorl. On third

or fourth whorl ribs suddenly weaker,

gradually fading away on following

whorls. Growth lines curved, fine.

About 4 or 5 rather weak spiral

cords on uppermost whorl. From sec-

ond postnuclear whorl on, a fine inter-

calated secondary spiral thread appears

between each pair of primary cords. On

following whorls numerous fine and in-

conspicuous tertiary spiral threads ap-

pear. Lower whorls covered by con-

spicuously fine spiral sculpture, axial

sculpture obsolete.

Aperture large, oval, pointed above,

white. Parietal callus smooth and thin.

Outer lip thin, simple, without internal

lirae. Columellar folds absent. Siphonal

canal shorter than aperture, rather

broad, strongly curved and widely

opened.

(Right page) Figures 76-88. Radulae. 76: Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911),

reproduced from SCHEPMAN (1911: fig. 10). 77: Fusinus (Chryseofusus) subangulatus (von Martens,

1901); 78: Fusinus (Chryseofusus) bradneri (Drivas and Jay, 1990), juvenile specimen; 79: Fusinus (Chry-

seofusus) kazdailisi Fraussen and Hadorn, 2000; 80: Fusinus (Chryseofusus) acherusius sp. nov.; 81: Fusinus

(Chryseofusus) jurgeni Hadorn and Fraussen, 2002; 82: Fusinus (Chryseofusus) artutus sp. nov.; 83: Fusinus

(Chryseofusus) riscus sp. nov.; 84: Fusinus (Chryseofusus) cadus sp. nov.; 85: Fusinus (Chryseofusus) alisae

sp. nov.; 86: Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911); 87: Fusinus (Chryseofusus) gra-

ciliformis (Sowerby, 1880), Holotype Fusinus valdiviae Hadorn and Fraussen, 1999 (junior synonym);

88: Fusinus (Chryseofusus) scissus sp. nov.

(Página derecha) Figuras 76-88. Rádulas. 76: Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911),

reproducido de SCHEPMAN (1911: fig. 10); 77: Fusinus (Chryseofusus) subangulatus (von Martens,

1901); 78: Fusinus (Chryseofusus) bradneri (Drivas y Jay 1990), juvenil; 79: Fusinus (Chryseofusus)

kazdailisi Fraussen y Hadorn, 2000; 80: Fusinus (Chryseofusus) acherusius sp. nov.; 81: Eusinus

(Chryseofusus) jurgeni Hadorn y Fraussen, 2002; 82: Fusinus (Chryseofusus) artutus sp. nov.; 83: Fusinus

(Chryseofusus) riscus sp. nov.; 84: Fusinus (Chryseofusus) cadus sp. nov.; 85: Fusinus (Chryseofusus)

alisae sp. nov.; 86: Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911); 87: Fusinus (Chryseo-

fusus) graciliformis (Sowerby 1880), Holotipo Fusinus valdiviae Hadorn y Fraussen, 1999 (sinónimo

junior); 88: Fusinus (Chryseofusus) scissus sp. 20%.

236

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

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Iberus, 21 (1), 2003

Operculum typical of genus, corneous,

brown, ovate, pointed at lower end, shape

and size corresponding to aperture, with

terminal nucleus. Outer side ornamented

with numerous concentric growth lines.

Periostracum and radula unknown.

Range and habitat: Only known from

western Australia. WILSON (1994)

reported this species as Siphonofusus

chrysodomoides (Schepman, 1911) from

off Rowley Shoals to Rottnest Island

between 300-500 m deep.

Comparison: F. westralis has hitherto

been identified and offered to collectors

as “Siphonofusus chrysodomoides”. Authors

figured this species as S. chrysodomoides

(KoSsuGE, 1985; WILSON, 1994) and as

Fusinus chrysodomoides (HADORN AND

BIOGEOGRAPHICAL DISCUSSION

The distribution of the 16 species of

Chryseofusus probably partly reflects ac-

tual distribution patterns and sampling

efforts. One species is known from the

East Pacific (kazdailisi) and Western Aus-

tralia (westralis), and these two species

are not known from elsewhere. Five

species are known from East and South-

east Asia (chrysodomoides, graciliformis,

hyphalus, artutus and dapsilis), of which

two (hyphalus, dapsilis) are not known

from elsewhere. Six species are known

from East Africa and the southwest In-

dian Ocean (chrysodomoides, graciliformis,

subangulatus, bradneri, acherusius and jur-

geni), of which three (subangulatus, brad-

neri and jurgeni) are not known from

elsewhere. Finally, eight species are

known from the SW. Pacific

(chrysodomoides, acherusius, artutus,

riscus, cadus, alisae, scissus and wareni), of

which five (riscus, cadus, alisae, scissus,

wareni) are not known from elsewhere.

The highest diversity is encountered

in New Caledonia, where all eight

species known in the southwest Pacific

co-occur, of which four (riscus, cadus, ali-

sae and scissus) can be regarded as local.

In the North of New Caledonia, up to

three species (cadus, alisae, wareni) poten-

tially occupy the same bathymetric

horizon between 300 and 545 m

238

FRAUSSEN, 1999). Wilson noted the

generic allocation of this species provi-

sional. Hadorn and Fraussen placed this

species in Fusinus based on conchological

resemblance with F. valdiviae. After

studying the Indonesian type material of

Fusus chrysodomoides we describe the

western Australian shells as F. westralis.

F. westralis differs from FH.

chrysodomoides by the larger shell size, the

more slender and more extended spire tip,

the larger number of whorls, the more

concave shoulder slope especially on the

lower whorls, the narrower interspaces

between axial ribs on upper whorls, and

finally by the much finer spiral sculpture

and the much larger number of very fine

intercalated spiral threads.

(whereas chrysodomoides occupies

slightly deeper water), but none of them

occurs syntopically with another species.

In the South and on Norfolk Ridge, up to

four species (chrysodomoides, cadus, riscus,

scissus) potentially occupy the same

bathymetric horizon between 401 and

675 m, and there is a single occurrence

where two species have been taken

together in the same haul (SMIB 1 stn

DW2: chrysodomoides, scissus). The high

diversity in the New Caledonia region

thus apparently reflects high environ-

mental heterogenity as well as individ-

ual specific ecological preferences (Table

D. The data for the southwest Indian

Ocean are more scanty but tend to

support the same conclusion: up to four

(chrysodomoides, graciliformis, bradneri and

jurgeni) occupy the same bathymetric

horizon between 350 and 600 m in the

Mozambique Channel, but we have no

documented case of syntopy.

ACKNOWLEDGMENTS

P. Bouchet (MNHN), B. Richer de

Forges (IRD, Nouméa) and A. Warén

(SMND) collectively collected most of the

material on which this paper is based. V.

Héros (MNHN) arranged for the loan of

HADORN AND FRAUSSEN: Indo-Pacific radiation of Fusinus (Chryseofusus subgen. nov.)

Table I. Geographical and bathymetrical distribution of species of Chryseofusus in the New Caledo-

nia region.

Tabla 1 . Distribución geográfica y batimétrica de las especies de Chryseofusus en la región de Nueva

Caledonta.

Species Coral Sea North New Caledonia New Coledonia proper Norfolk Ridge Loyality Ridge

chrysodomoides 610-705 415 577-600

acherusius 1850-1900

artutus 435

cadus 525 460-675

alisae 355-400 300-545

TÍSCUS 401-430

SCISSus 410-580

wareni 480-490 151-755

additional MNHN material, A. Warén

prepared radulae and SEM illustrations,

and P. Bouchet revised an earlier draft of

the manuscript. We thank R. Moolenbeek

(ZMA), J. Goud (RMNDH)), E. Gittenberger

(RMNH), K. Way (BMNH), J. Pickering

(BMND), R. Kilburn (NM), H. Saito

(NSMT), M. Gosteli (NMBE) and M.

Glaubrecht (ZMB) for the loan of type

material, S. Kosuge (IMT) for Japanese

literature and translation, R. Houart

(Belgium) for comments, corrections and

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O Sociedad Española de Malacología —_—_—_——T— Iberus, 21 (1):241-272, 2003

The Cystiscidae (Caenogastropoda) from upper reef for-

mations of New Caledonia

Los Cystiscidae (Caenogastropoda) de las formaciones superiores

de coral de Nueva Caledonia

Franck BOYER*

Recibido el 16-1-2003. Aceptado el 14-111-2003

ABSTRACT

The species of Cystiscidae from the New Caledonian upper reef formations are studied on

the ground of recent intensive recolts managed by Paris Museum, mainly the ones from

Noumea 1992-93, from the Expedition Montrouzier 1993 and from the Expedition Lifou

2000. The taxonomic analysis is focused on the morphs sampled off New Caledonia

mainland (northern and southern areas). The marginellids collection of Bavay recently

deposited in Paris Museum is used for the revision of several taxa.

Twenty eight morphs are recognized, among which five are attributed to species previ-

ously described, namely Gibberula lifovana (Crosse, 1871), Cystiscus goubini (Bavay,

1922), C. montrouzieri [Bavay, 1922), C. bougei (Bavay, 1917) and Plesiocystiscus

tomlini (Bavay, 1917), the last one with much reserve. The syntypes of Gibberula pulchella

(Kiener, 1834) are pictured and the species is compared to a New Caledonian relative.

Eighteen species are described as new: Gibberula squamosa sp. nov., G. cincta sp. nov.,

Crithe caledonica sp. nov., C. gofasi sp. nov., Cystiscus viridis sp. nov., C. punctatus sp.

nov., C. aurantivs sp. nov., C. marshalli sp. nov., C. boucheti sp. nov., C. camelopardalis

sp. nov., C. minor sp. nov., C. pardus sp. nov., C. deltoides sp. nov., C. caeruleus sp.

nov., C. fricinctus sp. nov., C. pseudoaurantius sp. nov., C. cooverti sp. nov., Plesiocystis-

cus bavayi sp. nov.

Five morphs are recorded as potential new species and provisionally referred as Gib-

berula sp. aff. philippii [Monterosato, 1878) (morphs A, B and S), Cystiscus sp. 1 and Ple-

siocystiscus sp. aff. bavayi sp. nov.

Despite the high diversity reported about the group Crithe/Cystiscus in shallow waters of

New Caledonia mainland, this diversity is appreciated as remaining much underesti-

mated. This diversity cannot be considered as restricted to the New Caledonian waters

and equivalent intensive collecting efforts in other places from West Pacific might possibly

yield equivalent results.

The genera Crithe and Cystiscus show as belonging to a continuous complex of related

forms, but the taxonomic unification of the group waits for a better knowledge of its total

diversity, allowing to propose appropriate subdivisions.

On the ground of the concrete limits met with the taxonomic interpretation, it is suggested

that the managing of systematic observations in the field about micro-habitats and about

the variability of the soft parts would highly increase the scientific profitability of intensive

malacological samplings.

* 110 chemin du Marais du Souci, 93270, Sevran, France. e-mail: Franck.Boyer6Gwanadoo.fr

241

Iberus, 21 (1), 2003

RESUMEN

Las especies de Cystiscidae de la parte superior de las formaciones arrecifales de Nueva

Caledonia son estudiadas en base a las recientes recolecciones intensivas dirigidas por el

Museum de Paris, principalmente las de Noumea 1992-93, la Expedition Montrouzier

1993 y la Expedition Lifou 2000. El análisis taxonómico se centra principalmente en los

morfos obtenidos fuera de New Caledonia [areas del norte y sur). La colección de margi-

nélidos de Bavay, recientemente depositada en el Museum de Paris ha sido usada para la

revisión de varios taxones.

Se reconocen veintiocho morfos, entre los cuales cinco son atribuidos a especies previa-

mente descritas, como Gibberula lifovana (Crosse, 1871), Cystiscus goubini [Bavay,

1922), C. montrouzieri [Bavay, 1922), C. bougei (Bavay, 1917) y Plesiocystiscus tomlini

(Bavay, 1917), aunque este último con muchas reservas. Los sintipos de Gibberula pulche-

lla (Kiener, 1834) son mostrados y la especie es comparada con una similar de Nueva

Caledonian.

Diez y ocho especes son descritas como nuevas: Gibberula squamosa sp. nov., G. cincta

sp. nov., Crithe caledonica sp. nov., C. gofasi sp. nov., Cystiscus viridis sp. nov., C. punc-

tatus sp. nov., C. aurantivs sp. nov., C. marshalli sp. nov., C. boucheti sp. nov., C. camelo-

pardalis sp. nov., C. minor sp. nov., C. pardus sp. nov., C. deltoides sp. nov., C. caeru-

leus sp. nov., C. tricinctus sp. nov., C. pseudoaurantius sp. nov., C. cooverti sp. nov., Ple-

siocystiscus bavayi sp. nov.

Cinco morfos son presentados como potenciales especies nuevas y provisionalmente des-

critos como Gibberula sp. aff. philippii [Monterosato, 1878) (morphs A, B y S], Cystiscus

sp. 1 y Plesiocystiscus sp. aff. bavayi sp. nov.

A pesar de la alta diversidad citada para el grupo Crithe/Cystiscus principalmente en

aguas superficiales de Nueva Caledonia, probablemente esta diversidad esté todavía

infravalorada. Sin embargo, esta diversidad no puede ser considerada restringida a las

aguas de Nueva Caledonia ya que una recolección intensiva equivalente en otras zonas

del Pacífico occidental posiblemente podría dar similares resultados.

Los géneros Crithe y Cystiscus muestran como vienen a ser un complejo continuo de

formas relacionadas entre sí, pero la unificación taxonómica del grupo queda pendiente

de un mayor conocimiento de su completa diversidad, permitiendo así proponer adecua-

das subdivisiones.

Basándonos en los límites encontrados con las interpretaciones taxonómicas, se sugiere

que la realización sistemáticas de las observaciones de campo acerca de los micro-hábi-

tats y de la variabilidad de las partes blandas aumentaría el provecho científico de los

muestreos intensivos.

KEY WORDS: Cystiscidae, Gibberula, Crithe, Cystiscus, Plesiocystiscus, New Caledonia, reef formations, sublit-

toral zone, diversity, soft parts chromatism.

PALABRAS CLAVE: Cystiscidae, Gibberula, Crithe, Cystiscus, Plesiocystiscus, Nueva Caledonia, formaciones

arrecifales, zona sublitoral, diversidad, cromatismo de partes blandas.

INTRODUCTION

The malacologic prospecting of

New Caledonia began around 1850

with the recolts made by the R. P. Mon-

trouzier and was developed through

the contribution of cultured local collec-

tors, often in link with experimented

242

European students. In these conditions,

the malacological fauna from New

Caledonia had become at the beginning

of the XX' century one of the best

known, as far as tropical latitudes are

concerned.

BOYER: The Cystiscidae from upper reef formations of New Caledonia

After a period of lower activity

begun around 1910, this malacologic

prospecting was revived at a high level

since the seventies, first about landsnails

and littoral marine mollusks, soon about

deep bathyal levels, leading to the dis-

covery of a highly diversified fauna.

The boosting of this malacologic

prospecting of New Caledonia is mainly

due to the sustained initiatives of Paris

Museum (MNHN), and of the local

branch of IRD (ex-ORSTOM). Besides

the managing of regular campaigns of

sampling at bathyal levels, the organiza-

tion of two expeditions devoted to the

fauna from upper reef formations

yielded intensive recolts from Northern

New Caledonia (Expedition Mon-

trouzier 1993 to Touho, NE of New

Caledonia, and Koumac, NW of New

Caledonia) and from Lifou Island (Expe-

dition Lifou 2000). These important

recolts, complementary to that ones

made earlier around Noumea, are

stored now in Paris Museum and allow

the revision of the New Caledonian sub-

littoral mollusks on the ground of a rep-

resentative material.

This article is devoted to the study of

Cystiscidae (Caenogastropoda) from

upper sublittoral levels of the New

Caledonian mainland, among which the

genus Cystiscus displays an original

high diversity.

The marginelliform gastropods from

New Caledonia were not subject to a

revision work since the article of BAVAY

(1922) devoting a first section (p. 57-65)

to the “Marginelles de l'archipel calé-

donien” (including Loyalty Islands).

BAVaAY (1922) did recognize the occur-

rence of eight morphospecies of Cystisci-

dae, among which only three were

attributed to species originally described

from the New Caledonian archipelago.

On the ground of the bicolor banded

animals of two sibling species of Cystis-

cus respectively recorded from Mas-

carene Islands and from Society Islands,

and of the original radula observed in

the first one, BOYER (in press b) sug-

gested that several supraspecific natural

groups may well be distinguished

within the genus Cystiscus. The docu-

mentation displayed in the present

article may help to future tentatives

leading to a new taxonomic organiza-

tion of the much diversified group

Crithe / Cystiscus.

Due to the imprecise limits recog-

nized between the genera Crithe and

Cystiscus, and to the lack of control of

the radulae which would allow to sepa-

rate the Plesiocystiscus species on the

ground of their full rachiglossan tri-

plated radular system, the attributions

proposed herein to these three genera

must be considered as provisional. The

systematics of the family Cystiscidae is

here understood sensu COOVERT AND

COOVERT (1995).

MATERIAL AND METHODS

The studied material belongs to the

MNHN collections and comes princi-

pally from three sources:

The recolts made by P. Bouchet

(MNHN) around Noumea (SW of New

Caledonia) during the years 1992-93.

The recolts made by the Reef Biodi-

versity Workshop organized in Touho

and Koumac from August to October

1993 (Expedition Montrouzier, NE and

* NW of New Caledonia).

The recolts made by a second

worshop organized in Lifou Island

(Expedition Lifou 2000, Loyalty, east off

New Caledonia).

Noumea, Touho and Koumac are

mainland sites with a coral lagoon

within a barrier reef, which is lacking in

Lifou. Koumac and Noumea are on the

leeward side of the mainland island

whereas Touho is on the windward side,

with much resulting differences in terms

of hydrodynamism and sediment types

(P. Bouchet, pers. comm.).

The sampling was mainly based on

advanced technics used by diving, like

succion-pipe (air system) and brushing

boulders and slabs on a basket-box.

These technics were specially applied to

the collect of the cryptic micro-fauna

inhabiting the coral formations, which

appear as sheltering the largest diversity

at the upper sublittoral levels (0-40 m).

243

Iberus, 21 (1), 2003

Some dredgings and trawlings have also

been performed in 20-100 m.

A part of the collected material was

observed in live state. Many species of

Cystiscidae were drawn by P. Bouchet

(Noumea) and by S. Gofas (Touho and

Koumac). A selection of the Gofas

sketches Was presented without tenta-

tive identifications in BOUCHET (1994).

Bouchet' s drawings are adapted herein

by the author; Gofas” drawings are

reproduced in their original state.

Except for few specimens conserved

in alcohol, the material was dried

station by station. Most of the stations

were sorted out later by morphospecies.

G. Coovert examined most of the mater-

ial from Touho and Koumac, and did try

a further selection, labelling provisional

determinations.

The northern area of “Grande Terre”

(mainland of New Caledonia) must be

considered on the whole as the most

documented by these campaigns, the

surroundings of Noumea being not so

intensively sampled, and the recolt from

Lifou being not the subject of live obser-

vations and drawings. So, the present

work is focusing on the data at hand

from the New Caledonian mainland.

In most cases, the environment field

data given for each station do not allow

to infer what is the micro-habitat in

which the subjects were sampled. Some

environment labels express a somewhat

homogeneous habitat and can be used

as such; that is the case for instance in

definitions such as: “fine sand”, “detritic

sand”, “sandy silt on rocky floor”,

“brushing of boulders”. But in the main

cases, the environment labels mention

ambiguous habitat situations (such as

” 18 ,I” 14

“outer slope”, “slopes with silt”, “verti-

cal slopes and overhang”, “succion-pipe

on hard bottoms”, in which the sam-

pling may concern coral alveolus as well

RESULTS

as detritic or sandy pockets) or hetero-

geneous habitat compositions (such as

“hard bottoms, grass”, “boulders, sand,

grass”, “sand, detritic domes”, “succion-

pipe on rocks, sargasses collect”). In

these ambiguous or heterogeneous situ-

ations, the habitat of the subject at hand

is considered as “non-recorded”.

In the course of this work, several

types of Bavay, considered as lost (ROTH

and COAN, 1973), were found by the

author within the Desjardins collection,

deposited in Paris Museum on Septem-

ber 1999 and containing the whole

Bavay' s marginellid collection (pur-

chased by M. Desjardins to Géret' s

widow during the thirties). The redis-

covery of these types does allow to

reassess some cystiscid species whose

status remained uncertain.

Abbreviations:

NC: New Caledonia

AMS: Australian Museum, Sydney

IRD: Institut de Recherche pour le

Développement, Paris (ex-ORSTOM)

IRSNB: Institut Royal des Sciences

Naturelles de Belgique, Bruxelles

MNHG: Muséum national d'Histoire

naturelle de Geneve

MNHN: Muséum national d'Histoire

naturelle, Paris

NMNZ: National Museum of New

Zealand

NSMT: National Science Museum,

Tokyo

FBC: author” s collection

MDC: Maxime Desjardins collection

ad: adult

fr: fragment

juv: juvenile

sh: dead collected subject

spm: live collected subject

stn: station

subad: subadult

Family CYSTISCIDAE Stimpson, 1865

Genus Gibberula Swainson, 1840

Type species: Gibberula zonata Swainson, 1840 = Volvaria oryza Lamarck, 1822, by monotypy.

244

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Gibberula squamosa sp. nov. (Figs. 5-8, 23)

Type material: Holotype (Figs. 5, 6, 23) in MNHN Touho, stn 1269. Numerous paratypes

(ad /juv, spm/sh) from the type locality in MNHN; 2 ad spm in AMS, 2 ad spm in NMNZ; 2 ad

spm in NSMT.

Other material studied: Noumea, 1992: stn 1339, 22* 21.9 S, 166” 15.4” E, 20 m, 6 ad spm, 1 juv

spm; Great Reef Aboré, 22” 22.26' S, 166” 15.92” E, 12-37 m, 4 ad spm, 1 juv spm;, stn 1355, 22”

18.9 S, 166* 26.6' E, 7-10 m, 1 ad sh.

Expedition Montrouzier, Touho, 1993: stn 1237, 20” 46.9 S, 165” 13.8” E, 0-1 m, 4 ad spm, 2 juv

spm, 1 ad sh; stn 1251, 20 46.0'-20* 46.5' S, 165” 13'-165* 14.5' E, 6-15 m, 1 ad sh; stn 1270, 20? 45'

S, 165” 16.5' E, 10-35 m, numerous spm and sh; stn 1272, 20” 49.5' S, 165” 19.6' E, 10 m, numerous

spm; stn 1273, 20 50.4” S, 165” 22.8' E, 20 m, numerous spm and sh.

Expedition Montrouzier, Koumac, 1993: stn 1312, 20? 40.4” S, 164” 14.9 E, 26-40 m, 1 ad spm, stn

1319, 20* 44.7' S, 164? 15.5' E, 15-20 m, 1 juv sh; stn 1323, 20” 40.9 S, 164” 14.8' E, 82-120 m, 1 juv sh.

Expedition Lifou 2000: stn 1435, 20” 55.2” S, 167” 00.7" E, 5-30 m, 8 ad sh (Figs. 7, 8), 3 juv sh.

Type locality: Touho area, Doiman Reef, stn 1269, 20? 35.1 S, 165” 08.1' E, 15-20 m, outer slope.

Etymology: From the scale aspect presented by the shell decoration of rounded and connected

axial convolutions.

Shell description (Figs. 5, 6): Short,

solid, rounded oval. Spire flat, faintly

umbilicate, aperture narrow, moderate

siphonal notch, four distinct columellar

plaits and a faintly distinct fifth upper

one. Lip not distinctly denticulate, fine

spiral lirations on the inner wall of the

outer lip. Outer lip angular, not mar-

ginate.

Axial decoration of very sinuous

brown lines on a creamy ground,

making six blunted convolutions ori-

ented towards the left. Axial lines thick-

ened at the level of the central intervals,

suggesting a darker central spiral zone.

Less distinct dark spiral zones occur at

the upper and lower level of the axial

convolutions, which are merging into

whitish zones at both tips.

Size: 4.25 x 2.80 mm.

Animal description (Fig. 23): Bifur-

cated head, long tentacles and siphon,

foot large and flat. Head, siphon and

foot decorated with large yellowish

patches and small orange stains. The

centre of the head lobes and the central

part of the metapodium are greenish.

Eyes black. The inner mantle shows

alternate zones of yellow, orange and

greenish colour.

Distribution: Known from the whole

NC archipelago, alive from the intertidal

to 26, empty shells to 82 m. Abundant in

Touho, uncommon in Noumea and

Koumac, rare in Lifou.

Habitat: Collected on hard bottoms

with patchy sediments or sandy film, in

silt with dead shells, in fine sands with

grass.

Remarks: G. squamosa sp. nov. shows

close similarities with G. pulchella

(Kiener, 1834) from Australia, which

has however a larger, more slender and

oblong shell, bearing a decoration of

more numerous and sharper axial con-

volutions. The two syntypes of G. pul-

chella pictured herein (Figs. 1-4)

measure respectively 7.9 x 4.5 mm and

7.4 x 4.1 mm (MHNG n* 1152/64),

whereas G. squamosa does not exceed 5

mm in length. G. pulchella seems to be

sympatric in its type locality of Sydney

with at least one other zigzag ornated

species (FBC) of smaller size which

looks as closely related to G. squamosa

and ranges up to South West Australia

(FBC). The complex of zigzag ornated

Gibberula presenting with evidence a

great number of species only distin-

guishable on the basis of subtile differ-

ences (BOYER, in press a), we provision-

ally reserve the name G. squamosa to the

populations represented in NC, charac-

terized by short shells with a limited

number of convolutions, and propose

to report the similar Indo-Pacific popu-

lations as G. cf. squamosa. Shells looking

as very similar to the types of G. pul-

chella are found in Norfolk Island

(EBC)

245

Iberus, 21 (1), 2003

Gibberula lifouana (Crosse, 1871) (Figs. 9-12)

Marginella lifouana Crosse, 1871, p. 205-206. Type figures in Crosse, 1872, pl. II, fig. 2.

Type material: 1 syntype (Fig. 9) in MNHN, here selected as lectotype Label: “Marginella lifouana

Crosse I. Lifou (Loyalty) M. Marie par M.E. Marie typus”.

Other material studied: Expedition Montrouzier, Touho, 1993: stn 1242, 20” 46.2” S, 165” 14.5' E,

tide, 1 ad spm (Fig. 12), 7 juv spm.

Expedition Lifou 2000: stn 1419, 20? 55.6' S, 167” 04.5” E, 5 m, numerous spm and sh, stn 1422, 20?

47.1" S, 167” 07.4' E, 4 m, numerous spm and sh, stn 1424, 20? 54.9 S, 167* 03.0” E, 4m, numerous

spm and sh (Figs. 10, 11); stn 1425, 20? 46.8” S, 167” 07.2” E, 4-5 m, numerous spm and sh, stn

1453, 20? 54.6' S, 167” 02.1' E, 21-30 m, 1 ad spm, 4 ad and juv sh.

Type locality: Lifou Island, Loyalty.

Shell description (Fig. 9): Short, sub-

pyriform. Spire flat, protoconch pro-

duced, aperture moderately narrowed,

widening to the base, deep siphonal

notch, four distinct columellar plaits

and faint upper lirations running along

the apertural wall. Outer lip smooth,

flexuous, angular, not marginate. Axial

decoration of sharp angular zig-zag

mustard yellow lines on a white ground.

Size: 4.5 x 2.8 mm.

Animal: Unknown.

Distribution: G. lifouana is abundant

in upper sublittoral levels from Lifou

Island. The species seems to be very

scarce Off NC mainland (one present

record from Touho). Alive in 0-21 m,

empty shells in 4-21 m.

Habitat: G. lifouana seems to be

restricted to soft bottoms, but it is

recorded from rough sands as well as

fine silty sediments.

Remarks: G. lifovana shows a more

pyriform shell than the one of

G.squamosa, with a slightly wider aper-

ture, a deeper siphonal notch, and a

small teat-like pointed protoconch, not

visible in G.squamosa. Both species are

principally separable on the ground of

their shell decoration, as G. lifovana has

very angular pointed axial zigzag lines, -

whereas G. squamosa has rounded and

blunted convolutions. Any kind of

intergrades does not exist, despite the

fact that both species have been found

sympatrically.

Gibberula cincta sp. nov. (Figs. 13-16)

Type material: Holotype (Figs. 13, 14) in MNHN, Noumea, stn 1347. Paratypes: 20 ad spm

(Figs. 15, 16) + sh and 8 juv spm + sh, in MNHN, 2 ad sh in AMS, 2 ad sh in NMNZ; 2 ad sh in

NSMT.

Other material studied: Noumea, 1993: stn 1367, 22” 24.3' S, 1667 20.7" E, 10 m, 1 ad sh, 3 juv sh;

stn 1368, 22” 24.3' S, 1667 20.7" E, 10 m, 3 ad spm, 1 ad sh, 1 juv sh.

Type locality: Noumea area, Great Reef Aboré, stn 1347, 22? 23.6' S, 1667 20.1' E, 10 m, silty sand

on rock floor.

Etymology: From the spiral ranks of dots ornating the shell.

Shell description (Figs. 13, 14): Short,

solid, subpyriform. Spire almost absent,

small teat-like apex, aperture narrow,

deep siphonal notch, four distinct col-

umellar plaits and a faintly distinct fifth

upper One. Lip not distinctly denticu-

late, no visible spiral lirations on the

inner wall of the outer lip. Outer lip

angular, not marginate.

246

Spiral decoration of six ranks of small

regularly spaced out yellowish brown

dots. A brown continuous line underlines

the suture and reachs the protoconch.

Size: 4.2 x 2.9 mm.

Animal: Unknown.

Distribution: Known only from the

Noumea area, SW coast of NC. Alive

and empty shells in 10 m.

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Habitat: Apparently restricted to soft

bottoms, in white sands and silty sand.

Remarks: G. cincta sp. nov appears as

closely allied to G. lifouana. The “dotted

phase” is also known as occurring as an

intraspecific variation in some zigzag

ornated species of Gibberula. That is for

instance the common phase in Gibberula

deburghi (A. Adams, 1864) from South

West Australia, whereas zigzag ornated

and intergrading subjects may occur in

this species, specially in the northern part

of the species distribution (pers. obs.).

The shell outline of G. cincta is squat-

ter and more rounded than in G.

lifouana, with a less flexuous labrum.

The number of dots on each spiral rank

in G. cincta is higher than the number of

left (or right) oriented convolutions tips

in G. squamosa, and the brown subsu-

tural line doesnot exist in this species.

No kind of intergrading form is

observed in sympatric material.

G. cincta must be accepted as a

species belonging to the zigzag ornated

Gibberula complex. In the case of G.

cincta, the tips of convolutions thickness

are conserved as perfect points, whereas

other species may have conserved

homologous marks as “arrow-pointed

accents” [that is the case for instance in

Gibberula thomensis (Tomlin, 1918) from

the oceanic islands situated off the

coasts of Gabonl.

Gibberula sp. aff. philippii (Figs. 17-22, 24-26)

Sorting out the small white-shelled

Gibberula from Touho and Koumac, G.

Coovert separated three morphospecies

corresponding to the drawings of live

animals made in Touho by S. Gofas:

Morph A corresponds to the shell

pictured in Figures 17 and 18, whose

own animal is pictured in fig. 24 (NC

309). "Two similar pictures (NC 136 and

NC 310) were performed.

Morph B corresponds to the shell pic-

tured in Figures 19 and 20, whose own

animal is pictured in Figure 25 (NC 360).

Morph S corresponds to the shell pic-

tured in Figures 21 and 22, whose own

animal is pictured in Figure 26 (NC 614).

G. Coovert separated as such many

lots of shells from Koumac, and some

few from Touho. A re-assessment of this

arrangement led to distinguish several

homogeneous morphs in some stations,

but also to find many intergrades

linking these morphs in other places (for

shell size as well as shell morphology).

A fourth morph from Koumac separated

by G. Coovert as morph R, presenting a

medium-sized ogival shell, also looks as

a possible intergrade between the

morphs A, Band S.

Even if several species are possibly

occurring here, the informations given

by the field sketches of live animals are

not sufficient in the present state to

define specific unities. All these

morphs belong to the “G. philippii

group” [from the mediterranean G.

philippii (Monterosato, 1878)], whose

species currently show a high vari-

ability of the soft parts. For instance

morph A, as represented in figures 17

and 18, and in figure 24, perfectly

matches the most common form of G.

philippii, by its shell morphology and

size as well as by the chromatism of its

soft parts (see in GOFas, 1990: 129-131,

138). The drawings NC 136 and NC

310 show the same pattern of the soft

parts.

On the other hand, the chromatism

of the soft parts represented in morph

B (Fig. 25) is also found as a frequent

variant of the G. philippi1 chromatism,

and exceptional cases of melanism may

occur too, looking like the sketch

attributed to morph S (Fig. 26).

A resolution of the diversity pattern

within the “G. philippii group” ranging

in NC requires a deeper study of the

variability of the soft parts correlated

with the variability of the shell mor-

phology, on the ground of further field

observations.

247

Iberus, 21 (1), 2003

Genus Crithe Gould, 1860

Type species: Crithe atomaria Gould, 1860, by monotypy.

Crithe caledonica sp. nov. (Figs. 27, 36, 45, 63, 64, 66, 67)

Type material: Holotype (Figs. 27, 36) in MNHN, Noumea, stn 1340. 5 paratypes (4 ad spm, 1

juv spm) in MNHN. 1 ad spm in AMS; 2 ad spm in NMNZ; 1 ad spm in NSMT.

Other material studied: Noumea, 1992: Great Reef Aboré, 22” 22.21' S, 166* 16.15' E, 15-35 m, 5

ad spm (NC 73), 1 juv spm.

Expedition Montrouzier, Touho, 1993: stn 1259, 20” 44.6' S, 165” 13.7" E, 15-35 m, 1 ad spm (Fig.

67); stn 1269, 20? 35.1" S, 165” 08.1' E, 15-20 m, 5 ad spm (Figs. 45, 64), 1 juv spm.

Expedition Montrouzier, Koumac, 1993: stn 1316, 20” 40' S, 164* 11.2 E, 12 m, 16 ad spm and sh

(Fig. 63); stn 1318, 20” 41.4 5, 164” 14.8” E, 20-30 m, 2 ad spm (Fig. 66), lad sh, 6 juv spm; stn

1319, 20” 44.7' S, 164? 15.5” E, 15-20 m, 11 ad spm and sh, 7 juv spm and sh; stn 1331, 20? 40'-20*

40.6' S, 164” 11.2'-164* 12.1” E, 55-57 m, 7 ad sh, 1 juv sh.

Expedition Lifou 2000: stn 1429, 20* 47.5'.5, 167” 07.1” E, 8-18 m, 7 ad sh, 4 juv sh; stn 1436, 20"

55.5 S, 167” 04.2 E, 47 m, 2 ad sh; stn 1449, 20* 45.8' S, 167* 01.65' E, 17 m, 1 ad sh; stn 1450, 209

45.8' S, 167* 01.65 E, 27-31 m, 2 ad sh; stn 1451, 20* 47.3 S, 167* 06.8' E, 10-21 m, 2 ad sh; stn

1454, 20 56.65' S, 167" 02.0' E, 15-18 m 2 ad sh.

Type locality: Noumea area, Tetembia Reef, stn 1340, 22” 21.0' S, 166” 14.0' E, 30 m, outer slope.

Etymology: From the widespread distribution of the species around NC mainland.

Shell description (Fig. 27): White,

short, solid, inflated, rounded subpyri-

form. Spire excavated with a flat top,

aperture widely opening to the base,

three distinct anterior columellar plaits

and four upper columellar varix on the

parietal wall. Inner lip faintly denticu-

late, shoulder of the outer lip low.

Size: 1.9 x 1.3 mm.

Animal description (Fig. 36): Bifur-

cated head with long frontal lobes, foot

small and triangular. Frontal lobes and

lateral sides ahead of the red eyes are

yellow, lateral sides of the head behind

the eyes are reddish, borders of the

central slit are orange in their frontal

part and lighter behind. Eyes red. The

foot is whitish, with mottled white

patches on the metapodium, the inner

mantle is whitish.

Distribution: Known from Noumea,

Touho, Koumac and Lifou, the species

does range all around the NC. area.

Alive from 12 to 30 m, empty shells

from 8 to 55 m.

Habitat: Not accurately established.

Live specimens seem to be associated to

reef coral formations in the whole upper

sublittoral zone..

Remarks: By its very rounded and

solid shell and by the original pattern of

its spire, C. caledonica sp. nov. shows as

much distinct from all the other species

belonging to the Crithe/Cystiscus

(Right page) Figures 1-4: Gibberula pulchella, syntypes, 7.9 x 4.5 mm and 7.4 x 4.1 mm, Sydney

(MHNG). Figures 5-8: G. squamosa; 5, 6: holotype, 4.25 x 2.80 mm, Touho, stn 1269 (MNHN);

7,8: 4.0 x 2.5 mm, Lifou, stn 1435. Figures 9-12: G. lifouana; 9: holotype, 4.5 x 2.8 mm, Lifou

(MNHN); 10, 11: 4.2 x 2.7 mm, Lifou, stn 1424; 12: 4.2 x 2.8 mm, Toubho, stn 1242. Figures 13-

16: G. cincta; 13, 14: holotype, 4.2 x 2.9 mm, Noumea, stn 1347 (MNAHN); 15, 16: paratype, 4.50

x 3.05 mm, Noumea, stn 1347 (MNHN).

(Página derecha) Figuras 1-4: Gibberula pulchella, sintipos, 7,9 x 4,5 mm and 7,4 x 4,1 mm, Sydney

(MHNG). Figuras 5-8: G. squamosa; 5, 6: holotipo, 4,25 x 2,80 mm, Toubo, stn 1269 (MNAN); 7,

8: 40 x 2,5 mm, Lifou, stn 1435. Figuras 9-12: G. lifouana; 9: holotipo, 45 x 2,8 mm, Lifou

(MNHN); 10, 11: 42 x 2,7 mm, Lifou, stn 1424; 12: 42 x 2,8 mm, Toubho, stn 1242. Figuras 13-

16: G. cincta; 13, 14: holotipo, 4,2 x 2,9 mm, Noumea, stn 1347 (MNAN); 15, 16: paratipo, 450

x 3,05 mm, Noumea, stn 1347 (MNHN).

248

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Iberus, 21 (1), 2003

complex. The shell size and morphology

are very constant.

Whereas the chromatism of the head

seems to be somewhat constant (Figs.

36, 63, 64, 66, 67), the ground color of

the foot and the inner mantle can vary

noticeably. The inner mantle ranges

from dirty white to creamy yellow, light

yellow and red, or light orange-red. The

foot ranges from dull white to light yel-

lowish or very light reddish.

The species seems to be abundant all

around the NC mainland and in Loyalty

Islands.

Crithe gofasi sp. nov. (Figs. 46, 65)

Type material: Holotype (Figs. 46, 65) in MNHN, Touho, stn 1270.

Type locality: Touho area, Great Reef Mengalia, stn 1270, 20” 45' S, 165” 16.5' E, 10-35 m, outer slope

Etymology: For Serge Gofas, University of Malaga (Spain), who was a pioneer of the revival of

the study of marginelliform gastropods in the recent times, and who devoted to an accurate pic-

turing of New Caledonian marginellids in the field.

Shell description (Fig. 46): White,

short, solid, inflated, rounded subpyri-

form. Spire slightly excavated with an

undulate top, outer lip much arched

with a shoulder much elevated, aper-

ture widely opening to the base, three

distinct anterior columellar plaits. Inner

lip smooth.

SrzeL.o9 xi i0mm:

Animal description (Fig. 65): Bifur-

cated head with medium size frontal

lobes, foot small and triangular.

Head hyalinous with light orange

borders, except around and behind the

red eyes. The borders of the central slit

are darker orange. The foot is whitish

translucent, mottled white on the

metapodium with a reddish axis. The

inner mantle is mottled light reddish

and yellow.

Distribution: Only known by the

holotype from Touho, NE coast of NC

mainland. Alive in 10 m.

Habitat: Not recorded.

Remarks: C. gofasi sp. nov. can be con-

sidered as a sibling species of C. cale-

donica. The distinct features of C. gofasi

are a smaller size, a more elevated and

arched outer lip, a discoloured zone

around and behind the eyes, a reddish

axis on the metapodium.

As these special features were not

observed in the numerous shells of C.

caledonica and in the different specimens

live pictured from Noumea, Touho and

Koumac, and considering the fact that

the holotype of C. gofasi was collected in

close vicinity of C. caledonica specimens,

a specific distinction can play validly

here.

Genus Cystiscus Stimpson, 1865

Type species: Cystiscus capensis Stimpson, 1865 (non Marginella capensis Krauss, 1848) = Mar-

ginella cystiscus Redfield, 1870 (nom. nov.), by monotypy.

Cystiscus viridis sp. nov. (Figs. 28, 37)

Type material: Holotype (Figs. 28, 37) in MNHN, Noumea, stn 1340. 1 paratype (ad spm) from

the type locality, in MNHN.

Type locality: Noumea area, Tetembia Reef, stn 1340, 22? 21.0' S, 166" 14.0', 30 m, outer slope.

Etymology: From the dominant green colour of the animal.

Shell description (Fig. 28): Translucent

white, short, solid, inflated, rounded

subpyriform. Spire very small, rounded

250

protoconch slightly protuberant, aper-

ture moderately widening to the base,

three distinct anterior columellar plaits

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Figures 17-22. Gibberula sp. aff. philippiz. 17, 18: morph A, 1.75 x 1.20 mm, Touho, stn 1259; 19,

20: morph B, 2.90 x 1.75 mm, Touho, stn 1271; 21, 22: morph S, 2.35 x 1.50 mm, Touho, stn

1268.

Figuras 17-22. Gibberula sp. aff. philippii. 17, 18: morfo A, 1,75 x 1,20 mm, Touho, stn 1259;

19, 20: morfo B, 2,90 x 1,75 mm, Toubo, stn 1271; 21, 22: morfo S, 2,35 x 1,50 mm, Toubo,

stn 1268.

Iberus, 21 (1), 2003

Figure 23: Gibberula squamosa, holotype, Touho, stn 1269 (NC 334). Figure 24: G. sp. aff. philip-

pii morph A, Touho, stn 1259 (NC 309). Figure 25: G. sp. aff. philippii morph B, Touho, stn 1271

(NC 360). Figure 26: G. sp. aff. philippii morph S, Touho, stn 1268 (NC 614).

Eigura 23: Gibberula squamosa, holotipo, Touho, stn 1269 (NC 334). Figura 24: G. sp. afk. philip-

pii morfo A, Touho, stn 1259 (NC 309). Figura 25: G. sp. af£. philippii morfo B, “Touho, stn 1271

(NC 360). Figura 26: G. sp. aff£. philippii morfo S, Touho, stn 1268 (NC 614).

252

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Figure 27: Crithe caledonica, holotype, 1.9 x 1.3 mm, Noumea, stn 1340 (MNHN). Figure 28: Cystiscus

viridis, holotype, 1.85 x 1.10 mm, Noumea, stn 1340 (MNHN). Figure 29: C. punctatus, holotype, 1.5

x 1.0 mm, Noumea, stn 1336 (MNAHN). Figure 30: C. aurantius, holotype, 1.65 x 1.15 mm, Noumea,

sen 1340 (MNHN). Figure 31: C. montrouzieri, lectotype, 1.55 x 0.95 mm, Lifou (MNHN). Figure 32:

C. montrouzieri, 1.45 x 0.95 mm, Noumea, stn 1360. Figure 33: C. marshalli, holotype, 1.8 x 1.0 mm,

Noumea (MNHN). Figure 34: C. boucheti, holotype, 1.4 x 0.8 mm, Noumea, stn 1343 (MNHN). Figure

35: C. camelopardalis, holotype, 1.3 x 0.7 mm, Noumea, stn 1336 (MNHN).

Figura 27: Crithe caledonica, holotipo, 1,9 x 1,3 mm, Noumea, stn 1340 (MNAN). Figura 28: Cys-

tiscus viridis, holotipo, 1,85 x 1,10 mm, Noumea, stn 1340 (MNAN). Figura 29: C. punctatus, holotipo,

1,5x 1,0 mm, Noumea, stn 1336 (MNAN). Figura 30: C. aurantius, holotipo, 1,65 x 1,15 mm, Noumea,

sn 1340 (MNHAN). Figura 31: C. montrouzieri, lectotipo, 1,55 x 0,95 mm, Lifou (MNHN). Figura

32: C. montrouzieri, 1,45 x 0,95 mm, Noumea, stn 1360. Figura 33: C. marshalli, holotipo, 1,8 x 1,0

mm, Noumea (MNAN). Figura 34: C. boucheti, holotipo, 1,4 x 0,8 mm, Noumea, stn 1343 (MNAN).

Figura 35: C. camelopardalis, holotipo, 1,3 x 0,7 mm, Noumea, stn 1336 (MNAHN).

ZSS

Iberus, 21 (1), 2003

and six upper visible low columellar

varix on the parietal wall. Labrum is

somewhat right in its lower part, two

profile breakings in the upper part,

inner border smooth, low shoulder.

Size: 1.85 x 1.10 mm.

Animal description (Fig. 37): Bifur-

cated head with long frontal lobes, foot

small and triangular. Head and foot

light green. Eyes red. Inner mantle

greenish with creamy white spots, exter-

nal mantle dark green.

Distribution: Only known by two

specimens from the same sampling in

Noumea, SW coast of NC mainland.

Alive in 30 m.

Habitat: Not recorded.

Remarks: C. viridis sp. nov. is distinct

from its relatives by its large tear-

shaped shell and by the green chroma-

tism of its animal.

C.viridis may be closely related to

Cystiscus ¡ota (Hedley, 1899) from Funa-

futi (Ellice Islands), which presents a

more slender and narrowed shell, a

more sinuous and somewhat elevated

labrum, and a longer and sinuous first

columellar plait.

Cystiscus punctatus sp. nov. (Figs. 29, 38, 73)

Type material: Holotype (Figs. 29, 38) in MNHN, Noumea, stn 1336. Paratypes: 1 subad and 2

juv spm from the type locality, in MNHN.

Other material studied: Noumea, 1992: Great Reef Aboré, 22* 22.21' S, 166* 16.15' E, 15-35 m, 2

ad spm.

Expedition Montrouzier, Koumac, 1993: stn 1318, 20” 41.4' S, 164?” 14.8' E, 20-30 m, 2 juv spm (Fig. 73).

Type locality: Noumea area, lle aux Canards, stn 1336, 22* 19.0' S, 1667 26.1” E, 18 m, boulders

and sargasses.

Etymology: From the decoration of the inner mantle of the animal, made of black spots on an

orange ground.

Shell description (Fig. 29): Translucent

white, short, solid, moderately inflated,

faintly subpyriform. Spire very short,

slightly angular, aperture moderately

widening to the base, three distinct ante-

rior columellar plaits. Outer lip very

arched in its upper part, straighter in its

lower part, inner lip straight and rather

smooth.

Size: 1.5 x 1.0 mm.

Animal description (Fig. 38): Bifur-

cated head with long frontal lobes, foot

small and triangular. Head and foot

deep orange. Eyes red. Inner mantle

orange with black spots.

Distribution: Only known from two lots

from Noumea and one lot from Koumac,

both localities situated on the west coast

of NC mainland. Alive from 15 to 20 m.

Habitat: Not recorded.

Remarks: C. punctatus sp. nov. is dis-

tinct from its relatives by its orange ani-

mal spotted with black on the inner man-

tle. The shell, even if smaller, shows some

similarities with that ones of C. viridis

and of C. goubini (Bavay, 1922) (cf. infra).

Cystiscus aurantius sp. nov. (Figs. 30, 39)

Type material: Holotype (Figs. 30, 39) in MNHN, 1.65 x 1.15 mm. Noumea, stn 1340. 1 paratype

(ad spm) from the type locality, in MNHN.

Other materiel studied: Noumea, 1992: Great Reef Aboré, 22* 22.21' S, 166* 16.15' E, 15-35 m, 4

ad and 1 subad spm.

Type locality: Noumea area, Tetembia Reef, stn 1340, 22* 21.0' S, 166" 14.0' E, 30 m, outer slope.

Etymology: From the general orange shade of the animal.

Shell description (Fig. 30): White,

short, solid, subtriangular. Spire hidden,

254

top slightly concave, aperture moder-

ately opening to the base, outer lip

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Figure 36: Crithe caledonica, holotype, Noumea, stn 1340 (NC 22). Figure 37: Cystiscus viridis,

holotype, Noumea, stn 1340 (NC 23). Figure 38: C. punctatus, holotype, Noumea, stn 1336 (NC

12). Figure 39: C. aurantius, holotype, Noumea, stn 1340 (NC 24). Figure 40: C. montrouzieri,

Noumea, stn 1360 (NC 78). Figure 41: C. marshalli, holotype, Noumea (NC 75). Figure 42: C.

boucheti, holotype, Noumea, stn 1343 (NC 52). Figure 43: C. sp. 1, Noumea (NC 14). Figure 44:

C. camelopardalis, holotype, Noumea, stn 1336 (NC 13).

Figura 36: Crithe caledonica, holotipo, Noumea, stn 1340 (NC 22). Figura 37: Cystiscus viridis,

holotipo, Noumea, stn 1340 (NC 23). Figura 38: C. punctatus, holotipo, Noumea, stn 1336 (NC

12). Figura 39: C. aurantius, holotipo, Noumea, stn 1340 (NC 24). Figura 40: C. montrouzieri,

Noumea, stn 1360 (NC 78). Figura 41: C. marshalli, holotipo, Noumea (NC 75). Figura 42: C.

boucheti, holotipo, Noumea, stn 1343 (NC 52). Figura 43: C. sp. 1, Noumea (NC 14). Figura 44:

C. camelopardalis, holotipo, Noumea, stn 1336 (NC 13).

235

Iberus, 21 (1), 2003

arched in its upper part, with a highly

produced shoulder, lower lip rather

straight, inner lip almost smooth, three

distinct anterior columellar plaits and

one small columellar varix just above

the third plait.

Animal description (Fig. 39): Bifur-

cated head with medium sized frontal

lobes, foot small. Head and foot deep

orange. Eyes red. Inner mantle creamy

orange.

Distribution: Only known from two

stations off Noumea, in upper sublit-

toral. Alive from 15 to 30 m.

Habitat: Not recorded.

Remarks: C. aurantius sp. nov. is dis-

tinct from C. punctatus by its

heartshaped shell as well as the uniform

creamy orange inner mantle of its

animal, contrasting with the deep

orange black spotted inner mantle

found in C. punctatus.

Cystiscus montrouzieri (Bavay, 1922) (Figs. 31, 32, 40)

Marginella montrouzieri Bavay, 1922, p. 62-63, pl. 1, figs. 4-5.

Type material: Lectotype (Fig. 31) and 6 paralectotypes (5 ad and 1 juv sh) in MNHN (ex-MDC).

Labels: “Ma. Montrouzieri By 1. Lifou sables”, from the hand of Bavay, and “TYPE” in typed red letters.

The specimen recorded as “holotype” (1.5 x 1.0 mm) by ROTH AND CLOVER (1973: 212) is a juv sh,

whereas the type figures in BAVAY (1922: pl. 1, figs. 4, 5) show clearly an adult subject. Further-

more, any label does not certify a status of holotype for the shell represented in “Collection du

Journal”, and this shell may as well belong to a different species that the one described and pic-

tured by Bavay. So this specimen can only be considered as a syntype, and it is designated here

as paralectotype n” 7, MNHN. |

Other material studied: Noumea, 1993: Mberé Reef, stn 1360, 22* 19.9' S, 166" 13.2 E, 10-15 m, 1

ad (Figs. 32, 40) and 1 subad spm.

Type locality: Lifou.

Shell description (Fig. 31): White,

short, narrow, suboval outline tapering

to the base, left side much arched, right

side angular in its upper part. Spire

hidden, aperture high and narrow, inner

lip straight, smooth, two strong anterior

columellar plaits and six upper small

plaits or lirations ranging all along the

columellar border.

Size: 1.55 x 0.95 mm.

Animal description (Fig. 40): Bifur-

cated head with medium sized frontal

lobes, foot small and triangular. Head

reddish-orange, eyes red. Foot yellow

with a reddish axis on he metapodium.

Inner mantle light yellowish with small

blackish stains.

Distribution: Described from Lifou.

The provisional attribution to the

species of the form observed in Noumea

deserves to be confirmed by a control of

the shell variability and overall of the

animal chromatism occurring in Lifou.

In Noumea, alive at 10-15 m. BAVAY

(1922: 63) records a “very analogous

form” from Kermadec Islands. Similar

forms were not observed in other places

from NC mainland. ]

Habitat: Brushed alive on coral boul-

ders.

Remarks: As far as the shell morphol-

ogy is concerned, C. montrouzieri is prin-

cipally comparable to C. bougei (Bavay,

1917) (see infra). It must be underlined

that the ad spm from Noumea (Fig. 32)

is noticeably different from the typical

form, mainly due to the lower rupture

of the outer profile of the labrum.

However this feature might be represen-

tative of an intraspecific variability.

Cystiscus marshalli sp. nov. (Figs. 33, 41, 55, 76)

Type material: Holotype (Figs. 33, 41) in MNHN. Noumea, Great Reef Aboré. 1 paratype (ad

spm) from the type locality, in MNHN.

256

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Figure 45: Crithe caledonica, 1.75 x 1.30 mm, Touho, stn 1269. Figure 46: C. gofasi, holotype, 1.55

x 1.10 mm, Touho, stn 1270 (MNHN). Figure 47: Cystiscus minor, holotype, 1.2 x 0.8 mm, Touho,

sen 1271 (MNHN). Figures 48-50: C. goubini; 48: ex auct, 2.3 x 1.4 mm (MNHN); 49: 2.00 x 1.35

mm, Touho, stn 1269; 50: 1.90 x 1.25 mm, Koumac, stn 1318. Figure 51: C. pardus, holotype, 1.55

x 1.00 mm, Koumac, stn 1318 (MNHN). Figure 52: C. deltozdes, holotype, 1.50 x 0.95 mm, Koumac,

sin 1331 (MNHN). Figure 53: C. caeruleus, holotype, 1.7 x 1.0 mm, Touho, stn 1270 (MNHN).

Figura 45: Crithe caledonica, 1,75 x 1,30 mm, Toubo, stn 1269. Figura 46: C. gofasi, holotipo, 1,55

x 1,10 mm, Touho, stn 1270 (MNHN). Figura 47: Cystiscus minor, holotipo, 1,2 x 0,8 mm, Touho,

stn 1271 (MNAN). Figuras 48-50: C. goubini; 48: ex auct, 2,3 x 1,4 mm (MNHN); 49: 2,00 x 1,35

mm, Touho, stn 1269; 50: 1,90 x 1,25 mm, Koumac, stn 1318. Figura 51: C. pardus, holotipo, 1,55 x

1,00 mm, Koumac, stn 1318 (MNHN). Figura 52: C. deltoides, holotipo, 1,50 x 0,95 mm, Koumac,

sn 1331 (MNAN). Figura 53: C. caeruleus, holotipo, 1,7 x 1,0 mm, Toubo, stn 1270 (MNAN).

237

Iberus, 21 (1), 2003

Other material studied: Expedition Montrouzier, Touho, 1993: stn 1271, 20* 52.7" S, 165* 19.5' E,

5-25 m, 6 ad spm (Fig. 55); stn 1272, 20* 49.5' S, 165” 19.6' E, 10 m, 1 ad spm (Fig. 76).

Expedition Montrouzier, Koumac, 1993: stn 1318, 20” 41.4” S, 164” 14.8' E, 20-30 m, 1 ad spm.

Type locality: Noumea area, Great Reef Aboré, 22” 22.21' S, 166” 16.15' E, 15-35 m, outer slope.

Etymology: For Bruce Marshall (NMNZ) who cooperated with P. Bouchet during the Noumea

Expedition in November 1992, in which this species was first collected.

Shell description (Fig. 33): Translucent

white, slender, light, suboval. Spire

merged into the insertion of the much

elevated labrum. Aperture regularly

opened, inner lip smooth, one large first

columellar plait and two upper packed

small ones.

Size: 1.8 x 1.0 mm.

Animal description (Fig. 41): Bifur-

cated head with medium sized frontal

lobes, foot small. Head yellow with a

long axial reddish band running from

each frontal lobe to the base of the head,

passing between the eye and the central

slit. Eyes red. Foot light yellow. Inner

mantle uniformly deep yellow.

Distribution: Known from Noumea,

Touho and Koumac, the species is sup-

posed to range all around the NC main-

land. Alive from 5 to 20 m.

Habitat: Not recorded.

Remarks: The shell as well as the

animal of C. marshalli show as very con-

stant. The animal of C. montrouzieri pre-

sents some similarities. However the

shells of both species are noticeably

divergent in several respects, and a close

relationship is not evident.

Cystiscus boucheti sp. nov. (Figs. 34, 42)

Type material: Holotype (Figs. 34, 42) in MNHN. Noumea, stn 1343. Paratypes 7 ad and 1 subad

spm from the type locality, in MNHN.

Other material studied: Noumea, 1992: Lagoon, Maitre Islet, 22” 20.41” S, 166" 25.69 E, low tide,

10 ad spm.

Type locality: Noumea area, Senez Reef, stn 1343, 22” 17.8' S, 166” 19.9 E, 7 m, inner slope.

Etymology: For Philippe Bouchet (MNHN), who led the coastal campaigns which procured the

material studied in this article, and who pictured valuable descriptive sketches of live cystiscids

in Noumea.

Shell description (Fig. 34): Translucent

white, light, regular slender oval outline,

narrow. Aperture very long and narrow,

contracted in its mid-part, no apex. Outer

lip thin and smooth, straight in its lower

2/3 parts and arched upper. Two first col-

umellar strong plaits, a third one faint,

four small columellar varix occupy the

half part of the remaining parietal wall.

Size: 1.4 x 0.8 mm.

Animal description (Fig. 42): Bifur-

cated head with long frontal lobes, foot

small and triangular. Head light pinkish

orange, eyes red. Foot pink with a

darker axis on the metapodium. Inner

mantle coloured by large packed pinky

white and chocolate brown layers.

Distribution: Only known from two

stations off Noumea. Alive from inter-

tidal to 7 m.

Habitat: Not recorded.

Remarks: According to the field notes

by P. Bouchet, the colour pattern of the

animal is much constant, even if the

inner mantle is more patchy in some

specimens.

Cystiscus sp. 1 (Fig. 43)

In his drawing NC 14 (Noumea area,

lle aux Canards, 18 m, 1992-06-30), P.

Bouchet pictured an animal comparable

to C. boucheti, but showing also some

258

constant differences even in the one

station where both forms were observed

as living in micro-sympatry (Senez Reef,

7 m, 1992-09-07). The corresponding

BOYER: The Cystiscidae from upper reef formations of New Caledonia

0 (y) (y)

Figure 54: Cystiscus tricinctus, holotype, 1.60 x 0.95 mm, Touho, stn 1269 (MNHN). Figure 55: C.

marshallí, 1.65 x 0.95 mm, Touho, stn 1271. Figure 56: C. pseudoaurantius, holotype, 1.9 x 1.3

mm, Touho, stn 1255 (MNHN). Figure 57: C. cooverti, holotype, 1.40 x 0.85 mm, Koumac, stn

1312 (MNHN). Figure 58: C. bougez, lectotype, 1.4 x 1.0 mm, Wallis (MNHN). Figure 59: C.

bougeí, 1.35 x 0.85 mm, Koumac, stn 1279.

Figura 54: Cystiscus tricinctus, holotipo, 1,60 x 0,95 mm, Toubo, stn 1269 (MNHN). Figura 55: C.

marshalli, 1,65 x 0,95 mm, Touho, stn 1271. Figura 56: C. pseudoaurantius, holotipo, 1,9 x 1,3 mm,

Touho, stn 1255 (MNHN). Figura 57: C. cooverti, holotipo, 1,40 x 0,85 mm, Koumac, stn 1312

(MNHN). Figura 58: C. bougei, lectotipo, 1.4 x 1.0 mm, Wallis (MNHN). Figura 59: C. bougel,

1,35 x 0,85 mm, Koumac, stn 1279,

specimens from both stations were

packaged separately in the field, but the

lots corresponding to Cystiscus sp. seem

to have been lost later.

Animal description (Fig. 43): Bifur-

cated head with medium size frontal

lobes, foot small. Head and foot pink.

Eyes red. Inner mantle made of a large

alternated zone of chocolate brown and

pink, this last zone being bordered by a

narrow white fringe.

Remarks: In his field notes, P. Bouchet

insists on the fact that the colour pattern

of the inner mantle is constant in this

form, and that it can be distinguished

from C. boucheti by several features: its

size is smaller, its foot is more uniformly

pink coloured, the clear zones of its inner

mantle are pink with a white fringe, and

not uniformly pinky white.

In these conditions, it can be

assumed that two sibling species are

occurring here. Waiting for the rediscov-

ery (or new recolts) of Cystiscus sp. 1

specimens, this morph is kept as an

unnamed species.

LID

Iberus, 21 (1), 2003

Cystiscus camelopardalis sp. nov. (Figs. 35, 44)

Type material: Holotype (Figs. 35, 44) in MNHN. Noumea, stn 1336. 3 paratypes: 2 ad and 1 juv

spm, in MNHN.

Type locality: Noumea area, lle aux Canards, stn 1336, 22” 19.0' S, 166" 26.1” E, 18 m, boulders

and sargasses.

Etymology: From the chromatism of the soft parts, with black subquadrate patches on a yellow

ground, similar to the coat decoration of the giraffe (Latin form = camelopardalis).

Shell description (Fig. 35): Translucent

white, slender cylindrical. Small pro-

duced spire, teat-like apex, aperture

narrow, moderately widened to the

base, labrum straight, inner lip smooth,

two long anterior columellar plaits and

a faintly produced third one.

Size: 1.0 x 0.7 mm.

Animal description (Fig. 44): Bifurcated

head with long sized frontal lobes, foot

very small and narrow. Head and foot

deep yellow, eyes red. Inner mantle deep

yellow with deep black irregular slabs.

Distribution: Only known by one lot

from Noumea area. Alive in 18 m.

Habitat: Not recorded.

Remarks: By its shell as well as its

animal chromatism, C. camelopardalis sp.

nov. shows as very distinct from all the

other species of Cystiscus.

Cystiscus minor sp. nov. (Figs. 47, 68, 69)

Type material: Holotype (Fig. 47) in MNHN. Touho, stn 1271. Paratypes: 8 ad and 1 juv spm

from the type locality, in MNHN.

Other material studied: Expedition Montrouzier, Touho, 1993: stn 1259, 20” 44.6' S, 165” 13.7' E,

15-35 m, 1 ad spm (Fig. 68); stn 1270, 20* 45' S, 165* 16.5' E, 10-35 m, 4 ad spm, stn 1271, 20* 52.7

S, 1657 19.5' E, 5-25 m, 1 ad spm (Fig. 69).

Expedition Montrouzier, Koumac, 1993: stn 1310, 20? 39.7" S, 164” 14.9 E, 15 m, 3 ad and 1 juv

spm; stn 1312, 20? 40.4” S, 164* 14.9 E, 26-40 m, 21 ad and 1 juv spm, 2 ad sh; stn 1318, 20* 41.4' S,

164” 14.8' E, 20 -30 m, 2 ad and 1 juv spm.

Type locality: Touho area, Tié shallows, stn 1271, 20* 52.7' S, 165* 19.5' E, 5-25 m, cliffs, sand on

rocky floor.

Etymology: From the small size of the species.

Shell description (Fig. 47): Translucent

white, short, subtriangular. Spire faintly

produced, teat-like protoconch, labrum

elevated and arched in its upper part,

straight in its central and lower part.

Aperture narrow, faintly widening to

the base, inner lip smooth, three anterior

columellar plaits and seven faint pari-

etal varix.

Size: 1.2 x 0.8 mm.

Animal description (Fig. 68): Bifur-

cated head with short frontal lobes, foot

small and triangular. Head whitish

fringed with light orange, more reddish

around the central slit. Eyes red. Foot

mottled opaque white. Inner mantle

whitish with beige shades.

Distribution: Known from Touho and

Koumac, in north of the NC mainland.

Alive from 5 to 26 m, empty shells at 26 m.

260

Habitat: Not recorded.

Remarks: In its northern distribution,

C. minor sp. nov. seems to be rather

common and live in somewhat dense

populations.

The shell morphology is constant,

but the chromatism of the animal some-

what variable (Figs. 68, 69). The light

orange zones on the head periphery

may be replaced by creamy yellow

zones restricted to the frontal lobes. The

reddish fringes may be very narrowed

and also restricted to the frontal lobes.

The foot may be mottled light

yellow better than white, with a fine red

fringe ahead. The inner mantle may be

deep white.

About specimens observed from Man-

galia Reef (Touho), S. Gofas noted that the

head may be “more or less orange”.

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Figure 60: Plesiocystiscus tomlini, lectotype, 2.20 x 1.25 mm, Ouvea, Wallis Is. (MNHN). Figures

61, 62. P bavayi. 61: holotype, 3.30 x 1.95 mm, Touho, stn 1261 (MNHN); 62: 3.10 x 1.70 mm,

Touho, stn 1271.

Figura 60: Plesiocystiscus tomlini, lectotipo, 2.20 x 1.25 mm, Ouvea, Wallis ls. (MNAN). Figuras 61,

62. P. bavayi. 61: holotipo, 3.30 x 1.95 mm, Toubo, sin 1261. (MNHN); 62: 3.10 x 1.70 mm, Touho,

sn 1271. cd

261

Iberus, 21 (1), 2003

Cystiscus goubini (Bavay, 1922) (Figs. 48-50, 70, 71)

Marginella goubini Bavay, 1922, p. 60-61, pl. L, fig. 10

Type material: Holotype said to be lost (ROTH and CLOVER, 1973: 211).

Other material studied: MDC: 11 ad sh (Fig. 48), labelled “Goubini” from the hand of Bavay.

Expedition Montrouzier, Touho, 1993: stn 1269, 20* 35.1” S, 165” 08.1” E, 15-20 m, 3 ad spm (Fig.

49, 70); stn 1271, 20? 52.7' S, 165” 19.5' E, 5-25 m, 3 juv spm.

Expedition Montrouzier, Koumac, 1993: stn 1299, 20* 34.4” S, 164” 13.0” E, 12-14 m, 1 ad spm; stn

1312, 20? 40.4' S, 164” 14.9 E, 26-40 m, 5 ad and 2 juv spm 2 ad sh; stn 1316, 20? 40' S, 164” 11.2 E,

12 m, 10 ad and 2 juv spm, 4 ad sh; stn 1318, 20” 41.4" S, 164” 14.8' E, 20-40 m, 4 ad and 3 juv spm,

1 ad sh, (Figs. 50, 71); stn 1319, 20” 44.7" S, 164” 15.5' E, 15-20 m, 5 ad and 2 juv spm;, stn 1331, 20*

40-20? 40.6' S, 164* 11.2'-164* 12.1 E, 55-57 m, 1 ad sh.

Expedition Lifou 2000: stn 1429, 20 47.5' S, 167” 07.1' E, 8-18 m, 1 ad spm, 2 ad sh; stn 1436, 20%

55.5' S, 1677 04.2” E, 10-20m, 4 ad sh; stn 1442, 20? 46.4” S, 167” 02.0' E, 47 m, 2 ad sh.

Type locality: Lifou.

Shell description (Fig. 48): White, solid,

egg-shaped outline, inflated, faintly

pyriform. Aperture long and narrow,

much widening to the base. Outer lip

thick and rounded in its upper part, thin

and flexuous in its lower part. Two

strong anterior columellar plaits, two

next ones fainter, and six regular spaced

lirations along the parietal border.

Size: 1.9 to 2.3 mm x 1.25 to 1.40 mm.

Animal description (Figs. 70, 71): Bifur-

cated head with medium sized frontal

lobes, foot medium sized and triangular.

The borders of the head are light orange,

more pronounced along the central slit, on

a whitish to yellow background. Eyes red.

The foot is opaque greyish to dirty white, a

fine orange vein may run ahead. Inner

mantle greyish blue, paving pattern of

irregular figures. External mantle is mot-

tled greyish blue with an irregular surface.

Distribution: Known from Lifou

(type locality) and from northern NC

(Touho and Koumac). Alive from 5 to 26

m, empty shells from 8 to 55 m.

Habitat: Hard bottoms.

Remarks: The shell of C. goubini

(Bavay, 1922) is distinct by its large

size and its bulbous shape. The animal

chromatism differs noticeably from all

the other NC species, except from the

species considered hereunder.

Due to the poor datum joined to

the lot of C.goubini found in the

Bavay's collection, and to the absence

of a “TYPE” label, the status of this lot

as type material is not ascertained. The

designation of a neotype is provision-

ally refrained, waiting for the possible

rediscovery of such a type lot within a

public institution, for instance in the

Dautzenberg Collection, IRSNB.

Cystiscus pardus sp. nov. (Figs. 51, 72)

Type material: Holotype (Figs. 51, 72) in MNHN. Koumac, stn 1318. 1 paratype (ad spm) from

the type locality, in MNHN.

Other material studied: Expedition Montrouzier, Koumac, 1993: stn 1319, 20” 44.7' S, 164* 15.5'

E, 15-20 m, 3 ad spm.

Type locality: Koumac area, Barrier Reef, stn 1318, 20? 41.4' S, 164? 14.8' E, 20-30 m, outer slope.

Etymology: From the leopard patterned inner mantle of the animal.

Shell description (Fig. 51): Translu-

cent white, egg-shaped, inflated, faintly

pyriform. Spire flat, labrum arched,

inner lip smooth, three anterior col-

umellar plaits.

Size: 1.55 x 1.00 m.

262

Animal description (Figs. 72): Bifur-

cated head with short frontal lobes, foot

small and triangular. Head fringed with

light orange on a whitish and yellowish

ground. Eyes red. Foot covered with

large clouds of light yellowish to

BOYER: The Cystiscidae from upper reef formations of New Caledonia

G E )

68x, A 1

) Ñ

) |

Figures 63, 64: Crithe caledonica. 63: Koumac, stn 1316, (NC 657); 64: Touho, stn 1269 (NC 328).

Figure 65: C. gofasi, holotype, Touho, stn 1270 (NC 329). Figures 66, 67: C. caledonica. 66: Koumac,

sin 1318 (NC 648); 67: Touho, stn 1259 (NC 301). Figures 68, 69: Cystiscus minor. 68: Touho, stn

1259 (NC 302); 69: Touho, stn 1271 (NC 372).

Figuras 63, 64: Crithe caledonica. 63: Koumac, stn 1316, (NC 657); 64: Touho, stn 1269 (NC 328).

Figura 65: C. gofasi, holotype, Touho, stn 1270 (NC 329). Figuras 66, 67. C. caledonica. 66: Koumac,

stn 1318 (NC 648); 67: Toubo, stn 1259 (NC 301). Figuras 68, 69. Cystiscus minor. 68: Touho, stn

1259 (NC 302); 69: Toubo, stn 1271 (NC 372).

263

Iberus, 21 (1), 2003

opaque white. Inner mantle light yel-

lowish decorated with packed and dis-

organized small leopard-patterned

greyish blue marks. The zone under the

spire is light brown, surrounded by

light yellow.

Distribution: Only known from 2 sta-

tions off Koumac, NW of NC mainland.

Alive from 15 to 20 m.

Habitat: Not recorded.

Remarks: C. pardus sp. nov. is very

similar to C. goubini. However, both

species were collected and observed in

micro-sympatry (stn 1318) and they

display constant differences. The shell of

C. pardus is smaller and more rounded.

The foot is yellowish to opaque white

better than dirty white. The inner

mantle has a yellow ground versus a

greyish-blue one, and shows irregular

fragmented greyish blue marks versus

rounded whitish grey-blue ones. The

head decoration is identical in both

species, suggesting a close relationship.

Cystiscus deltoides sp. nov. (Fig. 52)

Type material: Holotype (Fig. 52) in MNHN. Koumac, stn 1331. 1 paratype (ad sh) from the type

locality, in MNHN.

Type locality: Koumac area, Great Reef, stn 1331, 20” 40'-20* 40.6' S, 164” 11.2'-164* 12.1” E, 55-57

m, outer slope.

Etymology: From the subtriangular outline of the shell.

Shell description (Fig. 52): Opaque

white, light, slender, subtriangular,

upper part inflated, tapering to the base.

Spire flat, upper half part of the labrum

arched, shoulder slightly elevated, lower

half of the labrum oblique, inner lip

straight and smooth. Three distinct ante-

rior columellar plaits, eight visible small

upper parietal varix. Aperture moderatly

narrowed, slightly widening to the base.

Size: 1.50 x 0.95 mm.

Animal: Unknown.

Distribution: Only known by one

lot of two shells dredged off

Koumac, NW of NC mainland. Alive

in 55 m.

Habitat: Not recorded.

Remarks: C. deltoides sp. nov. shows

some similarities with C. punctatus as

well as with the species described

hereunder, as far as shell features are

concerned. C. deltoides has however a

more triangular outline than C.puncta-

tus, and has a less slender outline and

a less elevated labrum than the follow-

ing species. C. deltoides may have a

deeper bathymetric distribution than

the other species studied herein.

Cystiscus caeruleus sp. nov. (Figs. 53, 74)

Type material: Holotype (Fig. 53, 74) in MNHN. Touho, stn 1270.

Type locality: Touho area, Great Reef Mangalia, stn 1270, 20” 45' S, 165” 16.5” E, 10-35 m, outer

slope.

Etymology: From the blue inner mantle of the animal.

Shell description (Fig. 53): Translucent

white, light, slender, subtriangular,

upper part inflated. Spire flat, upper

part of the labrum arched and much ele-

vated, central and lower labrum

straight. Three distinct anterior columel-

lar plaits, two small upper parietal

varix. Aperture much narrow, hardly

opened to the base.

264

Size: 1.7 x 1.0 mm.

Animal description (Fig. 74): Bifur-

cated head with short frontal lobes, foot

small. Head and foot orange, inner

mantle blue.

Distribution: Only known by 1 spm

from Touho, NE of NC mainland. Alive

in 10 m.

Habitat: Not recorded.

BOYER: The Cystiscidae from upper reef formations of New Caledonia

Remarks: The shell of C. caeruleus

sp. nov. shows some similarities with

C. deltoides and with the species de-

scribed hereunder. Its animal presents

a much original bicoloured chroma-

tism.

Cystiscus tricinctus sp. nov. (Figs. 54, 75)

Type material: Holotype (Fig. 54, 75) in MNHN. Touho, stn 1269. 4 paratypes (ad spm) from the

type locality, in MNHN.

Type locality: Touho area, Doiman Reef, stn 1269, 20? 35.1' S, 165” 08.1' E, 15-20 m, outer slope.

Etymology: From the tribanded decoration of the inner mantle.

Shell description (Fig. 54): Translu-

cent white, slender, light, suboval. Spire

very small, low teat-like protoconch,

upper labrum arched, shoulder moder-

ately elevated, central and lower

labrum straight, aperture narrow, faint-

ly widening to the base, three columel-

lar plaits.

Size: 1.60 x 0.95 mm.

Animal description (Fig. 75): Bifur-

cated head with short massive frontal

lobes, foot small and triangular. Head

light green, eyes red. Foot mottled

opaque white. Inner mantle yellow with

three black spiral bands, the anterior

one being U-shaped. External mantle

translucent.

Distribution: Only known from one

station in Touho, NE of NC mainland.

Alive in 15 m.

Habitat: Not recorded.

Remarks: The shell of C. tricinctus sp.

nov. shows some similarities with that

ones of C. caeruleus and of C. punctatus.

However, it is more oval and less shoul-

dered than C. caeruleus and it is more

slender than C. punctatus, with a nar-

rower aperture. The animal of C. tricinc-

tus is also much different from that one

of these two species.

Cystiscus pseudoaurantius sp. nov. (Figs. 56, 77)

Type material: Holotype (Fig. 56, 77) in MNHN. Touho, stn 1255. 1 paratype (ad spm) from the

type locality, in MNHN.

Other material studied: Expedition Montrouzier, Touho, 1993: stn 1270, 20* 45' S, 165” 16.5' E,

10-35 m, 5 ad spm, stn 1271, 20? 52.7' S, 165* 19.5' E, 5-25 m, 2 ad spm.

Expedition Montrouzier, Koumac, 1993: stn 1316, 20? 40' S, 164” 11.2 E, 12 m, 5 ad and 1 subad spm.

Type locality: Touho area, Ouao Islet, stn 1255, 20? 43' S, 165 08' E, 11 m, sand, detritic domes.

Etymology: As distinct from C. aurantius a congeneric species which has also an orange coloured animal.

Shell description (Fig. 56): Translucent

white, solid, subtriangular, wide. Spire

hardly pronounced, upper labrum

arched, shoulder slightly elevated,

lower labrum straight, inner lip straight

and smooth, aperture moderately

narrow, faintly widening to the base,

three strong anterior columellar plaits

and a tiny fourth upper one.

Size: 1.9 x 1.3 mm.

Animal description (Fig. 77): Bifur-

cated head with long frontal lobes, foot

medium sized, triangular. Head light

orange with a wide red fringe along the

central slit, frontal lobes red, eyes red.

Foot and external mantle light orange,

inner mantle beige.

Distribution: Known from Touho and

Koumac, northern NC mainland. Alive

from 5 to 12 m.

Habitat: The type material comes

from detritic sandy places.

Remarks: The shell of C. pseudoauran-

tius sp. nov. is distinct from its relatives

by its large size, wide and massive pro-

portions, subtriangular rounded outline.

The animal is distinct from C. auran-

tius by the red marks on its head and by

the deeper orange colour ground of

both head and foot.

265

Iberus, 21 (1), 2003

Figures 70, 71. Cystiscus goubini. 70: Touho, stn 1269 (NC 330); 71: Koumac, stn 1318 (NC 646). 3

Figure 72: C. pardus, holotype, Koumac, stn 1318 (NC 647).

Figuras 70, 71. Cystiscus goubini. 70: Toubo, stn 1269 (NC 330); 71: Koumac, stn 1318 (NC 646).

Figura 72: C. pardus, holotipo, Koumac, stn 1318 (NC 647).

266

BOYER: The Cystiscidae from upper reef formations of New Caledonia

A AN

EN IAS E

AA:

uN E > ¡DN

Figure 73: Cystiscus punctatus, Koumac, stn 1318 (NC 649). Figure 74: C. caeruleus, holotype, Touho,

stn 1270 (NC 390). Figure 75: C. tricinctus, holotype, Touho, stn 1269 (NC 331). Figure 76: C.

marshalli, Touho, stn 1272 (NC 348). Figure 77: C. pseudoaurantius, holotype, Touho, stn 1255 (NC

322). Figure 78: Plesiocystiscus bavayi, Touho, stn 1271 (NC 359).

Figura 73: Cystiscus punctatus, Koumac, stn 1318 (NC 649). Figura 74: C. caeruleus, holotipo, Touho,

stn 1270 (NC 390). Figura 75: C. tricinctus, holotipo, Touho, stn 1269 (NC 331). Figura 76: C. mar-

shalli, Touho, stn 1272 (NC 348). Figura 77: C. pseudoaurantius, holotipo, Touho, stn 1255 (NC 322).

Figura 78: Plesiocystiscus bavayi, Touho, stn 1271 (NC 359).

267

Iberus, 21 (1), 2003

Cystiscus cooverti sp. nov. (Fig. 57)

Type material: Holotype (Fig. 57) in MNHN. Koumac, stn 1312. 2 paratypes (ad spm) from the

type locality, in MNHN.

Other material studied: Expedition Montrouzier, Koumac, 1993: stn 1310, 20? 39.7" S, 164” 14.9

E, 15 m, 3 ad spm; stn 1318, 20* 41.4' S, 164” 14.8' E, 20-30 m, 2 ad and 1 subad spm, stn 1319, 20*

44.7' S, 164? 15.5' E, 15-20 m, 1subad spm.

Type locality: Koumac area, Pass, east cliff, stn 1312, 20? 40.4' S, 164” 14.9 E, 26-40 m, hard bottoms.

Etymology: For Gary Coovert, who devoted deep studies in marginellid gastropods during the

last twenty years, and who made a first selection work on the material studied here.

Shell description (Fig. 57): Translucent

white, light, slender oval, spire merging

into the insertion of the outer lip, high

shouldered arched upper labrum, lower

part sinuous, narrow aperture slightly

widening to the base, two anterior oblique

columellar plaits, one upper subvertical

third one.

Size: 1.40 x 0.85 mm.

Animal: Unknown.

Distribution: Only known from four

stations in Koumac, NW of NC main-

land. Alive from 15 to 26 m.

Habitat: Hard bottoms.

Remarks: Despite the unknown ani-

mal, the shell characters are sufficient

here for a distinction at the specific level,

based on the moderate size, the slender

oval ouline, the sinuous elevated labrum

and the subvertical third columellar plait.

Cystiscus bougei (Bavay, 1917) (Figs. 58, 59)

Marginella bougei Bavay, 1917, p. 103-104, pl. IL fig. 3 (uncorrect picture). Bavay, 1922, p. 58, pl. L

Figs. 6-7 (correct pictures).

Type material: Lectotype (Fig. 58) in MNHN. Labels: “Marginella (Granula) Bougei cotypes. If

Wallis”; “Marginella Bougei. l£ Wallis. types”.

The lectotype was selected by ROTH and CLOVER (1973: 209), under ICZN (74b). Five other sh

belonging to the same lot were later separated by G. Coovert as two Cystiscus ¡ota (Hedley, 1899)

and three Cystiscus sp.

One lot of 43 sh and fr was found in MDC (MNHN) with the labels: “Ma Granula Bougei 1. Wallis

Sables”, from the hand of Bavay, and “TYPE” in typed red letters. These shells are now joined to

the type lot as paralectotypes.

Other material studied: Expedition Montrouzier, Koumac, 1993: stn 1277, 20” 34' S, 164” 16' E, 0-

2 m, lad and 1 juv spm; stn 1279, 20* 35' S, 164” 15.5' E, tide, 10 ad and 4 juv spm (Fig. 59); stn

1282, 20” 33.5' S, 164” 13' E, tide, 1 ad spm; stn 1289, 20” 29.2” 5, 164? 10.2' E, tide, 23 ad and 6 juv

spm, 7 ad and 2 juv sh; stn 1292, 20” 22.4” S, 164” 06.8' E, tide, 2 ad and 1 juv sh; stn 1300, 20* 35.6"

S, 164? 15.2” E, 10-11 m, 6 ad and 1 juv spm;, stn 1303, 20? 37.7'-20* 38.8' S, 164” 15.9-164* 17.1' E,

0-8 m, 1 ad sh; -Koumac area, Rat Islet, 20? 33.7' S, 164” 11' E, tide, 20 ad and 1 juv sh.

Type locality: Ouvea, Wallis Islands.

Shell description (Fig. 58): White,

light, slender egg-shaped outline, taper-

ing to the base. Outer lip sloping down

in its upper part, thickened and bulging

in its central part, slightly flexuous in its

lower part. Long aperture, moderately

widening to the base, three oblique col-

umellar plaits.

Size: 1.4 x 1.0 mm.

Animal: Unknown. One of the spm

from the stn 1279 is labelled as corre-

268

sponding to the drawing NC 639 by S.

Gofas. However, this drawing shows an

animal with well-pronounced rounded

pustules on the external mantle, this char-

acter being common in Granulina, but not

recorded in Cystiscus. The soft parts chro-

matism pictured in NC 639 matchs the one

of a species attributable to Granulina

(BOYER, in press c). Furthermore, the foot

of the dried animal said to be pictured in

NC 639 clearly shows a festoon of black

BOYER: The Cystiscidae from upper reef formations of New Caledonia

patches, whereas such a decoration is not

represented in NC 639. So, it seems evident

that an inversion of sketches references

led to the loss of the picture of C. cf bougel.

Distribution: North West NC to

Wallis Islands. Alive from intertidal to

10 m, empty shells in intertidal. BAVAY

(1917: 103) recorded a smaller form as

ranging in Tonga Islands.

Habitat: Live spm are recorded from

soft bottoms with grass.

Remarks: C. bougel appears as common

and somewhat abundant in very shallow

waters of Koumac, whereas it seems to be

lacking in low tide brushings made off

Noumea and Touho. However, tide sam-

plings seem to have been more frequent

in Koumac. The shell variability observed

in Koumac overlaps in the main the shell

variability observed in Wallis.

The distribution of a species of Cys-

tiscus from northern NC to Wallis and

Tonga cannot be considered as impossi-

ble, even if the occurrence of the species

in eastern NC, in Loyalty and in the

New Hebrids remains to be verified.

The control of the animal chromatism in

the different places would help greatly

to precise the taxonomic status of the

populations attributable to C. bougel.

Genus Plesiocystiscus Coovert and Coovert, 1995

Type species: Marginella jewetti Carpenter, 1857, by original designation.

Plesiocystiscus tomlini (Bavay, 1917) (Fig. 60)

Marginella tomlini, Bavay, 1917, p. 102-103, pl. IL fig. 7.

Type material: Lectotype (Fig. 60) and 3 paralectotypes (3 ad sh) in MNHN (ex-MDOC). Labels:

“Marginella Tomlini By Ouvea. I. Wallis, sables”, from the hand of Bavay, and “TYPE” in typed

red letters. No type material was found in MNHN by Clover (ROTH and CLOVER, 1973: 214).

Other material studied:

Expedition Montrouzier, Koumac, 1993: stn 1312, 20” 40.4” S, 164” 14.9 E, 26-40 m, 1 ad sh; stn

1318, 20? 41.4' S, 164” 14.8' E, 20-30 m, 1 ad sh.

Type locality: Ouvea, Wallis Islands.

Shell description (Fig. 60): White,

solid, slender egg-shaped outline, spire

short and faintly bulging, outer lip

straight, aperture long and narrow,

moderately widening to the base, three

much oblique columellar plaits.

LEIZZOX TZ DA:

Animal: Unknown.

Distribution: Only known from

Wallis Islands (type locality) and by the

two comparable shells from Koumac,

NW of NC mainland, sampled as empty

shells from 20 to 26 m.

Habitat: Unknown.

Remarks: The two shells sampled

off Koumac are slightly larger (L: 2.4

and 2.5 mm) than the types of P.

tomlini, their columellar border is

slightly more convex, and overall, they

both present a small fourth columellar

plait whereas the four types do not

have.

Despite the fact that all the other

shell features are similar in both

phenae, they are probably distinct at a

specific level.

Plesiocystiscus bavayi sp. nov. (Figs. 61, 62, 78)

Type material: Holotype (Fig. 61) in MNAHN. Touho, stn 1261. 2 paratypes (1 subad spm and 1

ad sh) from the type locality, in MNHN.

Other material studied: Expedition Montrouzier, Touho, 1993: stn 1271, 20” 52.7' S, 165” 19.5' E,

5-25 m, 1 subad spm (Figs. 62, 78).

269

Iberus, 21 (1), 2003

Expedition Montrouzier, Koumac, 1993: stn 1311, 20” 40.4” S, 164” 14.9 E, 10-60 m, 1 ad spm;, stn

1314, 20? 39.8' S, 164” 15.3” E, 30-63 m, 2 ad sh; stn 1318, 20? 41.4' S, 164” 14.8' E, 20-30 m, 3 ad

spm; stn 1319, 20? 44.7' S, 164” 15.5' E, 15-20 m, 1 ad spm, stn 1322, 20? 45.2' S, 164? 15.2 E, 53-71

m, 2 ad and 1 subad sh; stn 1331, 20? 40'-20* 40.6' S, 164” 11.2'-164* 12.1” E, 55-57 m, 2 ad sh.

Expedition Lifou 2000: stn 1442, 20” 46.4” S, 167” 02.0' E, 47 m, 1 ad sh; stn 1449, 20? 45.8' S, 167"

01.65 E, 17 m, 1 ad sh.

Type locality: Touho area, channel, stn 1261, 20” 46'-20* 47" S, 165* 15'-165* 16.5' E, 45-56 m,

detritic sand.

Etymology: For Arthur Bavay (1840-1923), french conchologist who specially devoted to the

study of small marginelliform gastropods in the earl XX'* Century, and who performed the first

revision of cystiscids from New Caledonia.

Shell description (Fig. 61): White,

slender tear-shaped, upper part inflated,

spire very small, rounded, aperture

narrow, slightly widening to the base,

inner lip long, straight and smooth, four

distinct oblique columellar plaits, a fifth

subvertical one is suggested.

Size: 3.30 x 1.95 mm.

Animal description (Fig. 78): Bilobed

head, with two long tentacles, black

eyes at their base on a swollen peduncle.

Siphon wide and produced. Foot large

and subrectangular. The back tip of the

metapodium is truncated. Mentum dis-

tinct ahead of the foot.

Head, siphon and foot whitish

translucent. A small orange stain and a

black one at the level of the eye, a

CONCLUSION

In the course of this study, 28

morphs were recognized as belonging to

the sublittoral Cystiscidae from NC

mainland. Among them, 5 are referred

to species previously described (one

being referred with much reserves), 18

are described and named as new

species, and 5 are recorded as potential

new species. As such, the known diver-

sity of the sublittoral Cystiscidae from

NC is increased by a factor higher than

xD:

Nineteen of these 28 morphs belong

to the group Crithe /Cystiscus. lt can be

assumed that much more species

belonging to this group remain to be

discovered at infralittoral levels from

NC, by the fact that most of the species

are represented by a very limited

number of lots and subjects within the

270

yellow patch at the centre of the siphon.

Inner mantle light reddish with three

spiral rings of yellow dots.

Distribution: Known from Touho and

Koumac, northern NC mainland. Alive

from 5 to 45 m, empty shells from 17 to

59 m.

Habitat: Detritic to silty sand as well

as hard bottoms.

Remarks: One larger shell from

Koumac (stn 1322, 53-71 m), sizing 3.75

mm in length, shows noticeably distinct

features (more slender and cylindric out-

line, less opened aperture to the base,

more concave first columellar plait, more

produced spire), and must be provision-

ally considered as belonging to a differ-

ent species, as P. sp. aff. bavay,.

important material checked here. As far

as the group Crithe/Cystiscus is con-

cerned, eight morphs are represented by

only one lot (six morphs by only one or

two subjects), three morphs by only two

lots, and only eight morphs among

nineteen are represented by more than

two lots.

In other words, the majority of the

morphs belonging to Crithe / Cystiscus

are represented by only one or two lots,

and 30% of the total morphs are repre-

sented by only one or two subjects.

From these elements, it can be inferred

that the “sampling saturation” is far to

be reached, and that deeper investiga-

tions within the sampled areas will very

probably procure many new morphs.

The species composition does not

show any evident correspondence with

BOYER: The Cystiscidae from upper reef formations of New Caledonia

the special environmental features rep-

resented within the four studied sites,

and the results suggest that the

endemism is very developed in NC

Crithe/Cystiscus: among the nineteen

studied morphs, ten are represented

only from northern NC, six only from

SW NC, and only three are represented

in northern as well as southern stations

(overlapping rate of only 16%). Even if it

can be supposed that the number of

widespread morphs is underestimated,

the results allow to infer that some of

the species may have a distribution

restricted to few hundred kilometres of

coast. As a consequence, each of the

wide coastal zones which were not

intensively checked (central West Coast,

central and southern East Coast, Ile des

Pins, far north lagoon and Belep Island)

may shelter a set of endemic species.

In these conditions, the diversity of

Crithe/Cystiscus displayed in this article

cannot be considered as representative

of the real diversity of this group in NC

waters, which is probably much higher.

It must be only considered as an indica-

tor demonstrating that the group

Crithe/Cystiscus is highly diversified in

NC waters, and as a stimulating pretext

to develop intensive samplings in other

places of the NC coast, to intensify

locally the sampling for a measure of

effort /diversity output, and to correlate

the faunal diversity at hand with the

diversity of its sheltering micro-habitats

in view to infer the real faunistic diver-

sity.

The best use of the observations

about the diversity is probably to play it

as an argument in the reconstruction of

an evolutionary history or as explana-

tory of a biogeographic context: this

supposes appropriate ways and degrees

of observations. In the case studied here,

the principal deficiencies of the informa-

tion seem to lie in the fact that micro-

habitats attached to each species are not

recorded (assessment of the sampling

impact correlated to the micro-habitat

diversity) and in the relative few

number of live animal pictures com-

pared to the high number of specimens

collected (assessment of the global vari-

ability of the species and control of the

specific diversity). In the case of Cystis-

cus/Crithe, made of numerous species

with very similar shells but with very

different and non-variable animals, the

knowledge of the animal chromatism

and external morphology is decisive for

estimating the diversity at the specific

level. In the case of the small white

shelled Gibberula, that is the important

variability of the shells and of the

animals of some species which makes

necessary to have recourse to the sys-

tematic observation of the live animals.

Consequently, the best exploitability of

the intensive samplings seems to require

the notation in the field of precise data

about the sampled micro-habitats, and

the organization of intensive picturing

of live animals.

The diversity of Crithe/Cystiscus

observed in NC cannot be considered no

more as a special occurrence restric-

tively attached to this area. In fact, the

situation revealed in NC may as well be

representative of a high diversity of

Crithe/Cystiscus in Central West Pacific.

The intensive samplings realized

recently in NC constitute an innovation,

and the rest of the Central West Pacific

(even the intertropical australian coasts)

waits for such a performance.

On the ground of the various

morphs studied here, the genera Crithe

and Cystiscus do not show as clearly dis-

tinct. It better seems that they both

belong to a wide complex of forms

which deserves to be unified under a

common taxonomic group. However,

this wide group will probably require

taxonomic subdivisions corresponding

to natural affinities, and it is conceivable

that the generic unification of the group

will be more correctly stated in the

frame of a general reorganization ren-

dering the diversity pattern working

here.

ACKNOWLEDGEMENTS

Thanks are due to the members of

the Expeditions Montrouzier 1993 and

Lifou 2000 field parties, and in particu-

271

Iberus, 21 (1), 2003

lar to Philippe Bouchet (Paris Museum)

and to Serge Gofas (University of

Malaga) for performing accurate obser-

vations and precious sketches about live

animals in the field.

I am also indebted to Yves Finet

(Geneva Museum) for the pictures of

Gibberula pulchella syntypes, to Philippe

Maestrati (Paris Museum) for the cre-

ation of the digital plates, and to Robert

BIBLIOGRAPHY

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sables littoraux de divers pays. Journal de

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BAVaY, A., 1922. Coquilles des sables littoraux

marins. Journal de Conchyliologie, 64 (1): 57-

65, pl. 1.

BOUcHET, P., 1994. Atelier Biodiversité

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novembre 1993. Rapports de missions,

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BOYER, F., in press a. Le groupe Gibberula fru-

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Mascarene Islands. Journal of Conchology.

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dans l'infralittoral de la Nouvelle

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and Nicole Hasselot (Jouques, France)

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Guido Poppe (Antwerpen, Belgium)

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A special acknowledgement is due to

Maxime Desjardins (Paris, France), who

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lection in Paris Museum, allowing to revise

several taxa in appropriate conditions.

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19 (3): 201-206.

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de Conchyliologie, 20 (3): 218-222, pls 2-13.

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(Marginellidae) en Méditerranée. Lavori

della Societa Italiana di Malacología, 23: 113-

139.

HEDLEY, C., 1899. The Mollusca of Funafuti,

Part I: Gasteropoda. Memoirs of Australian

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fils, Paris.

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of the Marginellidae described by Bavay,

1903-1922. The Veliger, 16 (2): 207-215.

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Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W. EF.

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lo considera conveniente, podrá solicitar a los autores otras modificaciones que considere oportunas. Si el trabajo es

aceptado, el autor deberá enviar una copia impresa del mismo corregida, acompañada por una versión en disco flexi-

ble (diskette), utilizando procesadores de texto en sus versiones de DOS o Macintosh. La fecha de aceptación figura-

rá en el artículo publicado.

* Las pruebas de imprenta serán enviadas al autor responsable, EXCLUSIVAMENTE para la corrección de erratas, y

deberán ser devueltas en un plazo máximo de 15 días. Se recomienda prestar especial atención en la corrección de las

pruebas.

e De cada trabajo se entregarán gratuitamente 50 separatas. Aquellos autores que deseen un número mayor, deberán

hacerlo constar al devolver las pruebas de imprenta, y NUNCA POSTERIORMENTE. El coste de las separatas adi-

cionales será cargado al autor.

INSTRUCTIONS TO AUTHORS

* Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. Papers are manus-

cripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. Monographs should exceed

50 pages of the final periodical, and will be published as Supplements. Authors wishing to publish monographs should

contact the Editor. Manuscripts are considered on the understanding that their contents have not appeared or will not

appeared, elsewhere in substantially the same or any abbreviated form.

+ Manuscripts and correspondence regarding editorial matters must be sent to: D. Gonzalo Rodríguez Casero, Editor

de Publicaciones, Apartado 156, 33600 Mieres, Asturias, Spain.

e Manuscripts may be written in any modern language.

* When a paper exceeds 20 pages, extra pages will be charged to the author(s) at full cost.

+ Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side on A-4

(297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted. When a paper has joint

authorship, one author must accept responsability for all correspondence.

* The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate.

* Papers should conform the following layout:

First page. Vhis must include a concise but informative title, with mention of family of higher taxon when appropriatte,

and its Spanish translation. It will be followed by all authors” names and surnames, their full adress(es), an abstract (and

its Spanish translation) not exceeding 200 words which summarizes not only contents but results and conclusions, and

a list of Key Words (and their Spanish translation) under which the article should be indexed.

Following pages. These should content the rest of the paper, divided into sections under short headings. Whenever pos-

sible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, Conclusions,

Acknowledgements and References. Unusual abbreviations used in the text must be grouped in one alphabetic sequence

after the Material and methods section.

* Notes should follow the same layout, without the abstract.

* Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical Nomencla-

ture must be strictly followed. The first mention in the text of any taxon must be followed by its authority including

the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN FULL, including the perio-

dical, in an abbreviate form, where they were described, and the type localities in square brackets when known. Follow

this example (please note the punctuation):

Dendrodoris limbata (Cuvier, 1804)

Synonyms

Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Type locality: Marseille].

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275.

These references must not be included in the Bibliography list, except if referred to elsewhere in the text. I£a full list

of references of the taxon is to be given immediately below it, the same layout should be followed (also excluding those

nowhere else cited from the Bibliography list).

Only Latin words and names of genera and species should be underlined once or be given in ¿talics. No word must

be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In Spanish

manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper comma (/).

» References in the text should be written in small letters or SMALL CAPITALS: Fretter and Graham (1962) or FRETTER

AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must include all of them

[Smith, Jones 82 Brown (1970)], thereafter use et al. [Smith et al. (1970)]. Ifan author has published more than one

paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit.

The references in the reference list should be in alphabetical order and include all the publications cited in the text but

only these. ALL the authors of a paper must be included. These should be written in small letters or SMALL CAPITALS.

The references need not be cited when the author and date are given only as authority for a taxonomic name. Titles of

periodicals must be given IN FULL, not abbreviated. For books, give the title, name of publisher, place of publication,

indication of edition if not the first and total number of pages. Keep references to doctoral theses or any other unpu-

blished documents to an absolute minimum. See the following examples (please note the punctuation):

Fretter, V. and Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.

Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder, W. F. (Ed.):

Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166.

Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Miscelá-

nea Zoológica, 3 (5): 21-51.

. Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size

when designing figures. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high,

or be proportional to these sizes. Two columns format is recomended. It is desirable to print figures with their legend

below, so authors are asked to take this into account when preparing full page figures. If computer generated graphics

are to be included, they must be printed on high quality white paper with a laser printer. Photographs must be of good

contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are of uni-

form width. Remember that grouping photographs of varied contrast results in poor reproduction. Take account of

necessary reduction in lettering drawings; final lettering must be at least 2 mm high. In composite drawings, each figure

should be given a capital letter; additional lettering should be in lower-case letters. A scale line is recomended to indi-

cate size, magnification ratio must be avoided as it may be changed during printing. UTM maps are to be used ¡if neces-

sary. Figures must be submitted on separate sheets, and numbered with consecutive Arabic numbess (1, 2, 3,...), without

separating “Plates” and “Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and an Spa-

nish translation must be included. Follow this example (please note the punctuation):

Figure 1. Neodoris carví. A: animal crawling; B: rinophore; C: gills.

If abbreviations are to be used in illustrations, group them alphabetically after the Legends for Figures section.

Authors wishing to publish illustrations in colour will be charged with additional costs (30,000 ptas, 180 euros per

page). They should be submitted in the same way that black and white prints.

If the authors want to send Figures in digital format, CONTACT the Editor first.

* Tables must be numbered with Roman numbers (L, II, IL...) and each typed on a separate sheet. Headings should

be typed on a separate sheet, together with their English translation. Complex tables should be avoided. As a general

rule, keep the number and extension of illustrations and tables as reduced as possible.

+ Manuscripts that do not conform to these instructions will be returned for correction before reviewing.

+ Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the date the

manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two referees. Based of these

evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors will receive a copy of the refe-

rees* comments. If a manuscript is accepted, the Editorial Board may indicate additional changes if desirable. Accep-

table manuscripts will be returned to the author for consideration of comments and criticism; a finalized manuscript

must then be returned to the Editor, together with a floppy disk containing the article written with a DOS or Macin-

tosh word processor. Dates of reception and acceptance of the manuscript will appear in all published articles.

* Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special

attention to references and their dates in the text and the Bibliography section, and also to numbers of Figures and

Tables appearing in the text.

e Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs are

returned, and will be charged at cost. NO LATER orders will be accepted.

LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Junta Directiva desde el 14 de noviembre de 2000

Presidente Emilio Rolán Mosquera

Vicepresidente Diego Moreno Lampreave

Secretario Luis Murillo Guillén

Tesorero Jorge J. Otero Schmitt

Avda. de las Ciencias s/n, Campus Universitario, 15706 Santiago

de Compostela, España

Editor de Publicaciones Gonzalo Rodríguez Casero

Apartado 156, 33600, Mieres del Camino, Asturias, España

Bibliotecario Rafael Araujo Armero

Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez

Abascal 2, 28006 Madrid, España

Vocales Ramon M. Álvarez Halcon

Benjamín Gómez Moliner

Eugenia María Martínez Cueto-Felgueroso

Jesús Souza Troncoso

José Templado González

La Sociedad Española de Malacología se fundó el 21 de agosto de 1980. La sociedad se registró como una aso-

ciación sin ánimo de lucro en Madrid (Registro N* 4053) con unos estatutos que fueron aprobados el 12 de

diciembre de 1980. Esta sociedad se constituye con el fin de fomentar y difundir los estudios malacológicos

mediante reuniones y publicaciones. A esta sociedad puede pertenecer cualquier persona o institución interesada

en el estudio de los moluscos.

SEDE SOCIAL: Museo Nacional de Ciencias Naturales, c/ José Gutierrez Abascal 2, 28006 Madrid, España.

CUOTAS PARA 2003:

Socio numerario (en España): 33,06 euros

(en Europa) 36,06 euros

(fuera de Europa): 42,07 euros

Socio estudiante (en España): 18,03 euros

(en el extranjero): 24,04 euros

Socio Familiar: (sin recepcion de revista) 3 euros

Socio Protector: (mínimo) 42,07 euros

Socio Corporativo (en Europa): 42,07 euros

(fuera de Europa): 48,08 euros

INSCRIPCIÓN: 6,01 euros, además de la cuota correspondiente.

A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al

Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la

sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque-

lios socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins-

cripción se enviarán junto con el abono de una cuota anual al Tesorero.

Membe:ss living in foreing countries can deduce 6 euros if paid before 15 April.

Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que

se publiquen.

2 MAA

Iberus 21 (1) 2003

SNYDER, M. A. Four new species of Latirus (Gastropoda: Fasciolariidae) from the Philippine

Islands and the southern Caribbean

Cuatro nuevas especies de Latirus (Gastropoda: Fasciolarizáae) de Filipinas y el Caribe suriiod9

ÁviLa, S. P. The littoral molluscs (Gastropoda, Bivalvia and Polyplacophora) of Sáo Vicente,

Capelas (Sáo Miguel Island, Azores): ecology and biological associations to algae

Los moluscos litorales (Gastropoda, Bivalvia y Polyplacophora) de Sáo Vicente, Capelas (Sáo

Miguel Island, Azores): ecología y asociaciones biológicas con AÍgAS cc 11-33

La PERNA, R., LANDAU, B. AND SILVA, C. M. Da. The genus Granulina (Gastropoda, Marginelli-

dae) from the Atlantic Iberian Pliocene with description of a new species from Portugal

El género Granulina (Gastropoda, Marginellidae) en el Plioceno Atlántico Ibérico y descripción

de una especie nueva para Portugal 0 AN 35-42

GARCÍA-ÁLVAREZ, O. AND URGORRI, V. Solenogastres molluscs from the BENTART Collection

(South Shetland Islands, Antarctica), with a description of a new species

Moluscos Solenogastres de la Colección BENTART (Islas Shetland del Sur, Antártida), con la

descripción de una nueva especie a o a 43-56

SORIANO, S., SÁNCHEZ LIZASO, J. L. AND GUERRA, A. Cephalopod species collected in the upper

continental slope off Alicante (Western Mediterranean)

Especies de cefalópodos capturadas en el talud superior de Alicante (Mediterráneo occiden-

O 57-66

SALVINI-PLAWEN, L. VON. On the phylogenetic significance of the aplacophoran Mollusca

Sobre la significación filogenética de los moluscos aplacoforos Lino ns 67-97

ROLÁN, E. AND RYALL, P. The genus Pelycidion (Mollusca: Archaegastropoda) in West Africa

El género Pelycidion (Mollusca: Archaegastropoda) en Africa occidental ............. 99-104

AMOR PÉREZ, M2 J., RAMÓN, M. Y DURFORT 1 COLL, M. El conducto excretor de la glándula de la cáp-

sula de Bolinus brandaris (Gastropoda Prosobranchia): estudio estructural y ultraestructural

Structural and ultrastructural study of the excretor duct of the capsule gland of Bolinus branda-

ris (Gastropoda, Prosobranchta) 0 e 105-113

ROLÁN, E. MONTEIRO, A. AND FRAUSSEN, K. Four new Euthria (Mollusca, Buccinidae) from the

Cape Verde archipelago, with comments on the validity of the genus

Cuatro nuevas Euthria (Mollusca, Buccinidae) del archipiélago de Cabo Verde con comentarios

sobre la validez del género 0 a ds daa NN 115-127

MORENO, J. A., PEÑAS, A. Y ROLÁN, E. Contribución al conocimiento de la fauna malacológica

miocénica de La Pedrera, Sant Llorenc d'Hortons (NE de la Península Ibérica)

Contribution to the knowledge of the Miocenic malacological fauna from La Pedrera, Sant

Lloreng d'Hortons (NE of the Iberian Peninsula) 129-175

PEÑAS, A. Y GIRIBET, G. Adiciones a la fauna malacologica del litoral del Garraf (NE de la Penín-

sula Ibérica)

Additions to the malacological fauna of El Garraf (NE of the Iberian Peninsula) .... 177-189

ROLAN, E. Y MARTÍNEZ-ORTÍ, A. Nuevas especies de la familia Hydrobiidae (Mollusca, Orthogas-

tropoda) de la Comunidad Valenciana (España)

New species of the family Hydrobiidae (Mollusca, Orthogastropoda) from “Comunidad Valen-

CARA [SPA A 191-206 -

HADORN, R. AND FRAUSSEN, K. The deep-water Indo-Pacific radiation of Fusinus (Chryseofusus

subgen. nov.) (Gastropoda: Fasciolariidae)

La radiación Indo-Pacífica de aguas profundas del género Fusinus (Chryseofusus subgen. nov.)

(Gastropoda: Eaciólarida o 207-240

BOYER, E The Cystiscidae (Caenogastropoda) from upper reef formations of New Caledonia. >

Los Cystiscidae (Caenogastropoda) de las formaciones superiores de coral de Nueva Caledo-

A 241-272

ISSN 0212-3010