Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Comité DE REDACCIÓN (BOARD OF EDITORS)

EDITOR DE PUBLICACIONES (EDITOR-IN=CHIEF)

Serge Gofas

Universidad de Málaga, España

DIRECTOR DE REDACCIÓN (EXECUTIVE EDITOR)

Gonzalo Rodríguez Casero

Apdo. 156, Mieres del Camino, Asturias, España

EDITORA EJECUTIVA (MANAGING EDITOR)

Eugenia M* Martínez Cueto-Felgueroso

Apdo. 156, Mieres del Camino, Asturias, España

EDITORES ADJUNTOS (ÁSSOCIATE EDITORS)

Francisco Javier Conde de Saro

Benjamín Gómez Moliner

Ángel Antonio Luque del Villar

Emilio Rolán Mosquero

José Templado González

Jesús S. Troncoso

Embajada de España, Japón

Universidad del País Vasco, Vitoria, España

Universidad Autónoma de Madrid, Madrid, España

Universidad de Vigo, Vigo, España

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Universidad de Vigo, Vigo, España

ComiTÉ EDITORIAL (BOARD OF REVIEWERS)

Kepa Altonaga Sustacha

Eduardo Angulo Pinedo

Rafael Araujo Armero

Thierry Backeljau

Ridiger Bieler

Sigurd v. Boletzky

Jose Castillejo Murillo

Karl Edlinger

Antonio M. de Frias Martins

José Carlos García Gómez

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Edmund Gittenberger

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Yuri |. Kantor

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- Takashi Okutani

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Paul 6. Rodhouse

Joandoménec Ros ¡ Aragones

María Carmen Salas Casanovas

Gerhard Steiner

Victoriano Urgorri Carrasco

Anders Warén

PORTADA DE Jberus

Universidad del País Vasco, Bilbao, España

Museo Nacional de Ciencias Naturales, Madrid, España

Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

The Field Museum, Chicago, Estados Unidos

Laboratoire Arago, BanyulsssurMer, Francia

Universidad de Santiago de Compostela, Santiago de Compostela, España

Naturhistorisches Museum Wien, Viena, Austria

Universidade dos Acores, Acores, Portugal

Universidad de Sevilla, Sevilla, España

Harvard University, EE.UU.

National Natuurhistorisch Museum, Leiden, Holanda

Instituto de Investigaciones Marinas, CSIC, Vigo, España

Zoologische Staatssammlung Múnchen, Múnchen, Alemania

A.N. Severtzov Institute of Ecology and Evolution, Moscú, Rusia

Estación Agrícola Experimental, CSIC, León, España

Universidad de Barcelona, Barcelona, España

Dalhousie University, Halifax, Canada

Nihon University, Fujisawa City, Japón

Universitá di Roma “La Sapienza”, Roma, Italia

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

Australian Museum, Sydney, Australia

Universidad del País Vasco, Bilbao, España

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Instituto de Investigaciones Marinas, CSIC, Vigo, España

British Antarctic Survey, Cambridge, Reino Unido

Universidad de Barcelona, Barcelona, España

Universidad de Málaga, Málaga, España

Institut fir Zoologie der Universitút Wien, Viena, Austria

Universidad de Santiago de Compostela, Santiago de Compostela, España

Swedish Museum of Natural History, Estocolmo, Suecia

Iberus gualterianus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da

nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”.

Iberus

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

| CENTASO A

Ú MAR 1 6 2006

SUBRARIES_A

Vol. 23 (2) Oviedo, diciembre 2005

Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Iberus publica trabajos que traten sobre cualquier aspecto relacionado con la Malacología. Se

admiten también notas breves. /berus edita un volumen anual que se compone de dos o más números.

INSTRUCCIONES PARA LOS AUTORES

Los manuscritos deben remitirse a: Serge Gofas, Editor de Publicaciones, Departamento de Bio-

logía Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, España.

Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer

cuidadosamente las normas de publicación que se incluyen en cada número de la revista.

SUBCRIPCIONES

Iberus puede recibirse siendo socio de la Sociedad Española de Malacología, en cualquiera de sus

formas, o mediante intercambio. Aquellos socios que deseen adquirir números atrasados deberán diri-

girse al bibliotecario.

Los no socios deberán ponerse en contacto con BACKHUYS PUBLISHERS, PO. Box 321,

2300 AH Leiden, The Netherlands. Tel.: +31-71-51 70 208, Fax: +31-71-51 71 856, Correo Elec-

trónico: backhuysCeuronet.nl

Los resumenes de los artículos editados en esta revista se publican en Aquatic Science

and Fisheries Abstracts (ASFA) y en el Zoological Records, BIOSIS.

Contents list published in Aquatic Science and Fisheries Abstracts and Zoological Records,

BIOSIS.

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

O Sociedad Española de Malacología

Iberus, 23 (2): 1-6, 2005

A new species of Lodderena (Gastropoda: Skeneidae) from

the Bahamas

Una nueva especie de Lodderena (Gastropoda: Skeneidae) de las

Bahamas

Colin REDFERN* and Emilio ROLÁN**

Recibido el 26-1-2005. Aceptado el 30-VI-2005

ABSTRACT

A new species in the genus Lodderena (Gastropoda: Skeneidae) is described from the

Bahamas. The new species is compared to L. ornata (Olsson and McGinty, 1958).

RESUMEN

Se describe una nueva especie del género Lodderena (Gastropoda: Skeneidae) de las

Bahamas. La nueva especie se compara con L. ornata (Olsson and McGinty, 1958).

KEY WORDS: Skeneidae, Lodderena, Bahamas, new species.

PALABRAS CLAVE: Skeneidae, Lodderena, Bahamas, nueva especie.

INTRODUCTION

OLSSON AND MCGINTY (1958)

described Pachystremiscus as a subgenus

of Cyclostremiscus Pilsbry and Olsson,

1945. Included were two new species

described from the Caribbean coast of

Panama: C. ornatus and C. pulchellus, the

type species of Pachystremiscus. KAY

(1979) figured two species from the

Pacific. MOOLENBEEK (1996) discussed

the genus Lodderena Iredale, 1924, which

is considered the correct genus name for

this group of species, mentioning most

of the existing taxa and describing two

new species.

RUBIO, ROLÁN AND REDFERN (1998)

studied species of Lodderena occurring in

the Caribbean, including some speci-

mens from other areas that suggested a

wider distribution. They also described

* 7475 Estrella Circle, Boca Raton, Fl 33433, USA.

one new species, L. janetmayae from

Abaco, Bahamas.

In recent years the senior author has

continued to examine sediment samples

from the Bahamas. One of these con-

tained examples of the three known

Caribbean species together with exam-

ples of what appeared to be a fourth,

undescribed species. The study of this

material is the subject of this paper,

which includes the description of a new

species that is morphologically close to

L. ornata.

MATERIAL AND METHODS

This study is based on a 0.4 liter

sample of sediment collected at a depth

** Museo de Historia Natural, Campus Universitario Sur. 15782 Santiago de Compostela. e-mail: emilioro-

lanGinicia.es

Iberus, 23 (2), 2005

of 6 m at Lynyard Cay, Abaco, Ba-

hamas.

Abbreviations:

AMNH American Museum of Natural

History, New York

ANSP Academy of Natural Sciences,

Philadelphia

BMNH The Natural History Museum,

London

RESULTS

BMSM Bailey-Matthews Shell Museum,

Sanibel, Florida, USA

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Museum Nationale d'Histoire

Naturelle, Paris

ZMB Zoologisches Museum, Berlin

CCR Collection of Colin Redfern, Boca

Raton, Florida, USA

CER Collection of Emilio Rolán, Vigo

Family SKENEIDAE Clark, 1851

Genus Lodderena Iredale, 1924

Lodderena bunnelli n. sp. (Figs. 1-9)

Type material: Holotype (Figs. 1, 9), 0.56 mm, deposited at ANSP. One paratype deposited in each

of the following collections: MNCN (15.05 /46685) (Figs. 2, 8), 0.64 mm; MNHN (Fig. 3), 0.56 mm;

AMNH (Figs. 4, 7), 0.58 mm; BMNH (Fig. 5), 0.56 mm; ZMB (Fig. 6), 0.60 mm; BMSM. 3 paratypes

in CER and 21 in CCR. All type material consists of empty shells collected from the type locality in

August, 2004.

Type locality: Lynyard Cay, Abaco, Bahamas in 6 m.

Etymology: Named for Rodger R. Bunnell, who collected the sediment sample that contained the

type material.

Description: Shell (Figs. 1-6) very

small, rounded, dorsally planispiral,

whitish. Protoconch (Figs. 8, 9) approxi-

mately 200 um in diameter, with about

one whorl, nucleus rounded, surface

roughened, with 1-2 spiral lines of irreg-

ular granules. Separation from the teleo-

conch is rather well defined. Teleoconch

with only one whorl, sometimes a little

less. Sculpture is formed dorsally by

irregular undulating axial ribs that are

well separated initially but become nar-

rower, more numerous and crowded as

the whorl develops. The beginning of

the peripheral curvature is marked by a

narrow spiral cord, above which a series

of fine spiral threads starts to form

when the whorl is about half developed.

The strengthening spiral sculpture, sub-

sequently consisting of up to 9-10

threads, causes the axial ribs to shorten,

but they lengthen again just behind the

aperture. Sculpture on the rounded

periphery of the shell initially consists

of three weak, irregular spiral cords that

are connected by axial threads of similar

strength to form a net-like pattern of

ovoid or rectangular shapes (Fig. 7).

After the first half of the whorl this

pattern is replaced by spiral threads,

one of which is a little stronger and

sometimes forms a peripheral angula-

tion. Ventrally the sculpture is very

similar, but with the addition of short,

widely-spaced axial riblets adjacent to

the lower peripheral cord. Occasional

shells have a spiral row of nodules close

to the suture.

Aperture regularly rounded, with a

double peristome.

Microsculpture: Under high magnifi-

cation, small granules can be seen in the

peripheral area (Fig. 7), with very fine

axial lines elsewhere.

Dimensions: the holotype is 0.56 mm

in maximum diameter, and the largest

specimen studied is 0.64 mm.

Distribution: Lodderena ornata and L.

pulchella are widely distributed in the

Caribbean area and it is likely that L.

bunnelli n. sp. has a comparable range,

as indicated by the similarity of a shell

from the U. S. Virgin Islands figured as

Lodderena sp. by RUBIO ET AL. (1998, fig.

REDFERN AND ROLÁN: A new species of Lodderena from the Bahamas

Figures 1-9. Lodderena bunnelli n. sp., from Abaco, Bahamas (CCR). 1: holotype, 0.56 mm

(ANSP); 2: paratype, 0.64 mm (MNCN); 3: paratype, 0.56 mm (MNHN); 4-6: paratypes, 0.58,

0.56, 0.60 mm (AMNH, BMNH, ZMB); 7: microsculpture, paratype (AMNH); 8: protoconch,

paratype (MNCN); 9: protoconch, holotype (ANSP).

Figuras 1-9. Lodderena bunnelli, n. sp., de Abaco, Bahamas (CCR). 1: holotipo, 0,56 mm (ANSP); 2:

paratipo, 0,64 mm (MNCN); 3: paratipo, 0,56 mm (MNHN); 4-6: paratipos, 0,58, 0,56, 0,60 mm

(AMNH, BMNH, ZMB); 7: microescultura, paratipo (AMNH); 8: protoconcha, paratipo (MNCN);

protoconcha, holotipo (ANSP).

Iberus, 23 (2), 2005

Table I. Differences between Lodderena ornata and Lodderena bunnelli n. sp.

Tabla I. Diferencias entre Lodderena ornata y Lodderena bunnelli ». sp.

[. ornata

larger diam. between 0.66 — 0.92 mm

L. bunnelli n. sp.

smaller diam. between 0.56 — 0.64 mm

Dimensions n= 10 median= 0.78 mm n=10 median= 0.58 mm

Sp shell dorsally planispiral; periphery meets shell dorsally planispiral; periphery

e aperture near the centre meets aperture near upper dorsal border

Teleoconch l whorl + 1/8 1 whorl or a little less

3 very prominent nodulose Usually no nodulose cords; occasionally

Spiral sculpture :

cords on each side a small one ventrally close to the suture

Spiral sculpture at the periphery

three prominent cords; sometimes with

three weak, irregular cords connected by

(near beginning of teleoconch whorl) axial threads forming irregular rectangles — axial threads forming ovoid or rectangular shapes

Spiral sculpture at the periphery

(towards end of teleoconch whorl)

Thickness of outer lip

Nucleus of the protoconch

Microsculpture of the protoconch

Microsculpture of the teleoconch

fine

near the suture

41). The existence of L. bunnelli n. sp.

may have been overlooked elsewhere in

the Caribbean due to its small size and

possible confusion with L. ornata.

Remarks: The sediment sample that

contained the type material of L. bunnelli

n. sp. also yielded 12 examples of L.

janetmayae, 20 of L. pulchella and 486 of

E. ornata. Study of this material clearly

showed that there was no intergradation

between the four species.

MAYR (1963) stated that when two

different morphs are found sympatri-

cally without any intergradation

between them, they represent two dif-

ferent species. Problems for the taxono-

mist occur when two different morphs

share similar main characters. In the

present case the distinction between

Lodderena bunnelli n. sp. and L. ornata

(Figs. 10-14), the closest species, was

based on several different characters,

and importance was also attached to the

three prominent, laterally nodulose cords;

intermediate spiral threads only at the end

thicker, about a third of

the diameter of the aperture

slightly ovoid

irregular, mainly near the periphery

rarely with cords; numerous spiral

lines on a convex curvature

thinner, about a quarter of

the diameter of the aperture

rounded

some irregular granules spirally aligned

finer

lack of any intergradation between these

two sympatric species. Comparison of

the two species showed us a number of

differences, as summarized in Table 1.

Another similar species is Lodderena

striata (Kay, 1979) from Hawaii, but this

species is larger (up to 1 mm), and the

periphery ot the last whorl has only

spiral striae, lacking the net-like pattern

that is characteristic of L. bunnelli n. sp.

Records of L. ornata in RUBIO ET AL.

(1998) from the Bahamas, Cuba, Cape

Verde and Sáo Tomé, together with those

in the present work from the Bahamas,

Sáo Tomé and Easter Island, show that

this species probably has a wide distrib-

ution in tropical seas, and that the mor-

phological characters of the shell, includ-

ing features of the protoconch, remain

surprisingly constant throughout its

range. L. emeryi (Ladd, 1966) is a taxon

very similar to L. ornata and could even

be conspecific (MOOLENBEEK, 1996).

REDFERN AND ROLÁN: A new species of Lodderena from the Bahamas

200 um

50 um

Figures 10, 11. Lodderena ornata, 0.78 mm, Abaco, Bahamas (CCR). 10: shell; 11: protoconch.

Figure 12. Lodderena ornata, 0.75 mm, Sáo Tomé, West Africa (CER). Figures 13, 14. Lodderena

ornata, 0.80 mm, Easter Island (CER). 13: shell; 14: protoconch.

Figuras 10, 11. Lodderena ornata, 0,78 mm, Abaco, Bahamas (CCR). 10: concha; 11: protoconcha.

Figura 12. Lodderena ornata, 0,75 mm, Santo Tomé, África occidental (CER). Figuras 13, 14. Lod-

derena ornata, 0,80 mm, Isla de Pascua (CER). 13: concha; 14: protoconcha.

It is surprising that one species such

as L. ornata could be present in such

widely separated areas, especially con-

sidering that the protoconch is paucispi-

ral and does not have a long plank-

totrophic period that would allow wide

distribution. We think that there are two

possible explanations, the first of which

is that dispersal could be attributable to

human intervention, such as the dis-

charge of ballast water from ships.

Alternatively, this could be a case of

reproductive isolation involving more

than one species. Identification from

widely separated geographic areas has

been based solely on shell similarities,

and the biology, ecology and soft parts

are mostly unknown.

Iberus, 23 (2), 2005

ACKNOWLEDGEMENTS

We thank Rodger R. Bunnell, of

Sanibel, Florida, for providing the sedi-

ment sample that formed the basis of

BIBLIOGRAPHY

Kay, E. A., 1979. Hawaiian Marine Shells. Reef

and Shore Fauna of Hawaii, sect. 4: Mol-

lusca. B. P. Bishop Museum Special Publica-

tion, 64 (4), 653 pp.

MAYR, E., 1963. Animal Species and Evolution.

Belknap Press, Cambridge. 797 pp. (6th Ed.,

1979).

MOOLENBEEK, R. G., 1996. New skeneiform

species of the genus Lodderena Iredale, 1924

from the Sultanate of Oman (Gastropoda:

Skeneidae). Vita Marina, 44 (1-2): 21-28.

this study. We also thank Jesús Méndez

of the CACTT, University of Vigo, for the

scanning electron micrography.

OLssON, A. A. AND MCGINTY, T. L., 1958. Re-

cent marine mollusks from the Caribbean

coast of Panama with the description of some

new genera and species. Bulletins of American

Paleontology, 39 (177): 1- 58.

RUBIO, F., ROLÁN, E. AND REDFERN, C., 1998. The

genus Lodderena Iredale, 1924 (Achaeogas-

tropoda, Skeneidae) in the Caribbean. Ar-

gonauta, 11 (2): 39-48.

O Sociedad Española de Malacología —__—_——— Iberus, 23 (2): 7-24, 2005

Los caenogasterópodos terrestres (Mollusca, Orthogastro-

poda) de la Comunidad Valenciana (España)

The land caenogastropods (Mollusca, Orthogastropoda) of the

“Comunidad Valenciana” (Spain)

Alberto MARTÍNEZ-ORTÍ* y Fernando ROBLES**

Recibido el 8-I11-2005. Aceptado el 30-VI-2005

RESUMEN

El estudio de las especies terrestres del Superorden Caenogastropoda, realizado a partir

de la revisión bibliográfica, de colecciones museísticas y recolecciones propias, permite

dar a conocer la presencia en la Comunidad Valenciana de Cochlostoma [Obscurella)

martorelli, Platyla polita polita, Leonia mamillaris, Pomatias elegans, Tudorella sulcata sul-

cata y Truncatella subcylindrica. Se presentan datos sobre su registro fósil, los mapas de

distribución geográfica en el área de muestreo, las características de sus hábitats y su

estado de conservación en la Comunidad Valenciana. Además se amplía la distribución

conocida de P. elegans hacia el sur de la Península Ibérica.

ABSTRACT

The study of the land species of the Superorder Caenogastropoda, by means of biblio-

graphic examination, of collections from museums and our own collected samples, shows

the presence in the “Comunidad Valenciana” of Cochlostoma (Obscurella) martorelli,

Platyla polita polita, leonia mamillaris, Pomatias elegans, Tudorella sulcata sulcata and

Truncatella subcylindrica. The data of fossil registers of each species, geographical distri-

bution maps in the sample area, the characteristics of their habitats and the state of con-

servation in the “Comunidad Valenciana” are hereby presented. Furthermore, the known

distribution of P. elegans has been extended further South in the Iberian Peninsula.

PALABRAS CLAVE: Caenogastropoda, terrestre, Cochlostomatidae, Aciculidae, Pomatiidae, Truncatellidae,

Comunidad Valenciana, España.

KEY WORDS: Caenogastropoda, Land, Cochlostomatidae, Aciculidae, Pomatiidae, Truncatellidae,

“Comunidad Valenciana”, Spain.

INTRODUCCIÓN

Desde 1989 los autores están reali- presentes en el área, concretar su distri-

zando un muestreo sistemático de los bución geográfica en la misma y anali-

moluscos continentales de la Comuni- zar el estado de sus poblaciones, reco-

dad Valenciana con una triple finalidad: mendando medidas de protección para

obtener el censo completo de especies las que se consideran amenazadas.

* Museu Valencia d'Historia Natural. Passeig de la Petxina, 15. E-46008. Email: alberto.martinezGQuv.es

** Instituto «Cavanilles» de Biodiversidad y Biología Evolutiva y Departamento de Geología de la Universitat

de Valencia. c/ Dr. Moliner, 50. E-46100 Burjassot (Valencia). Email: roblesfGuv.es

Iberus, 23 (2), 2005

Tabla I. Muestras estudiadas de caenogasterópodos terrestres, bibliográficas, revisadas y propias, de

la Comunidad Valenciana (España).

Table 1. Samples studied of the land caenogastropods, proceeding from bibliographic data, re-examined

samples and our own, from the “Comunidad Valenciana” (Spain).

Publicadas Publicadas Recolección

no revisadas y revisadas autores

C. martorelli l l /

P p. polita 3 | /

L. mamillaris 9 19 34

P elegans 8 43 159

[. s. sulcata - - 4

[. subcylindrica 3 | |

Total 24 65 212

Simultáneamente se han revisado las

colecciones depositadas en diversos

museos y se ha recopilado y evaluado la

información bibliográfica disponible

sobre la materia (MARTÍNEZ-ORTÍ y

ROBLES, 2003).

Presentamos en este trabajo los

resultados obtenidos sobre las seis espe-

cies terrestres del Superorden Caenogas-

tropoda Cox 1960 que habitan en la

Comunidad Valenciana: Cochlostoma

(Obscurella) martorelli (Servain, 1880),

Platyla polita polita (Hartmann, 1840),

Leonta mamillaris (Lamarck, 1822), Poma-

tias elegans (O.F. Miller, 1774), Tudorella

sulcata sulcata (Draparnaud, 1805) y

Truncatella subcylindrica (Linnaeus,

1767). El estudio se completa con el aná-

lisis del registro fósil cuaternario de

estas especies en el mismo ámbito geo-

gráfico.

MATERIAL Y MÉTODOS

Los puntos de muestreo cubren todo

el ámbito geográfico de la Comunidad

Valenciana y corresponden a hábitats

muy variados: roquedos, cuevas, marja-

les, cultivos, etc. Las muestras examina-

das, en su mayoría, proceden de la

colección MARTÍNEZ-ORTÍ, depositada en

el Museu Valencia d'História Natural de

Valencia (MVHN) y aparecen reflejadas

también en su tesis doctoral de 1999.

Colecciones Registro

(inéditas) fósil Total

l 10

12 74

8 | 219

| 5

6 11

27 2 330

Además se han revisado muestras de las

colecciones ROBLES (Museo de Geología

de la Universitat de Valencia), BOSCÁ y

SIRO DE FEZ (MVHN), ROSELLÓ (Museo

de Ciencias Naturales de Valencia,

MCNV), AGUILAR-AMAT, BOFILL,

GASULL, MARTORELL y ROSALS (Museu

de Zoologia de Barcelona, MZB) y las de

AZPEITIA, HIDALGO y PAZ Y MEMBIELA

(Museo Nacional de Ciencias Naturales

de Madrid).

En total se han recopilado 330 regis-

tros cuya distribución se indica en la

Tabla 1. De ellos, 24 corresponden a citas

bibliográficas que no han podido ser

confirmadas por los autores mediante la

revisión del material original. 84 se han

recogido en la provincia de Castellón,

105 en la de Valencia y 141 en la de Ali-

cante. En el listado sólo se cita el autor

que señala una localidad por primera

vez, excluyendo aquellos autores que

repiten citas anteriores. Las muestras

procedentes de localidades bibliográfi-

cas que han podido ser revisadas, al

encontrarse depositadas en diversos

museos españoles, se indican con (R). La

posición geográfica de los puntos de

muestreo se sitúa por su coordenada

UTM, con precisión de 1 km para las

muestras recolectadas por los autores y

de 10 km para las bibliográficas. Para

cada especie se recopilan las citas

previas, se lista el material examinado

indicando la colección a la que perte-

MARTIÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

nece, se da a conocer su distribución

geográfica en la Comunidad Valenciana,

su hábitat y el estado de conservación

en las especies amenazadas, así como

las medidas de protección y la legisla-

ción medioambiental que les afectan.

Para la Sistemática y Nomenclatura

se ha utilizado el listado proporcionado

por el proyecto CLECOM I+II (Check

RESULTADOS Y DISCUSIÓN

En total, se han hallado seis especies

de caenogasterópodos terrestres, perte-

List of European Continental Mollusca)

(BANK, BOUCHET, FALKNER, GITTENBER-

GER, HAUSDORFE, VON PROSCHWITZ Y

RIPKEN, 2001; FALKNER, BANK y RIPKEN,

2001), teniéndose en cuenta las observa-

ciones y modificaciones contenidas en

BANK, FALKNER, NORDSIECK Y RIPKEN

(2001) y en FALKNER, RIPKEN Y FALKNER

(2002).

necientes a dos Ordenes, tres superfami-

lias y cuatro familias.

Familia COCHLOSTOMATIDAE Kobelt, 1802

Cochlostoma (Obscurella) martorelli (Servain, 1880) (Figs. 1, 11-13, 20)

Citas previas: ROSELLÓ (1934): Lucena

(R) (YK34). GaAsuLL (1981): Chodos

(YK35). Material inédito: Col. MARTÍNEZ-

OrTÍí: Castillo de Villamalefa. Fte. Tosca

(YK2450); La Pobla de Benifassáa. Font de

la Canaleta (BF6205). Vistabella. Fuente

coput (YK3262); Vistabella. Penyagolosa.

Canaleta S (YK2555); Vistabella. Penya-

golosa. Fte. Pegunta (YK2558); Xodos

(=Chodos). Barranco (YK3058). Col.

ROBLES: Xodos. Fte. Archivello (YK2959).

Col. SIRO DE FEZ: Lucena (YK34).

Dimensiones: Macho (m): 13,2 mm

hmx y 6,0 mm de Vmx; hembra (h): 14,3

mm hmx y 6,0 mm Ymx. Según GOFAS Y

BACKELJAU (1994) distinguir las especies

de este género resulta muy difícil, debido

a su escasa diferenciación y a la tendencia

a formar cada una de ellas poblaciones

locales. Si bien las características de nues-

tros ejemplares coincidían con los que

estos autores asignan a C. martorelli,

algunos ejemplares de la fuente de la

Pegunta han sido revisados por el Dr.

Gofas, quien ha confirmado la asignación

específica.

Aparato reproductor (Figs. 11-13): Las

genitalias, de ambos sexos, de nuestros

ejemplares coinciden con las descritas y

figuradas por GOFAS Y BACKELJAU (1994).

Distribución geográfica (Fig. 20):

Endemismo localizado preferentemente

en las montañas y valles de los Pirineos

orientales, franceses y españoles, con

algunas localidades aisladas en las mon-

tañas de Montserrat (provincia de Barce-

lona) y en la provincia de Tarragona

(ALTONAGA, GÓMEZ, MARTÍN, PRIETO,

PUENTE Y RALLO, 1994; BOFILL Y HAAS,

1920; GASULL, 1981; GOFAS Y BACKELJAU,

1994; KERNEY Y CAMERON, 1999; RAVEN,

1990; VILELLA, 1967). En la Comunidad

Valenciana se distribuye por la provin-

cia de Castellón, en las comarcas de l'Al-

calatén, el Alto Mijares y el Baix Maes-

trat (MARTÍNEZ-ORTÍ Y ROBLES, 2003).

Las nuevas localidades amplían la dis-

tribución de esta especie en el interior

de esta provincia, de donde únicamente

se conocían dos localidades (GASULL,

1981; ROSELLÓ, 1934). Estos enclaves

constituyen el área de distribución más

meridional de esta especie.

Hábitat: Ha sido recogida en lugares

de altitud superior a 800 m, en fuentes o

muros y roquedos calcáreos que en

general están cercanos a fuentes O

arroyos. La muestra de Penyagolosa-

Canaleta sur fue recogida a 1.600 m de

altitud, en un roquedo que general-

mente se encuentra envuelto por nieblas

y donde la humedad suele ser elevada.

Conservación: Especie propuesta para

su protección en la Comunidad Valen-

ciana por MARTÍNEZ-ORTÍ Y ROBLES

(2003).

Iberus, 23 (2), 2005

Familia ACICULIDAE J.E. Gray, 1850

Platyla polita polita (Hartmann, 1840) (Figs. 2, 20)

Citas previas: Fez (1961): Pego. San

Juan (R) (Y]50). BOETERS, GITTENBERGER

Y SUBAI (1989): Benirrama (= 6 km ssw

Pego) (YJ40); Dénia. Cueva de Benima-

quia (BD50); Jeresa (YJ32).

Material inédito: Col. Martínez-Ortí:

Benialí. Benirrama. Alto del Chap

(YJ4301); Benialí, ctra. a Pego km 37,5

(YJ4403); Gandia. Cova Xurra (YJ4117);

Pego. Bco. de los Frailes (YJ5200); Sueras.

Font de Castro (YK2424); Alzira, la Murta

(V. Escutia leg.). Col. Siro de Fez: Dénia.

Santa María del Mar (BD50).

Dimensiones: Las dimensiones máximas

de las conchas estudiadas son 3,55 mm de

altura y 1,25 mm de diámetro. GITTEN-

BERGER en SEDDON Y HOLYOAK (1993)

indica que los ejemplares de esta especie

que viven en nuestra área de estudio pre-

sentan un tamaño relativo mayor y un

ápice claramente romo, en comparación

con los de Europa central.

Distribución geográfica (Fig. 20): Espe-

cie paleártica atlántico-mediterránea oc-

cidental y europea, distribuida por va-

rios países mediterráneos: N de África,

España, Francia, S de Italia y Sicilia y

centroeuropeos hasta el N de Rusia (Bo-

ETERS ET AL., 1989; GASULL, 1975; GITTEN-

BERGER, 1990, 1991; KERNEY, CAMERON Y

JUNGBLUTH, 1983; SEDDON Y HOLYOAK,

1993; SHIKOV, 1984). Las localidades va-

lencianas son las únicas conocidas en la

Península Ibérica y quedan muy alejadas

de otros enclaves europeos (BOETERS ET

AL., 1989; PRIETO, MARTÍN Y GÓMEZ,

1887; PRIETO, MARTÍN, GÓMEZ Y LARRAZ,

1986). Se localizan en las tres provincias

de la Comunidad Valenciana, en la de

Castellón en la comarca de la Plana Alta,

en la de Valencia en la Safor y en la Ri-

bera Alta y en Alicante en la Marina Alta

(MARTÍNEZ-ORTÍ Y ROBLES, 2003).

Hábitat: No se ha encontrado ningún

ejemplar vivo. Las conchas estudiadas

se han localizado en cuevas y en la base

de roquedos, tamizando gran cantidad

de tierra (MARTÍNEZ-ORTÍ Y ROBLES,

2003).

Conservación: Especie propuesta para

su protección en la Comunidad Valen-

ciana por MARTÍNEZ-ORTÍ Y ROBLES (2003).

Familia POMATIDAE Newton, 1891

Leonia mamillaris (Lamarck, 1822) (Figs. 3, 14, 15, 20)

Citas previas: ROSSMASSLER (1853): Ali-

cante (=Alacant) (YH24). HIDALGO (1870):

Alicante (R). MARTORELL Y BOFILL (1888):

Alicante (RR). SAINT-SIMON (1891): Orihuela

(XH81). ROSELLÓ (1910): Alicante (R).

BOFILL Y AGUILAR-AMAT (1924): Hifac (R)

(BC48). PICARD (1949): Alicante. GASULL

(1975): Alicante. Cap Huertas (R) (YH24);

Alicante. Castillo Santa Bárbara (YH24);

Alicante. Ctra. a Elche km 73 (R) (YH14);

Aspe. Ctra. Elche km 3 (R) (X<H94); Beni-

dorm. Sierra Helada (R) (YH57); Benijófar

(R) H91); Campoamor (R) (XG99); Cre-

villente (R) (XH93); Elche. Pantano Vina-

lopó (R) ((H94); Faro de Santa Pola (R)

(YH13); Ifac. Peñón; Orihuela. Castillo (R)

(H81); Pinet. Ctra. Elche (R) (YH02); Santa

Pola. Ctra. Faro (R) (YH13); Torrevieja

(YHO00). IBÁÑEZ Y ALONSO (1980): Beni-

dorm. Cayola (YH47); Carretera de Santa

Pola a Elche (YH03); Cuevas de Canalo-

bre (YH26); Punta de Llomarit (YH25); vía

férrea al oeste de la estación de Benidorm.

Barranco de la Tapia (YH46). FRANK (1987):

La Marina (YHO02). TALAVÁN Y TALAVÁN

(2004): Islote de Benidorm (YH56).

Material inédito: Col. Azpeitia: Alicante;

La Vila Joiosa (YH46); Orihuela. Col. Bofill:

Alicante; Valencia. Col. Boscá: Alicante.

Col. Martínez-Ortí: Aguas de Busot. Bal-

neario (YH2965); Alacant, ctra. a Elx km

4 (YH1343); Alacant. El Arenal. Ermita

Ntra. S?. Rosario (YH1733); Alacant. El

Arenal. Playa (YH1640); Alacant. Serra de

Fontcalent (YH1047); Alacant. Tánger,

cementerio (YH2054); Alacant. Villafran-

queza, a 1 km (YH2054); Albatera. Monte

Alto (XH8135); Calp. Penyal d'Ifac

MARTÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

(BC4580); Campello, ctra. a Alacant.

Fábrica de Balaustres (YH2858); Crevillent.

Entrada desde Elx (XH9236); Elx. L'Altet

(YH1539); Guardamar. El Moncayo

(YHO0517); La Vila Joiosa. Embalse de Ama-

dorio (YH3969); La Vila Joiosa. Casas del

Cojo (YH3768); La Vila Joiosa, ctra. a Relleu

km 4,1 (YH4069); La Vila Joiosa. Plá Cal-

dereta (YH3463); Monforte del Cid. Casas

de Bautista (YH0546); Orihuela. Barranco

(Bco.) de la Cañada de la Estaca (XG9196);

Orihuela. Bco. al N del bco. de la Cañada

de la Estaca (XH9901); Orihuela. Bco. al S

del bco. de la Cañada de la Estaca

(XH9900); Orihuela. Cabezo la Pedrera

(X<H8511); Orihuela. Puerto de Rebate, km

20 (<H8603); Orihuela. Rincón de Bonanza

(XH7718); Orihuela. Torremendo

(X<H8707); Pilar de la Horadada. Cañada

de Matamoros (XG9597); Pilar de la Hora-

dada, ctra. a Rebate km 23 (XH8600); Pilar

de la Horadada, ctra. a Rebate km 24

(XG9493); Pilar de la Horadada. Dehesa

de Campoamor (XG9898); Pilar de la Hora-

dada. Paraje natural Río Seco (XG9196);

Pilar de la Horadada. Pueblo (XG9597);

Santa Pola. Cap de Santa Pola. Faro

(YH1732); Torrevieja. Barranco (YHO0508);

Torrevieja. Urbanización Los Balcones

(X<H9804). Col. Siro de Fez: Altea (YH57);

Alicante. Camino del Castillo; Santa Pola.

Cap de Santa Pola (YH13); Valencia.

Observaciones: Teniendo en cuenta la

distribución de esta especie, creemos que

la localidad de las muestras n“80-1105

(MZB) y n*354 (MVHN), “Valencia”, no

debe corresponder a la localidad de reco-

lección, sino que debe referirse a la

Comunidad Valenciana en general.

Dimensiones: Las medidas han sido

realizadas sobre 31 ejemplares recogidos

vivos (13 m y 18 h). M: 17,1-15,7 mm h y

10,3-9,1 mm WJmx; h: 20,0-17,4 mm h y

12,0-11,6 mm Ymx. Se aprecia un claro

dimorfismo sexual, en el que las

hembras presentan, en general, mayor

tamaño que los machos, coincidiendo

con los datos obtenidos por ALONSO E

IBÁÑEZ (1980) e IBÁÑEZ Y ALONSO (1980)

de la población de Bolnuevo (Almería).

Aparato reproductor (Figs. 14, 15): La

morfología de las genitalias de los ejem-

plares estudiados, de ambos sexos, coin-

ciden con las descritas y figuradas por

ALONSO E IBÁÑEZ (1980) e IBÁÑEZ Y

ALONSO (1980).

Distribución geográfica (Fig. 20): Especie

Ibero-nordafricana que se presenta en el

Norte de África desde la zona oriental del

Rif (Marruecos) a Orán (Argelia) y en la

Península Ibérica se conoce en las pro-

vincias de Alicante, Murcia, Almería

(ALONSO E IBÁÑEZ, 1980; GASULL, 1972,

1975; GIUSTI Y MANGANELLI, 1984; IBÁÑEZ

Y ALONSO, 1980; MERMOD, 1952; PICARD,

1949; SAccH1I, 1957; SACCHI Y Nos, 1958)

y de Punta de la Mona, La Herradura en

Almuñecar (UrM=305VF5271) y que

corresponde a la primera cita de esta

especie para la provincia de Granada.

Según SACCHI (1957) e IBÁÑEZ Y ALONSO

(1980) esta distribución podría indicar una

posible conexión terciaria entre la región

bética de la Península Ibérica y el Rif (N

de Marruecos). En la Comunidad Valen-

ciana vive en la provincia de Alicante en

las comarcas de la Marina Baixa, 1'Ala-

cantí, el Vinalopó Migja, el Baix Vinalopó

y la Vega Baja.

Hábitat: Especie asociada a suelos

calcáreos y pedregosos (GASULL, 1975;

IBÁÑEZ Y ALONSO, 1980). Ha sido encon-

trada en este tipo de suelos ligada a

pinares y matorral mediterráneo, hasta

una altura máxima de 600 m. Ha sido

recogida viva en 14 localidades. Habita

junto a Tudorella s. sulcata o a Pomatias

elegans en varias localidades. GASULL

(1972) la encuentra junto a ejemplares

fósiles de T. s. sulcata en Almería.

Pomatias elegans (O.F. Múller, 1774) (Figs. 4, 16, 17, 21)

Citas previas: HIDALGO (1871): Játiva

(11); Peñíscola (R) (BE77); Valencia (R)

(YJ27). RoseELLÓ (1910): Valencia (R).

PARDO (1920): Valencia. BOFILL Y AGUI-

LAR-AMAT (1924): Cim del Mondúber (1)

(YJ32). La Barraca de Valldigna. Bco. de

la Falzía (R) (YJ23). Mascarat (R) (BC38).

Montgó (R) (BC59). Sogorb (R) (YK11).

Xeresa (YJ42). Fez (1961): Pego. San Juan

(R) (YJ50). MADURGA (1973): Gandia. Ca-

Iberus, 23 (2), 2005

verna del Parpalló (fósil) (V-YJ42). GA-

SULL (1975): Ayora. Acequia Les Chichi-

les (R) (XJ62); Banyeres. Río (R) (YHO08).

Benidoleig. Cova les Calaveres (KR)

(YH59). Buñol. Cueva de Turche (R)

(XJ86). Callosa d'Ensarriá. El Algar (R)

(YH58). Cerda. Acequia de Ranes (R)

(YJ11); Corbera d'Alzira. Les Fontanelles

(R) (YJ24). Dénia. Montgó (R) (BC59).

Gandía. Mondúber (R) (YJ32). Garganta

de Gata (R) (BC49). Ifac. Peñon (R)

(BC48). Jarafuel (R) (XJ63). Játiva. Ace-

quia La Murta (La Vila) (R) (4J11). Játiva.

Alboy (YJ11); Játiva. Castillo (R) (YJ11).

Jeresa. Les Cingles (R) (YJ32); La Nucia.

La Favara (R) (YH57). Montaverner. Río

Clariano (R) (YJ10); Ontinyent. Pous

Clars (R) (YHO09); Pego. El Bodoix (YJ50).

Pego. San Juan (R) (YJ50). Tabernes de

Valldigna. Monte Umbría (R) (YJ32); Va-

llada. Castillo (R) (YJ00); GASULL (1981):

Campos de Arenoso (R) (YK04); La Jana

(R) (BE68); Lucena del Cid. Bco. (R)

(YK34); Lucena del Cid. Huertos (R)

(YK34); Montanejos (R) (YK13). Monta-

nejos. Cueva Negra (R) (YK13); Monta-

nejos. La Alquería (R) (YK13); Navajas.

Fte. la Luz (R) (YK11); Viver. Fte. San Mi-

guel (R) (YK02). FRANK (1987): La Ma-

rina, (YHO02). ROBLES (1991): Islas Colum-

bretes (fósil) (CE01). ALTONAGA ET AL.

(1994): Castelló de Rugat (YJ20); Peñón

de Ifach (BC48); Ayelo de Rugat (YJ30);

Xátiva: castillo (YJ11). HERRERO-BORGO-

ÑÓN Y GONZÁLEZ (1993): Palma de Gan-

dia. Cova del Blanquissal (YJ41).

Observaciones: GASULL (1975) publica

la localidad: “Játiva. Acequia de la

Murta”. Sin embargo en la etiqueta

aparece: “Játiva. Acequia de la Vila”.

Creemos que se trata de un error de

trascripción, siendo válida la localidad

indicada en la etiqueta.

Material inédito: Col. Hidalgo: Ollería

(YJ11); Orihuela (XH81). Col. Martínez-

Ortí: Adzaneta del Maestrat, cueva os-

cura (YK3753); Agres, cruce río Agres-

FFCC (YH1696); Aín. Bco. de la Caridad

(YK2719); Aín. Cueva del Gat (YK2820);

Alacant. El Arenal. Playa (YH1740); Al-

cala de Xivert. Corral de Capellanes

(BE6862); Alcalalí. Llosa de Camatxo

(YH5995); Alcoi. Parc Natural la Font

Roja. Pico Menejador (YH1482); Almedí-

jar. Collado del Cañar (YK2116); Alzira.

La Murta (YJ2834); Argelita. Bco. a 1 km

(YK2539); Artana. Bco. de Castro

(YK3519); Banyeres, cruce ctras.

(YH0189); Benafer. Bco. de Carlos

(YK0227); Benasal. Balneario (YK4372);

Bicorp, ctra. a Quesa km 11 (XJ9233); Ca-

lig. Bco. Río Seco (BE7782); Benasal. Pis-

cina municipal (YK4272); Benialí. Beni-

rrama (YJ]4403); Benialí. Benirrama. Alto

del Chap (YJ4301); Benialí, ctra. a Pego,

km 37,5 (YJ4403); Benichembla, río Xalo

(YH5193); Benimaurell (YH4995); Bejís.

Fte. los Cloticos (XK9322); Borriol. Río

Seco. Puente (YK4935); Calig. Salida

(BE7582); Callosa d'Ensarriá. Bolulla

(YH5185); Callosa d'Ensarria. Río Gua-

dalest (YH5080); Calp. Parc Natural del

(Página derecha) 1. Cochlostoma martorelli (Servain, 1880) (hembra), Vistavella, fuente coput (Cas-

tellón) (12,86 mm h). 2. Platyla polita polita (Hartmann, 1840), Benialí, ctra. a Pego km 37,5 (3,5

mm h). 3. Leonia mamillaris (Lamarck, 1822) (macho), Pilar de la Horadada, pueblo (Alicante)

(16,5 mm h). 4. Pomatias elegans (O.E. Miller, 1774) (macho), L Alcudia de Veo, camino rural

(Castellón) (14,6 mm h). 5. Tudorella sulcata sulcata (Draparnaud, 1805) (hembra), Orihuela,

barranco al N del barranco de la Cañada de la Estaca (Alicante) (20,78 mm h), 6-10. Ejemplar

adulto de Truncatella subcylindrica (Linnaeus, 1767), Valencia, puerto (MVHN n2817) (4,7 mm

h). 6: vista frontal; 7, 8: vista apical; 9, 10: detalles de la ornamentación.

(Right page) 1. Cochlostoma martorelli (Servain, 1880) (female), Vistavella, coput spring (Castellón)

(12.86 mm h). 2. Platyla polita polita (Hartmann, 1840), Benialí, road to Pego km 37.5 (3.5 mm

h). 3. Leonia mamillaris (Lamarck, 1822) (male), Pilar de la Horadada, village (Alicante) (16.5 mm

h). 4. Pomatias elegans (O.E Miller, 1774) (male), L'Alcudia de Veo, country lane (Castellón) (14.6

mm h). 5. Tudorella sulcata sulcata (Draparnaud, 1805) (female), Orihuela, the gully north of the

gully of la Cañada de la Estaca (Alicante) (20.78 mm h). 6-10. Adult specimen of Truncatella subcy-

lindrica (Linnaeus, 1767), Valencia, port (MVEN n*817) (4.7 mm h). 6: front view; 7, 8: apical

view; 9, 10: details of the sculpture.

lana

dos terrestres de la Comunidad Valenc

ópo

Caenogaster

MARTÍNEZ-ORTÍ Y ROBLES

SINS

ASS

Ny So

Iberus, 23 (2), 2005

Penyal d'Ifac (BC4580); Carretera La

Jana-Canet lo Roig km 2 (BE6690); Ca-

rretera La Cenia-Traiguera. Bco. de la

Cova Alta (BE7294); Castellnovo. Fte.

Marjalet (YK1816); Castelló de Rugat.

Collado del Raconet (YJ2802); Castillo

de Villamalefa. Fte. Tosca (YK2450);

Chelva. Fte. Berra (XK7000); Chelva.

Puente del Reatillo (XK7101); Chera.

Finca la Ermita (XJ7244); Chulilla. Fte.

de la Rinconada (XJ8289); Corbera d'Al-

zira. Cova Negra (YJ2935); Coves de

Vinromá. Font del Molinet (BE5772);

Domeño. Bco. del Agua (XJ7897); Do-

meño. Baños de Verche (XJ7897); Dos

Aguas. Fte. de San José (XJ8951); Dos

Aguas. Bco. del Bosque (XJ8955); En-

guera, camino a Casa Perereta (XJ9312);

Enguera, ctra. a Ayora km 2 (XJ9915);

Estivella. Fte. de Barraix (YJ2397); Fan-

zara. 2,1 km a la Cueva de la Mola

desde ctra. (YK2930); Fondeguilla. Bco.

San Juan (YK3312); Fuentes de Ayodar.

Bco. de Ayodar (YK2033); Gátova. Los

Costales (YK0906); Gestalgar. Fte. de la

Peña María (XJ8486); Gestalgar. Fte. los

Morenillos (XJ8485); Jarafuel. Fte. de las

Anguilas (XJ6633); L'Alcudia de Veo.

Camino rural (YK2125); L'Alcudia de

Veo. Pantano de Benitandús (YK2723);

L'Alcudia de Veo. Racó San Francés

(YK2924); La Pobla de Benifassa. Balles-

tar. Fte. Ballestar (BF6005); La Pobla de

Benifassa. Convent (BF6306); La Pobla

de Benifassá. Font de la Canaleta

(BF6205); La Pobla de Benifassá. Fredes.

Bco. del Salt (BF6210); La Pobla de Beni-

fassá. Font de Sant Pere (BF6805); La Po-

bla de Benifassá. Fredes. Fuente la Roca

(BF6010); La Pobla de Benifassa. Fredes.

Font del Teix (BF6110); La Pobla de Be-

nifassa. Molí del Abad (BF6705); La Po-

bla de Benifassá. Presa (BF6606); La Po-

bla del Duc. Bco. de Sara (YJ2307); La

Vila Joiosa. Plá Caldereta (YH3463);

['Orxa. Río Serpis (YJ3202); Millares.

Alto de la Cuesta (XJ9344); Millares.

Bco. del Hondo (XJ9245); Moixent. Bco.

en el pueblo (XJ9405); Moixent. Pozo

San Juan (XJ9703); Montanejos. Bco. de

la Maimona (YK0938); Morella. Fábrica

de Giner. Río Bergantes (YL4200); More-

lla. Puerto de Torremiró, km 74

(YL4706); Navarrés. Fte. del Río

(XJ9327); Olocau. Bco. de Pedralvilla

(YJ1494); Ontinyent, ctra. a Fontanars

km 9 (YH0196); Ontinyent. Fte. de la

Morera (YH0096); Orba. Campell. Cruce

ctra. a Orba (YH5397); Orba, ctra. a Vall

de Laguar km 3 (YH5397); Ortells. Bco.

de la Juncosa (YL3910); Parcent. Coll de

rates (YH5590); Pego. Bco. de los Frailes

(YJ5200); Pego, ctra. a Alcalá. Barranco

(YH4499); Pego. Frente finca San Juan

(YJ5101); Polop. Entrada (YH5075); Qua-

tretonda, barranco (YJ2719); Quesa. Río

Grande (XJ9129); Relleu. Río Amadorio

(YH3473); Rosell. Molí de Malany

(BF6903); San Antonio de Benagéber. La

Hoya Somera (XK6400); San Rafael del

Río. Masia de Canet (BE7399); Segorbe.

Río Palancia (YK1614); Serra. Fte. del

Berro (YJ1698); Serra. La Caseta del Re-

tor. Bco. Saraguillo (YJ1497); Siete

Aguas. Cueva Alta (XJ8370); Siete

Aguas. Ventamina (XJ8369); Simat, ctra.

a Barx (YJ3323); Sinarcas. Cueva de los

Castillejos (XK5703); Sot de Chera. Fte.

Masalucas (XJ7988); Sot de Ferrer. Río

Palancia. Puente nuevo (YK2109); Sue-

ras. Manantial de Castro (YK2424); Su-

macarcer. Font la Teula (YJ0429); Tales,

cruce de ctras. (YK2925); Tárbena, ctra.

km 28 (YH5288); Tárbena, ctra. a Callosa

km 34 (YH5286); Tarbena. Umbría de

Ferrer (YH5387); Teresa de Viver. Bco.

Uredilla (XK9918); Teulada. Ermita de

San Vicente (BC5089); Titaguas. Campa-

mento del Molino Quemado (XK5616);

Todolella. Bco. de Todolella (YL3203);

Torre Lloris. Río Albaida (YJ1723); To-

rrent. Depósito de agua (YJ1768); Trai-

guera. Bco. de Barranquet (BE7289); Vall

d'Almonacid. Río Chico (YK1617); Vall

d'Ebo, a 1,5 km (YH4599); Vall d'Ebo,

ctra. a Pego km 2 (YJ5001); Vallibona.

Fte. de las Rocas (BE4697); Vallibona.

Fte. Sta. Águeda (BE4897); Vallibona.

Les Moles (BE4997); Vallibona. Masia de

la Torre (BE5199); Vilafamés. Font del

Lleó (YK5042); Villahermosa del Río.

Bco. del Regajo (YK1753); Villar del Ar-

zobispo. Corral del Mosén (XK8702);

Vistabella. Fte. Coput (YK3262); Viver.

Fte. de la Salud (YK0521); Venta la Hi-

guera. Rambla San Mateo (BE5774); Vi-

naros. Playa del Surrac (BE8278); Vista-

bella. Fte. de la Pegunta (YK2558); Xa-

MARTÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

bia. Cala Blanca (BC5794); Xabia. Cap de

Sant Antoni (BC5698); Xabia, ctra. a Dé-

nia km 1,5 (BC5398); Xabia, ctra. al Cap

de la Nau (BC5694); Xabia. Platja del

Portitxol (BC5893); Xativa. Penya San

Diego (YJ1216); Xeresa. Bco. de Xeresa

(YJ3920); Xert. Bco. de la Fuente

(BE6088); Xixona. Río Torremanzanas

(YH1867); Yátova. Entre cola del pan-

tano y ctra. (XJ7959); Yátova. Mijares.

Fte. Ntra. Sra. Desamparados (XJ7661);

Yátova. Río Mijares. Puente (XJ7661);

Zorita. La Balma (YL3814); Zorita. Mo-

lino de Villar (YL4016). Col. MZB: Anna

(YJ02); Hifac (BC48). Col. Robles: Anna.

Cerca de la Albufera (YJO320); Benafer.

Fte. de los Nogales (YK0723); Campell.

Vall Laguart, ctra. al río (2 km)

(YH5196); Canet lo Roig. Font de la

Roca (BE6691); Fanzara. Fuente de 1'Al-

cudia (YK2832); Islas Columbretes (fósil,

CE0219); La Pobla de Benifassa. Font del

Convent (BF6306); Navajas. Fte. de la

(YK1317); Ribarroja. Fte. la Cisterna

(YJ0980); Rossell. Font del Baix (BF6400);

Salzedella. Fte. Ciurana (BE5974); Sarra-

tella. Font del Torrent (BE4766); Sot de

Chera. Río Reatillo (XJ7887); Teresa de

Cofrentes (XJ6830); Vallibona. Font del

Fou (BE5899); Vilanova d'Alcolea. Font

de la Vila (BE5057). Col. Siro de Fez: La

Barraca de Aigues Vives (YJ2830); Porta-

celi (YJ19); Valencia.

Dimensiones: M: 17,1 mm hmx; 11,9

mm QJmx; h: 185 mm hmx; 129 mm

Hmx.

Las medidas realizadas sobre las

conchas de ejemplares recogidos vivos

de ambos sexos indican que, aunque las

hembras pueden llegar a ser un poco

mayores que los machos, no se puede

observar un claro dimorfismo sexual.

Aparato reproductor (Figs. 16, 17): La

morfología de las genitalias, de ambos

sexos, de nuestros ejemplares coinciden

con las descripciones y figuras de CREEK

(1951), ALONSO E IBÁÑEZ (1977) y GIUSTI

ET AL. (1995).

Distribución geográfica (Fig. 21):

Especie europea, atlántico-mediterránea

occidental y centroeuropa (ADAM, 1960;

ALONSO E IBÁNEZ, 1977; AÁLTONAGA ET

AL., 1994; GITTENBERGER, BACKHUYS Y

RIPKEN, 1984; GROSSU, 1986; KERNEY ET

AL., 1983). Esta distribución puede venir

condicionada por su intolerancia a los

inviernos fríos, lo que hace que presente

su límite septentrional a lo largo de una

línea que sigue la isoterma de 2 *C en el

mes de enero (KILLEEN, 1992).

En la Península Ibérica se distribuye

por el área central de Portugal, la

Cornisa Cantábrica desde Galicia hasta

el País Vasco, Navarra, Valle del Ebro

hasta los Pirineos y por el Mediterráneo

se extiende por las Islas Baleares e Islas

Medas y por el continente desde Cata-

luña hasta Murcia (ALONSO E IBÁÑEZ,

1977; ALTIMIRA, 1969; ALTIMIRA Y

ALTABA, 1984; ALTONAGA ET AL., 1994;

ANGULO Y MARTÍN, 1985; CASTILLEJO,

1981; Faci, 1991; GASULL, 1974, 1975;

IBÁÑEZ Y ALONSO, 1980; LARRAZ Y EQUI-

SOAÍN, 1993; NOBRE, 1941; PICARD, 1949).

Recientemente han sido halladas dos

poblaciones en la provincia de Málaga:

Barranco de la Coladilla (UTM=

305VF2468) y en el Río de la Miel

(UTM= 305VF2768), ambas en Nerja,

que constituyen las primeras citas para

Andalucía. Sería necesario realizar más

muestreos en la región para confirmar

que no se trata de introducciones antró-

picas. En Aragón se distribuye preferen-

temente por el pre-pirineo, evita la

depresión del Ebro en la zona de los

Monegros y aparece en la Meseta Ibérica

(FacI, 1991). En la Comunidad Valen-

ciana se distribuye ampliamente por las

provincias de Castellón en todas sus

comarcas, Valencia, donde falta única-

mente en la comarca del Rincón de

Ademuz, y Alicante, donde se conoce de

las comarcas del Comtat, la Marina Alta

y la Baixa, l'Alcoia, 1'Alacantí y la Vega

Baja. Ha sido citada en las Islas Colum-

bretes, donde ROBLES (1991) señala que

esta especie debe darse como extin-

guida, tras los numerosos muestreos

realizados en los que únicamente se han

encontrado conchas. Los ejemplares

fósiles han sido datados del último

glaciar (-17.000 años).

Hábitat: Es una especie humícola,

detritícola y calcícola (ADAM, 1960;

ALTONAGA ET AL., 1994; GASULL, 1975,

1981; GIUSTI ET AL., 1995; IBÁÑEZ Y

ALONSO, 1980). Ha sido recolectada en

Iberus, 23 (2), 2005

ambientes de pinada, matorral medite-

rráneo y también, aunque en menor

medida, en encinares, riberas, arroyos y

cultivos. Se halló viva en 47 localidades.

Los ejemplares estaban escondidos

debajo de piedras, mantillo o en la base

del matorral, donde la humedad era

más abundante y, en general, se presen-

taban en colonias. Estos datos coinciden

con los de FaAcI (1991) y LARRAZ Y EQUI-

SOAÍN (1993). Se han encontrado ejem-

plares superando los 1.300 m de altitud

tanto en la provincia de Alicante como

en la de Castellón. Faci (1991) indica

que sus muestras fueron recolectadas

entre 300 y 1.000 m y añade que

MERMOD no encontró esta especie en

Suiza en alturas superiores a 1.000 m.

Tudorella sulcata sulcata (Draparnaud, 1805) (Figs. 5, 18, 19, 22; Tabla II)

Comentarios taxonómicos: T. s. sulcata

ha sido, frecuentemente, denominada

como Pomatias sulcatus; sin embargo

FALKNER ET AL. (2002), basándose en el

trabajo de VITTURI, CATALANO Y MACA-

LUSO (1986), asignan esta especie al

género Tudorella P. Fischer, 1885.

Citas previas: ROBLES Y MARTÍNEZ-

Ortí (1995): La Cañada de la Estaca, A-

XG99; Pilar de la Horadada. Paraje

natural de Río Seco, A-XH090.

Material inédito: Col. Martínez-Ortí:

Orihuela. Barranco al N del Barranco de

la Estaca (XH9901); Orihuela. Barranco

al S del Barranco de la Estaca (XH9900).

Col. Robles: Bacarot (fósil, YH1446).

Dimensiones: Esta especie presenta un

claro dimorfismo sexual, con las hembras

de tamaño generalmente superior al de

los machos. En la Tabla II queda patente

esta característica, de acuerdo con las

medidas de una muestra de 73 indivi-

duos adultos recogidos vivos en la locali-

dad del barranco al norte del barranco de

la Cañada de la Estaca (Orihuela).

Aparato reproductor (Figs. 18, 19): La

morfología de las genitalias, de ambos

sexos, que han sido recolectados en Ali-

cante coincide con las descritas y figura-

das por IBÁÑEZ Y ALONSO (1978, 1980) y

GIUSTI ET AL. (1995).

Distribución geográfica (Fig. 22): Tudo-

rella s. sulcata es una especie de distribu-

ción mediterránea occidental: SE de

Francia, Córcega, Cerdeña, Sicilia, Malta

y El Maghreb (FECHTER Y FALKNER, 1993;

GIUSTI Y MANGANELLI, 1984; GIUSTI ET

AL., 1995; HAas, 1929; IBÁÑEZ Y ALONSO,

1978, 1980; KERNEY Y CAMERON, 1999;

KERNEY ET AL., 1983; NOBRE, 1941; PA-

VÓN, 2005, en prensa). En la Comunidad

1ó

Valenciana se han encontrado cuatro po-

blaciones actuales en la Vega Baja, en el

sur de la provincia de Alicante (MARTÍ-

NEZ-ORTÍ Y ROBLES, 2003; ROBLES Y MAR-

TÍNEZ-ORTÍ, 1995) y una fósil atribuida al

Pleistoceno inferior.

En la Península Ibérica, además de las

localidades valencianas, T. s. sulcata vive

en otras dos localidades, en el Algarve

(sur de Portugal) y en Motril (Granada)

(IBÁNEZ Y ALONSO, 1978; NOBRE, 1941).

Los autores han constatado la existencia

de individuos vivos en ambas localida-

des en Enero de 2005. La distribución de

esta especie en yacimientos plio-cuater-

narios es más amplia: Palau Sacosta (pro-

vincia de Gerona) (BOFILL ET AL., 1921;

GASULL, 1972; HAAS, 1929; SACCHI, 1957),

La Pita Calataray y San Juan de Terreros

(provincia de Almería) (IBÁÑEZ Y

ALONSO, 1978), Casas del Rincón (provin-

cia de Albacete) (ALBERDI, ARIAS, BI-

GAZZI, BONADONNA, LEONE, LÓPEZ, MI-

CHAUX, MORALES, ROBLES Y SORIA, 1982),

Cañada de Murcia (provincia de Gra-

nada), Sierra de Quibas (Abanilla, pro-

vincia de Murcia) (ROBLES, 1989; ROBLES

Y MARTÍNEZ-ORTÍ, 1995) y Bacarot (Ali-

cante). La edad de tres de los yacimientos

es conocida, ya que se encontraron ma-

míferos fósiles: Casas del Rincón es del

Plioceno terminal y Cañada de Murcia y

Sierra de Quibas son del Pleistoceno infe-

rior. Bacarot puede correlacionarse con

Quibas. El yacimiento de Palau Sacosta

ha sido atribuido al “Cuaternario anti-

guo” (GASULL, 1972). La edad de los yaci-

mientos de Almería es pleistocena, sin

que sea posible precisarla más.

La comparación de la distribución

de esta especie en el Plioceno terminal y

MARTÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

Figuras 11-19. Aparatos reproductores. 11-13. Cochlostoma (Obscurella) martorelli (Servain, 1880),

Vistavella, Penyagolosa, fuente de la Pegunta (Castellón). 11: genitalia de una hembra; 12: genita-

lia de un macho; 13: detalle del pene. 14, 15. Leonia mamillaris (Lamarck, 1822), Pilar de la Hora-

dada, dehesa de Campoamor (Alicante). 14: genitalia de una hembra; 15: genitalia de un macho.

16, 17. Pomatias elegans (O.F. Miller, 1774), Alcudia de Veo, camino rural (Castellón). 16: geni-

talia de un macho; 17: genitalia de una hembra. 18, 19. Tudorella sulcata sulcata (Draparnaud,

1805), barranco al N del Barranco de la Cañada de la Estaca (Orihuela, Alicante). 18: genitalia de

un macho; 19: genitalia de una hembra. Escalas, 1 mm.

Figures 11-19. Reproductive systems. 11-13. Cochlostoma (Obscurella) martorelli (Servain, 1880), Vis-

tavella, Penyagolosa, Pegunta spring (Castellón). 11: female genitalia; 12: male genitalia; 13: detail of

the penis. 14, 15. Leonia mamillaris (Lamarck, 1822), Pilar de la Horadada, dehesa de Campoamor

(Alicante). 14: female genitalia; 15: male genitalia. 16, 17. Pomatias elegans (O.E Múller, 1774),

LAlcudia de Veo, country lane (Castellón). 16: male genitalia; 17: female genitalia. 18, 19. Tudorella

sulcata sulcata (Draparnaud, 1805), the gully north of the gully of la Cañada de la Estaca (Orihuela,

Alicante). 18: male genitalia; 19: female genitalia. Scale bars, 1 mm.

el Pleistoceno inferior con la distribu- la península de los yacimientos pliocua-

ción actual, permite asignar a los encla- ternarios, parece indicar un retraimiento

ves actuales un carácter relicto. Su posi- a refugios cálidos de su área original de

ción en las proximidades del mar, frente distribución, debida a los episodios fríos

a la amplia distribución en el interior de del Cuaternario.

Iberus, 23 (2), 2005

Tabla II. Dimensiones de Tudorella sulcata sulcata. Ejemplares procedentes del barranco al N del

Barranco de la Estaca (Orihuela, Alicante).

Table 11. Shell measurements of Tudorella sulcata sulcata. Specimens proceeding from the gully North of

the gully of la Cañada de la Estaca (Orihuela, Alicante).

T. sulcata sulcata Altura Diámetro

n=73 máx - X - min Or] máx -X - min Or]

Hembras 30 23,23 19601115 15 16,39-14,34-12,61 0,79

Machos 43 199017 1115/35 00/81 14,65-12,69-11,42 0,54

Hábitat: Vive en ambientes similares a

Leonia mamillaris, con la que se ha encon-

trado conviviendo en la Comunidad Va-

lenciana. Las poblaciones ibéricas viven

en áreas cercanas a la costa y a una altitud

que no supera los 100 m. Las poblaciones

alicantinas viven sobre suelos calcáreos y

pedregosos ligados a pinares de Pinus ha-

lepensis (pino carrasco) y matorrales ter-

momediterráneos y pre-estépicos, con es-

pecies como Chamaerops humilis (marga-

l1ló o palmito), Pistacia lentiscus (lentisco)

y Stipa tenacissima (esparto), entre la pi-

nocha, en la base de los tallos y debajo de

las piedras (MARTÍNEZ-ORTÍ Y ROBLES,

2003). Concretamente el barranco de la

Cañada de la Estaca presenta como ma-

torral dominante el Sideritido-Helianthe-

metum caput-felicis, cuyas especies priori-

tarias son Helianthemum caput-felis, Sideri-

tis murgetana subsp. littoralis y Thymus

hyemalis (DOCV, 3505/ 28/05/1999). Las

otras dos poblaciones ibéricas, la portu-

guesa y la granadina, se han encontrado

en la base de la vegetación de la cual pro-

bablemente también se alimenta: P. lentis-

cus, Rhamnus lycioides (espino negro), Fo-

eniculum vulgare (hinojo) y Genista sp., en

la primera y de Ch. humilis, tal y como se-

ñalan IBÁÑEZ Y ALONSO (1978), y Mayte-

nus senegalensis subsp. europaeus (espino

cambrón) en la segunda.

Conservación: La localidad del ba-

rranco de la Cañada de la Estaca y los

barrancos al norte y al sur, en la comarca

de La Vega Baja, están siendo actual-

mente urbanizados (MARTÍNEZ-ORTÍ Y

ROBLES, 2003). En un esfuerzo por ga-

rantizar su conservación se trasladaron

50 ejemplares a otra localidad cercana al

Cabo de Santa Pola en 1996 y cuyos te-

rrenos pertenecen a la Generalitat Valen-

ciana (MARTÍNEZ-ORTÍ Y ROBLES, 2003).

Tras examinar nuevamente esta locali-

dad, en Junio de 1999, se observó la pre-

sencia de algunos machos y hembras vi-

vos, lo que permite albergar esperanzas

de su supervivencia en la provincia de

Alicante. En la otra localidad alicantina,

“Pilar de la Horadada. Paraje natural

Río Seco”, no se han encontrado nunca

ejemplares vivos. En el Catálogo Valen-

ciano de Especies de Fauna Amenaza-

das, y tras los informes relativos al es-

tado de conservación de esta especie re-

alizados por los autores en los últimos

años, presentados a la Conselleria de Te-

rritori i Habitatge de la Comunidad Va-

lenciana, ha sido incluida con la catego-

ría de “vulnerable” (DOCV n” 4.705,

4/03/2004: p. 4.972). Además, esta espe-

cie ha sido propuesta por los autores (en

ALONSO, ALTONAGA, ÁLVAREZ, ARAUJO,

ARCONADA, ARRÉBOLA, BECH, BROS,

CASTILLEJO, GÓMEZ, IBÁÑEZ, LUQUE,

MARTÍNEZ-ORTÍ, MORENO, PRIETO,

PUENTE, PUJANTE, ROBLES, ROLÁN Y

TEMPLADO, 2001) para su protección e

inclusión en el Catálogo Nacional de es-

pecies Amenazadas, con la categoría de

“sensible a la alteración de su hábitat”.

KERNEY ET AL. (1983) señalan que las po-

blaciones del SE francés se encuentran

en proceso de extinción, mientras que

KERNEY Y CAMERON (1999) y PAVvóN

(2005) opinan que está en regresión. Éste

último señala que ha desaparecido del

departamento de los Alpes-Maritimes,

es muy escasa en el de Var y abundante,

aunque amenazado por la urbanización

y la degradación del litoral, en el depar-

tamento de Bouches-du-Rhóne.

MARTÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

xXHo2

Figuras 20-22. Distribución geográfica de los caenogasterópodos terrestres de la Comunidad

Valenciana. 20. Cochlostoma martorelli (Servain, 1880) (puntos), Platyla polita polita (Hartmann,

1840) (cuadros) y Leonia mamillaris (Lamarck, 1822) (círculos vacíos). 21. Pomatias elegans (O.F.

Miller, 1774). 22. Tudorella sulcata sulcata (Draparnaud, 1805) (puntos) y Truncatella subcylin-

drica (Linnaeus, 1767) (círculos vacíos).

Figures 20-22. Geographical distribution of the land caenogastropods of the “Comunidad Valenciana”

(Spain). 20. Cochlostoma martorelli (Servain, 1880) (black circles), Platyla polita polita (Hart-

mann, 1840) (squares) and Leonia mamillaris (Lamarck, 1822) (white circles). 21. Pomatias elegans

(O.E Múller, 1774). 22. Tudorella sulcata sulcata (Draparnaud, 1805) (black circles) and Truncate-

lla subcylindrica (Linnaeus, 1767) (white circles).

Familia TRUNCATELLIDAE J.E. Gray, 1840

Truncatella subcylindrica (Linnaeus, 1767) (Figs. 6-10, 22)

Citas previas: ROSELLÓ (1910, 1934): T.

microlena y T. truncatula var. laevigata.

Valencia (R) (YJ27). GASULL (1971):

Calpe. Salinas (BC48); Elche. Salinas de

Pinet (YHO02).

Observaciones: T. microlena Bourguig-

nat, 1884 es una forma de pequeña talla

de la variedad lisa de T. subcylindrica

(GERMAIN, 1931). T. truncatula (Drapar-

naud, 1801) es un sinónimo posterior de

T. subcylindrica (FRETTER Y GRAHAM,

1978; GERMAIN, 1931; GROSSU, 1986). Se

han revisado dos muestras de la colec-

ción ROSELLÓ (MCNV) comprobándose

que corresponden, efectivamente, a T.

subcylindrica.

Material inédito: Col. BoscÁ: Valen-

cia. Puerto. Col. MARTÍNEZ-ORTÍ: Torre-

blanca, prado pantanoso (BE6252). Col.

SIRO DE FEz: Dénia (BD50) (dos mues-

tras); Valencia (tres muestras).

Discusión: La ornamentación de esta

especie es muy variable. En las muestras

estudiadas predominan los ejemplares

con costulación bien desarrollada (Figs.

6, 9, 10), pero existen algunos en los que

este carácter es poco aparente (var. laevi-

gata Risso). FRETTER Y GRAHAM (1978:

138) señalan que “entre las costillas son

a veces visibles delicadas estrías espira-

les” mientras que BUTAKOV, CHUHCHIN,

CHERKASOVA Y LELEKOV (1997, sin pág.)

afirman que la “escultura espiral está

ausente”. La observación a elevados

aumentos de varias conchas, utilizando

el M.E.B., muestra la presencia de fila-

Iberus, 23 (2), 2005

mentos espirales muy finos, bien marca-

dos en el espacio comprendido entre las

costillas y muy próximos entre sí (Figs.

10).

Distribución geográfica (Fig. 22):

Especie con amplia distribución en el

Mar Mediterráneo, Mar Negro y Mar de

Azov. En el Océano Atlántico se

extiende a lo largo de la costa meridio-

nal europea hasta el S de Inglaterra y las

costas francesas del Canal de la Mancha.

Presente también en Azores, Madeira y

Canarias (BUTAKOV ET AL.1997; FRETTER

Y GRAHAM, 1978, 1994; GASULL, 1971;

GROSSU, 1986; IBÁÑEZ, ÁLONSO Y Luis,

2001; WHITE, 1999). Una antigua intro-

ducción en Newport (USA) no ha pros-

perado (CARLTON, 1992). Su distribución

en la Península Ibérica es mal conocida.

CONCLUSIONES

En la Comunidad Valenciana se han

identificado seis especies terrestres de

caenogasterópodos: Cochlostoma (Obscu-

rella) martorelli (Servain, 1880), Platyla

polita polita (Hartmann, 1840), Leonia

mamillaris (Lamarck, 1822), Pomatias

elegans (O.F. Miller, 1774), Tudorella

sulcata sulcata (Draparnaud, 1805) y

Truncatella subcylindrica (Linnaeus,

1767).

Desde los puntos de vista biogeográ-

fico y de su interés para la conservación,

la importancia de estas especies es muy

diferente: C. (O.) martorelli posee en esta

región el límite meridional de su área de

distribución. Esta característica, junto a

la escasez de localidades en la misma,

recomienda su inclusión en el Catálogo

Valenciano de Especies de Fauna Ame-

nazadas (CVEFA). P. polita polita está

representada en la Península Ibérica

solamente por las localidades valencia-

nas, muy alejadas de su área general de

distribución. Por ello se ha recomen-

dado su inclusión en dicho Catálogo y

probablemente debería incluirse, dada

su singularidad, en el Catálogo Nacio-

nal. T. sulcata sulcata presenta, en el sur

de la provincia de Alicante, varias

poblaciones relictas sometidas a fuerte

presión urbanística. En la Península

BECH (1990) recopila las citas de Cata-

luña. En la Comunidad Valenciana se

conoce de las tres provincias. En la de

Castellón, donde se cita por primera

vez, en la comarca de la Plana Alta, en la

de Valencia en la de l'Horta y en la de

Alicante en las de la Marina Alta y el

Baix Vinalopó en el sur.

Habitat: Vive en la zona supralitoral,

enterrada hasta 15 cm entre las raíces de

plantas, detritus vegetal y sedimentos

finos. Más rara en suelos fangosos, bajo

las rocas. Especie anfibia, vive preferen-

temente al aire libre, pero puede perma-

necer sumergida durante largos perio-

dos. Es frecuente también en el borde de

salinas litorales (GASULL, 1971; GROSSU,

1986; FRETTER Y GRAHAM, 1978, 1994;

WHITE, 1999).

Ibérica solamente se conocen otras dos

poblaciones, una en la provincia de

Granada y otra en el Algarve portugués.

Las poblaciones del SE francés, donde se

describió originalmente, se encuentran

en clara regresión. Por todo ello se ha

incluido en el CVEFA y se ha recomen-

dado su inclusión en el Catálogo Nacio-

nal. L. mamillaris y P. elegans son especies

abundantes en la Comunidad, aunque

su distribución geográfica es muy dife-

rente. La primera, iberomagrebí, sólo

ocupa el sur de la provincia de Alicante

mientras que la segunda, de amplia dis-

tribución europea, aparece bien repar-

tida por las tres provincias. Para ambas

se amplía su área de distribución hacia

el sur peninsular. Ninguna de ellas exige

medidas especiales de conservación. Por

último, T. subcylindrica es una especie de

distribución mal conocida en la Penín-

sula Ibérica, por lo que es difícil evaluar

su situación real en la misma.

AGRADECIMIENTOS

A los conservadores, Dr. Francesc

Uribe del Museu de Zoologia de Barce-

lona, Dr. Oscar Soriano del Museo

Nacional de Ciencias Naturales de

MARTÍNEZ-ORTÍ Y ROBLES: Caenogasterópodos terrestres de la Comunidad Valenciana

Madrid y Dra. Margarita Belinchón, del

Museo de Ciencias Naturales de Valen-

cia, por la cesión de las muestras que

han sido revisadas en este trabajo. Al Dr.

Serge Gofas por confirmar la asignación

específica de las poblaciones valencia-

nas de Cochlostoma martorelli. Al Dr. Juan

José Herrero Borgoñón por la determi-

nación de la vegetación de las localida-

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O Sociedad Española de Malacología —__—_——— Iberus, 23 (2): 25-38, 2005

Soft-bottom mollusc assemblages in the Ría de Ares-

Betanzos (Galicia, NW Spain)

Asociaciones malacológicas de substratos blandos de la Ría de Ares-

Betanzos (Galicia, NO España)

Jesús S. TRONCOSO*, Juan MOREIRA? and Victoriano URGORRI**

Recidido el 26-1-2005. Aceptado el 21-VII-2005

ABSTRACT

The composition and spatial distribution of the mollusc fauna on the subtidal soft bottoms of the Ría

de Ares-Betanzos (Galicia, NW Spain) were studied by means of semi-quantitative sampling and

multivariate analyses. The faunal distribution in the ría seems to be mainly conditioned by a depth

and grain size gradients, the latter defined by a increase in silt/clay and a decrease of coarser gra-

nulometric fractions from the mouth towards the margins and inner areas of the ría. Several assem-

blages were determined which could be defined according to the classic terms of *community' and

facies”. Sandy bottoms showed a “Venus fasciata community” in coarser sediments of the outer ría,

while a “Venus gallina community was found in fine sand at the center of the ría. The shallower and

muddier sediments in the inner ría showed a mix of typical species from the “Abra alba' and the

"Venus gallina" communities. However, two facies could be distinguished: the “facies of Ringicula

auriculata-Pandora inaequivalvis' in sediments with a greater fine sand content, and the “facies of

Nassarius pygmaeus-Dentalium novemcostatum' in the muddiest sediments.

RESUMEN

Se estudia la distribución espacial de los Moluscos en los fondos blandos de la Ría de Ares y

Betanzos (Galicia, NO, España) utilizando muestras semi-cuantitativas y técnicas de análisis

multivariante. La distribución de la fauna parece estar condicionada principalmente por la pro-

fundidad y por los gradientes en el tamaño de grano, definidos por un incremento de la frac-

ción pelítica y una disminución de las fracciones gruesas de sedimento desde la boca hasta

los márgenes y zonas internas de la ría. Las agrupaciones halladas pueden ser definidas den-

tro de los términos clásicos de comunidades” y “facies”. Los fondos de arena en la parte más

externa de la ría donde los sedimentos son más gruesos presentan la 'comunidad de Venus

fasciata', mientras que la “comunidad de Venus gallina” fue encontrada en fondos de arena

fina en la parte central de la ría. Los fondos fangosos se encuentran en zonas más someras

del interior de la ría y presentan una mezcla de especies típicas de las comunidades de “Abra

alba” y “Venus gallina”. Sin embargo dos facies pueden ser distinguidas: la “facies de Ringi-

cula auriculata-Pandora inaequivalvis' en sedimentos con altos contenidos de arenas finas y la

facies de Nassarius pygmaeus-Dentalium novemcostatum' en los sedimentos más fangosos.

KEY WORDS: Soft-bottom, molluscs, distribution, Ría de Ares-Betanzos.

PALABRAS CLAVE: Substratos blandos, moluscos, distribución, Ría de Ares-Betanzos.

* Departamento de Ecoloxía e Bioloxía Animal, Facultade de Ciencias, Campus de Lagoas-Marcosende s/n,

Universidade de Vigo, 36200 Vigo, Spain

** Departamento de Bioloxía Animal, Laboratorio de Zooloxía Mariña, Facultade de Bioloxía, Universidade de

Santiago, 15706 Santiago de Compostela, Spain

Correspondence: J. S. Troncoso; e-mail: troncosoCuvigo.es

Iberus, 23 (2), 2005

INTRODUCTION

During the last thirty years, there

has been an ongoing interest in the 'rías”

of Galicia (NW Spain), which are a

special kind of estuarine system. The

rías originated from flooded river

valleys and have a high primary pro-

ductivity due to upwellings and regular

inflows of nutrients (NOMBELA, VILAS

AND EVANS, 1995). The great economic

and social importance of these systems

(fisheries, bivalve culture on rafts, shell-

fish resources) would greatly benefit

from a scientific study of the environ-

ment, especially that of the benthic com-

munities, which are good indicators of

the conditions of marine bottoms

(BELLAN, 1967; PEARSON AND ROSEN-

BERG, 1978; WARWICK, 1988).

The Ría de Ares-Betanzos is the

largest ría of north-west Galicia and is

located between the Ría de Coruña and

the Ría de Ferrol (Golfo Ártabro). This

ría is a double estuarine system with

depths ranging between 2 and 43 m

(SÁNCHEZ-MATA, GLÉMAREC AND MORA,

1999). Over the last years, several

papers have been devoted to its benthic

macrofauna (SÁNCHEZ-MATA, MORA,

GARMENDIA AND LASTRA, 1993; TRON-

COSO AND URGORRI, 1993a; GARMENDIA,

SÁNCHEZ-MATA AND MORA, 1998;

SÁNCHEZ-MATA AND MORA, 1999a; b).

Furthermore, the hard-bottom mollusc

fauna was studied by TRONCOSO,

URGORRI, PARAPAR AND LASTRA (1988)

and TRONCOSO, URGORRI AND OLABAR-

RÍA (1996) while TRONCOSO AND

URGORRKI (1992, 1993b) analyzed the ver-

tical distribution of infauna in the sedi-

ment. However, there is a lack of syne-

cological studies on soft-bottom malaco-

fauna. Thus, this paper deals with the

distribution of soft-bottom mollusc

assemblages and the relation with envi-

ronmental parameters in the subtidal

areas of the Ría de Ares-Betanzos. On

the other hand, this area was strongly

affected by the Aegean Sea oil spill

during 1993, and this paper can there-

fore serve as a baseline study for future

comparisons of molluscan fauna evolu-

tion.

MATERIAL AND METHODS

Sample collection: A total of 55 sta-

tions were sampled in subtidal soft

bottoms to cover adequately the exten-

sion Of the ría (Fig. 1). Sampling

program was carried out between Feb-

ruary and November 1986. Semi-quanti-

tative data were obtained using a natu-

ralist rectangular dredge. Sampled area

varied between stations depending on

nature of substrate; data were stan-

darized to 25 l of collected sediment

(maximum volume of dredge net).

Samples were sieved through a 0.5 mm

mesh; fauna was sorted in the labora-

tory after fixation in 10% buffered for-

malin. An additional sediment sample

was taken at each station to analyze the

granulometric composition, carbonates,

nitrogen (N), organic carbon (C) and

total organic matter (TOM) contents

(TRONCOSO AND URGORRI, 1993a). The

following granulometric fractions were

considered: gravel (GR, >2 mm), coarse

sand (CS, 2-0.5 mm), medium sand (MS,

0.5-0.25 mm), fine sand (FS, 0.25-0.063

mm), and silt/clay (<0.063 mm).

Median grain size (Q50), sorting coeffi-

cient (So) (Trask, 1932) and C/N ratio

were also determined for each sample.

Sedimentary types were characterized

according to RODRIGUES AND QUINTINO

(1985) and JUNOY AND VIÉITEZ (1989).

Data analyses: Total abundance of

individuals (N) and number of species

(S) were calculated for each sampling

station. Mollusc assemblages were deter-

mined through non-parametric multi-

variate techniques as described by FIELD,

CLARKE AND WARWICK (1982) using the

PRIMER v5.0 (Plymouth Routines in

Multivariate Ecological Research) soft-

ware package (CLARKE AND WARWICK,

1994). A similarity matrix between sam-

pling stations was constructed by means

of the Bray-Curtis similarity coefficient

by first applying square root transforma-

tion on species abundance to down-

weight the contribution of the most

abundant species. From this matrix, a

classification of the stations was per-

formed by cluster analysis based on the

group-average sorting algorithm, as well

TRONCOSO ET AL.: Soft-bottom mollusc assemblages in the Ría de Ares-Betanzos

Figure 1. Locations of sampling stations in the Ría de Ares-Betanzos.

Figura 1. Localización de las estaciones de muestreo en la Ría de Ares-Betanzos.

as an ordination by means of non-metri-

cal multidimensional scaling (MDS).

Rare species (1.e., those wich appeared in

one station and /or with 1-2 inviduals)

were included in the final analyses

because preliminary trials showed that

their supression did not affect classifica-

tion and ordination of stations. The

SIMPER program was next used to iden-

tify species that greatly contributed to

similarity in a given group derived from

those analyses.

The possible relationship between

mollusc distribution and the measured

environmental variables were re-

searched using the BIO-ENV procedure

(belonging to the PRIMER package) and

the canonical correspondence analysis

(CCA) using the CANOCO v4.02

(Canonical Community Ordination)

package (BRAAK, 1988). Forward selec-

tion was employed in the latter to detect

which variables explained the most

variance in the species data. All vari-

ables expressed in percentages were

previously transformed by log (x+1) and

then normalised. Stations 16, 29 and 60

were discarded because of low sediment

quantity.

RESULTS

Sediments were mainly of a sandy

nature and muddy bottoms were

restricted to inner and sheltered areas.

Iberus, 23 (2), 2005

% Bray-Curtis Similarity

. 4B

10073

Figure 2. Mollusc assemblages in the Ría de

on Bray-Curtis similarity coefficient.

es-Betanzos as determined by cluster analysis based

Figura 2. Asociaciones malacológicas en la Ría de Ares-Betanzos determinadas por el análisis cluster

basado en el coeficiente de similaridad de Bray-Curtis.

Coarser sandy granulometric fractions

are greater at the mouth and in the outer

areas of the ría and there is a decrease in

grain size and an increase in organic

content towards the inner areas of the

ría (TRONCOSO AND URGORRI, 1993a).

A total of 8030 individuals of mol-

luscs belonging to 116 species were col-

lected, of which 62 were gastropods, 49

bivalves, three polyplacophorans, one

scaphopod, and one cephalopod. Gas-

tropods and bivalves were the dominant

groups in terms of abundance (48.37

and 46.54%, respectively), followed by

scaphopods (4.65%). A complete list of

the collected species is provided by

TRONCOSO, URGORRI AND PARAPAR

(1993). The gastropods Nassarius reticula-

tus (Linné, 1758), N. pygmaeus (Lamarck,

1822) and Ringicula auriculata (Ménard,

1811), the bivalves Chamelea striatula (da

Costa, 1778), Pandora inaequivalvis

(Linné, 1758) and Goodallia triangularis

(Montagu, 1803), and the scaphopod

Dentalium novemcostatum Lamarck, 1818

were the most abundant species in the

ría, accounting for 50% of the total

mollusc abundance. These species

reached their highest abundances in

finer sediments excepting G. triangularis

which prefers coarser sandy sediments

(coarse and medium sand).

Multivariate analysis: Cluster analysis

and MDS ordination revealed the pres-

ence of two large groups of stations at a

12% similarity level (Figs. 2, 3): group 1,

comprised sampling stations with coarse

sediments (St. 2, 4, 5, 8, 9, 10, 11, 14, 15,

16, 17, 21, 23, 24, 25), and group 2, com-

prised bottoms of finer granulometry (St.

7, 13, 18, 19,22, 26, 31,32, 19/31 1300)

37,38, 39, 40, 41, 42, 43, 45, 46, 47, 48, 49,

50, 51, 52, 53, 54, 55, 56, 57, 58, 59, 60, 61).

Group 2 could be split into two further

groups (25% similarity level): group 3,

fine sand bottoms (St. 7, 13, 18, 19, 31, 32,

35, 36, 37, 38), and group 4, sandy

bottoms with higher content in silt/clay

(St. 33, 34, 40, 41, 42, 43, 45, 46, 47, 48, 49,

51, 52, 53, 54, 55, 56, 97, 99, 99) 00) BUS

thermore, group 4 could be split in two

subgroups (32% similarity level): 4A,

which had a greater content of fine sand

(St. 42, 49, 51, 52, 53, 54, 58, 60), and 4B,

which was comprised of muddier sedi-

ments (St. 33, 34, 40, 41, 43, 45, 46, 47, 48,

55, 56, 57, 59). MDS ordination, however,

suggests that St. 43 has a greater affinity

TRONCOSO ET AL.: Soft-bottom mollusc assemblages in the Ría de Ares-Betanzos

Stress: 0.16

Figure 3. Non-metric multidimensional scaling (MDS) ordination of mollusc assemblages in the

Ría de Ares-Betanzos. Groups derived from cluster analysis are delimited by lines.

Figura 3. Ordenación MDS de las asociaciones malacológicas en la Ría de Ares-Betanzos. Los grupos

derivados del análisis cluster estan delimitados por líneas.

for subgroup 4A in opposition to that

showed in the dendrogram.

MDS ordination also revealed that

several sampling points were displaced

from the main groupings: St. 3 was a

muddy sand basin located in the middle

of coarse sands and shows a high abun-

dance of Acanthocardia paucicostata

(Sowerby, 1841); the samples from St. 20

and St. 30 were very poor in terms of

malacofauna; St. 29 had species which

are typical of rocky bottoms. On the

other hand, St. 22, 39 and 50 appeared

displaced from the two main groups

within group 2. In addition, St. 26 and

61 show affinities with group 3 but the

former is characterized by a dominance

of the bivalve Glycimeris glycimeris

(Linné, 1758) and the latter for the pres-

ence of the bivalve Donax trunculus

Linné, 1758.

Results of the SIMPER analysis are

shown in Table I. The bivalves Goodallia

triangularis, Clausinella fasciata (da Costa,

1778) and Gar1 tellinella (Lamarck, 1818)

are the species with a greater contribu-

tion to similarity (up to a cumulative

70%) for coarser sandy bottoms of

group 1. Group 3 is mainly determined

by Chamelea striatula and Nassarius retic-

ulatus. Group 4A is defined by Ringicula

auriculata, Pandora inaequivalvis, N. retic-

ulatus and C. striatula. Group 4B is char-

acterized by Nassarius pygmaeus, N. retic-

ulatus, Nucula nitidosa Winckworth,

1930, C. striatula, Dentalium novemcosta-

tum, R. auriculata, Corbula gibba (Olivi,

1792), P. inaequivalvis and Montacuta

phascolionis Dautzenberg and Fisher,

1925;

The BIO-ENV procedure (Table II)

showed that the best combinations of

environmental variables via the highest

correlations with faunistic data was that

composed of depth, gravel, fine sand

and silt/clay. Depth was the variable

with the best value when each variable

was considered alone (pw: 0.464). The

Iberus, 23 (2), 2005

Table I. Results of SIMPER analysis. Species were ranked according to their average contribution to

similarity between assemblages in the Ría de Ares-Betanzos. Average abundance, ratio value (simi-

larity/standard deviation, Sim./SD), and percentage of cumulative similarity were also included.

Tabla I. Resultados del análisis Simper. Los rangos de las especies están organizados de acuerdo con la contribución

a la similaridad entre los grupos de estaciones en la Ría de Ares-Betanzos. Se incluye la abundancia media, valor

del ratio” (similaridad/desviación estándar, Sim./SD), y el porcentaje de la similaridad acumulada.

Group 1 (average simil.: 26.01%) Av.Abund. Av.Sim. Sim./SD. Contrib.% Cum.%

Goodallia triangularis (Montagu, 1803) 28.53 1.78 0.88 29.92 29.92

Clausinella fasciata (da Costa, 1778) 13.87 7.40 1.00 28.44 58.36

Gari tellinella (Lamarck, 1818) 3.87 29 0.71 1019 69.55

Spisula elliptica (Brown, 1827) 0.73 0.95 0.35 3.65 73.20

Timoclea ovata (Pennant, 1777) 6.33 0.80 0.28 3.06 76.26

Tellina donacina Linné, 1758 2.13 0.77 0.29 2.98 79.24

Retusa mammillata (Philippi, 1836) 1.07 0.74 0.28 2.84 82.07

Caecum trachea (Kanmacher, 1798) 1278 0.60 0.27 34 84.40

Caecum glabrum (Montagu, 1803) 4.00 0.56 0.27 2.14 86.54

Euspira pulchella (Risso, 1826) 0.40 0.51 0.29 1.96 88.50

Obtusella intersecta (Wood, 1857) 173 0.48 0.22 1.86 90.36

Group 3 (average simil.: 28.74%) Av.Abund. Av.Sim. Sim./SD Contrib.% Cum.%

Chamelea striatula (da Costa, 1778) 21.33 13.01 1.34 45.27 45.27

Nassarivs reticulatus (Linné, 1758) 15.50 8.13 2.04 28.30 13.56

Thracia papyracea (Poli, 1791) 7.50 1.65 0.37 5.14 19.30

Tellina fabula Gmelin, 1791 3.50 1.44 0.46 5.01 84.31

Mysella bidentata (Montagu, 1803) 3 1.37 0.53 4.78 89.09

Turbonilla acuta (Donovan, 1804) 9.50 0.94 0.32 3.26 92.35

Group 4A (average simil.: 43.38%) Av.Abund. Av.Sim. Sim./SD Contrib.% Cum.%

Ringicula auriculata (Ménard, 1811) 49.10 12.76 1.59 29.42 29.42

Pandora inaequivalvis (Linné, 1758) 32.80 11.56 2.22 26.05 56.07

Nassarius reticulatus (Linné, 1758) 5.00 3.46 1.01 1.98 64.04

Chamelea striatula (da Costa, 1778) 12.80 3.3) 0.66 1.18 11.82

Mactra stultorum (Linné, 1758) 2.40 2.61 1.18 6.01 17.84

Turbonilla acuta (Donovan, 1804) 3.70 2.03 0.75 4.68 82.51

Spisula subtruncata (da Costa, 1778) 3.80 1.96 0.69 451 87.02

Nassarius pygmaeus (Lamarck, 1822) 11.40 1.37 0.37 9.15 90.17

Group 4B (average simil.: 44.86%) Av.Abund. Av.Sim. Sim./SD Contrib.% Cum.%

Nassarius pygmaeus (Lamarck, 1822) 34.58 5.98 1.27 13.33 13.33

Nassarius reticulatus (Linné, 1758) 26.58 5.18 2.43 11.55 24.88

Nucula nitidosa Winckworth, 1930 16.25 4.61 2.81 10.28 35.16

Chamelea striatula (da Costa, 1778) 15.83 3.94 133 8.77 43.94

Dentalium novemcostatum Lamarck, 1818 30.92 3.07 0.94 6.85 50.79

Ringicula auriculata (Ménard, 1811) 9.83 2.82 1.58 6.30 57.08

Corbula gibba (Olivi, 1792) 18.75 2.13 1.14 6.09 63.17

Pandora inaequivalvis (Linné, 1758) 8.42 2.21 0.89 4.92 68.09

Montacuta phascolionis Dautzenberg 8. Fisher, 1925 9.83 1.65 0.88 3.08 A

Odostomia unidentata (Montagu, 1803) 14.50 1.54 1.05 3.42 15.20

Chrysallida indistincta (Montagu, 1808) 10.58 1.46 0.92 3.26 18.46

Philine aperta (Linné, 1767) 6.42 1632 0.70 2.94 81.40

Volvulella acuminata (Bruguiére, 1792) 3.33 1.28 105 2.85 84.25

Acanthocardia paucicostata (Sowerby, 1841) 10.58 127 0.94 DE 86.97

Ihyasira flexuosa (Montagu, 1803) 4.97 0.66 0.60 1.47 88.44

Ondina diaphana (Jeffreys, 1848) 3.58 0.62 0.63 1.39 89.83

Abra alba (Wood, 1802) 3.67 0.60 0.58 1.34 91.17

TRONCOSO ET AL.: Soft-bottom mollusc assemblages in the Ría de Ares-Betanzos

Table II. Best combinations of variables obtained through BIO-ENV analysis according the values

of the Spearman's rank correlation (pw) for the Ría de Ares-Betanzos. GR, gravel; CS, coarse sand;

ES, fine sand; Q50, median grain size; C/N, Carbon-Nitrogen ratio.

Tabla 11. Mejores combinaciones de variables obtenidas a través del análisis BIO-ENV de acuerdo con

los valores del coeficiente de correlación de rango de Spearman (pw) para la Ría de Ares-Betanzos. GR,

grava; CS, arena gruesa; FS, arena fina; Q50, mediana del tamaño de grano; C/N, relación Carbono-

Nitrógeno.

Number of variables Correlation (pw)

2 0.508

3 0.511

4 0.518

0.515

5 0.516

0.516

0.514

' 0.516

0.515

0.512

All 13 variables: 0.410

Best variable combination

Depth-FS

Depth-GR-FS

Depth-GR-FS-Silt/Clay

Depth-Qso-GR-FS

Depth-Qso-GR-FS-Silt/Clay

Depth-GR-CS-FS=Silt/ Clay

Depth-C/N-CSFS<Silt/Clay

Depth-Qso-GR-ES-FSSSilt/Clay

Depth-C/N-GR-ESFS<Silt/Clay

Depth-Q50-C/N-GR-FS-Silt/Clay

Table IM. Summary of canonical correspondence analysis (CCA) for the Ría de Ares-Betanzos.

Tabla III. Resumen del análisis de correspondencias canónicas (CCA) para la Ría de Ares-Betanzos.

Axes |

Eigenvalues 0.701

Species-environment correlations 0.927

Cumulative percentage variance

of species data: 12.1

of species-environment relation 34.4

Sum of all unconstrained eigenvalues

Sum of all canonical eigenvalues

forward selection of CCA selected depth

and gravel as the variables explaining

most of the variance in the species data

(p<0.01), while fine sand and median

grain size were found significant only at

the 5% level. Axes I and II accumulate

16.8% of the species variance and 47.9%

of species-environment variance (Table

II). Depth, median grain size, fine sand

and silt/clay showed the highest corre-

lations with axis l; correlations with the

other axes were less significant. Sam-

pling stations appeared distributed from

the right to the left of axis I following an

[ 1 IV Total inertia

0.274 0.212 0.167 5.784

0.869 0.824 0.883

16.8 20.5 23.4

47.9 58.3 66.5

5.784

2.035

increase in content of fine sand and

silt/clay and a decrease in depth and

median grain size (Fig. 4). The same

pattern can be observed in the MDS

ordination with superimposed values of

depth, gravel, fine sand and silt/clay

(Fig. 5). Thus, the different analyses sug-

gested that distribution of fauna in the

study area is mainly related to a depth

and grain size gradient.

Description of assemblages: Distribu-

tion in the Ría de Ares-Betanzos of the

mollusc assemblages determined by

multivariate analyses are shown in

Iberus, 23 (2), 2005

31/39

42 ——— | 19 18274

4 22

56 so Gb 49 4 21

57 Carbonates Depth

53==33 4

EQ : 3

e y ; SMS 44 2 e

59 47

1044

TOM C 46

55)

Figure 4. Canonical correspondence analysis (CCA) ordination of stations and environmental vari-

ables relative to axes 1 and II for the Ría de Ares-Betanzos. GR, gravel; CS, coarse sand; MS,

medium sand; ES, fine sand; Q50, median grain size; So, sorting coefficient; TOM, total organic

matter; C, organic carbon; N, nitrogen; C/N, Carbon-Nitrogen ratio.

Figura 4. Ordenación de las estaciones de muestreo y variables ambientales de la Ría de Ares-Betanzos

para los ejes I y II del análisis de correspondencias canónicas (CCA). GR, grava; CS, arena gruesa; MS,

arena media; ES, arena fina; (Q50, mediana del tamaño de grano; So, coeficiente de selección; TOM,

materia orgánica total; C, carbono orgánico; N, nitrógeno; C/N, relación Carbono-Nitrógeno.

Figure 6 and their environmental and

faunistic characteristics in Table IV.

Group 1 is comprised of the deepest

bottoms of the ría; sediments are mostly

composed of coarse sand, medium sand

and gravel with a high carbonate

content. The assemblage is numerically

dominated by Goodallia triangularis,

Caecum trachea (Kanmacher, 1798) and

Clausinella fasciata; other characteristic

species were Gar tellinella, Timoclea ovata

(Pennant, 1777) and Caecum glabrum

(Montagu, 1803), which are distributed

almost exclusively in these bottoms.

In Group 3, sediment are composed

of fine sand and median sand and has a

higher amount of silt/clay and carbon-

ate content than group 1. The species

composition of these bottoms indicates a

transition between the fauna of coarser

sediments (group 1) and muddy

sand/muds (groups 4A, 4B). This

assemblage is characterized by a high

abundance of Chamelea striatula, Nassar-

tus reticulatus and Turbonilla acuta

(Donovan, 1804) and by the presence of

Thracia papyracea and Mysella bidentata.

Mollusc abundance per station is lower

than in coarse sands and total species

number is greater (49 vs 38). The MDS

ordination showed that stations 22 and

61 have certain affinities with this

TRONCOSO ET AL.: Soft-bottom mollusc assemblages in the Ría de Ares-Betanzos

Figure 5. Non-metric multidimensional scaling (MDS) ordination of mollusc assemblages in the

Ría de Ares-Betanzos with superimposed values of the abiotic variables selected by BIO-ENV

analysis. A: depth; B: gravel; C: fine sand; D: silt/clay. Circle size is proportional to values of vari-

ables in each sampling station.

Figura 5. Ordenación MDS de las asociaciones malacológicas en la Ría de Ares-Betanzos con los valores

superpuestos de las variables abióticas seleccionadas por el análisis BIO-ENV. A: profundidad; B: grava;

C: arena fina; D: limoslarcillas. El tamaño de los círculos es proporcional al valor de las variables en

cada estación de muestreo.

assemblage, although the presence of

Tellina fabula Gmelin, 1791 and Donax

trunculus also suggest a mix with the

faunal assemblage from intertidal sedi-

ments.

Group 4A is spread in shallower

bottoms which have a greater fine sand

and silt/clay content. Total abundance

and species number per station are

higher than in the previous groups. The

dominant species are Ringicula auriculata

and Pandora inaequivalvis, and several

species, such as Chamelea striatula and

Nassarius reticulatus, were shared with

group 3 although in lower abundances

and with a more irregular presence. On

the other hand, the bivalves Mactra stul-

torum and Spisula subtruncata were

mostly found in this assemblage.

Sediments in group 4B are slightly

muddier than those of group 4A and

have a greater organic content. Domi-

nant species in terms of abundance were

Nassarius pygmaeus, Dentalium novem-

costatum and Nucula nitidosa; other char-

acteristics species were Corbula gibba,

Nassarius reticulatus, Acanthocardia pauci-

costata, Chamelea striatula, Ringicula

auriculata and Pandora inaequivalvis,

although the latter two were less abun-

dant than in group 4A.

DISCUSSION

According to our analyses, the distri-

bution of the molluscan fauna in the Ría

de Ares-Betanzos seems to be primarily

determined by gradients in depth and

grain size. The latter was characterized

by an increase in fine sand and silt/clay

content from the mouth of the ría

towards the inner margins. This sedi-

mentary gradient is related to tidal

current systems and interactions

between Oceanic and continental water

(SÁNCHEZ-MATA ET AL., 1999). The pres-

ence of coarser sediments in the mouth

is due to a stronger hidrodynamism

(TRONCOSO ET AL., 1993) while a deposi-

Iberus, 23 (2), 2005

.o..

po .:

Rocky bottoms

Muddy sand basin

Group 1: Venus fasciata community

Group 3: Venus gallina community

Group 4: Abra alba community

Group 4A: Ringicula auriculata-

Pandora inaequivalvis facies

Group 4B: Nassarius pygmaeus-

Dentalium novemcostatum facies

e. .

...

. »

Figure 6. Spatial distribution of mollusc assemblages in the Ría de Ares-Betanzos as determined

through cluster analysis.

Figura 6. Distribución espacial de las asociaciones malacológicas en la Ría de Ares-Betanzos determi-

nadas por el análisis cluster.

tion of finer fractions occurs in inner,

sheltered areas. This relationship of

depth and sedimentary composition

with faunal distribution was also

observed by a large number of authors

(RHOADS AND YOUNG, 1970; GRAay, 1974;

EVANS AND TALLMARK, 1976; GLÉMAREC,

1978; 'TUNBERG, 1981).

According to their faunistic compo-

sition and environmental features, the

mollusc assemblages in the Ría de Ares-

Betanzos could be defined using the

classic terms of “community” and facies”

(PETERSEN, 1918; THORSON, 1957). Thus,

group 1 has a fauna that could be

included among the different varieties

of the “Branchiostoma lanceolatum-Venus

fasciata community” (IHORSON, 1957).

Several authors have reported the pres-

ence of similar faunal associations in

other areas of Galicia such as Ría da

Coruña (LÓPEZ-JAMAR AND MEJUTO,

1985) and Ensenada de Baiona

(MOREIRA, QUINTAS AND TRONCOSO,

2005). These bottoms have clean coarse

sediments with a high content of bio-

genic carbonates and are located at the

outer areas of the rías where the hydro-

dynamism is stronger (NOMBELA, VILAS,

RODRÍGUEZ AND ARES, 1987). The fauna

present in group 3 agrees with the

description of the “Venus gallina commu-

TRONCOSO ET AL.: Soft-bottom mollusc assemblages in the Ría de Ares-Betanzos

Table IV. Summary of biotic and physical characteristics of the four molluscan assemblages in the

Ría de Ares-Betanzos determined through cluster analyses (values: mean + standard deviation).

Dominant species in any given assemblage are those which account for 275% of total abundance.

Tabla IV. Resumen de las características bióticas y físicas de las cuatro asociaciones malacológicas de la

Ría de Ares-Betanzos determinadas a traves del análisis cluster (valores: media + desviación estándar).

Las especies consideradas como dominantes en cada una de las asociaciones fueron áquellas que represen-

taron 275% de la abundancia total.

Group | Group 3 Group 4A Group 4B

Dominant species Goodallia triangularis Chamelea striatula Ringicula auriculata—— Nassarius pygmaeus

Clausinella fasciata Nassarivs reticulatus Pandora inaequivalvis — Dentalium novencostatum

Caecum trachea Turbonilla acuta Chamelea striatula Nassarius reticulatus

limoclea ovata Thracia papyracea Nassarius pygmaeus — Corbula gibba

Chrysallida decussata Nucula nitidosa

Chamelea striatula

Odostomia unidentata

Calyptraea chinensis

Acanthocardia paucicostata

Chrysallida indistincta

Ringicula auriculata

Montacuta phascolionis

N 89.93+87.65 19.67453.61 172.22+114.38 286.50+179.15

S 8.86+5.08 8.67+3.70 12.44+3.68 22.9246.89

Depth 27.35+8.33 13.21+2.58 5.78+3.10 1.21+2.30

Y Gravel 18.29+23.63 1.25+2.56 0.07+0.10 1.85+2.14

Y Coarse sand 52.04+25.62 9.12+16.16 2.29+2.59 1.3348.17

% Medium sand 18.76+19.65 20.39+16.16 9.84+14.77 10.09+8.17

% Fine sand 10.51+18.33 60.53+17.82 73.20+15.70 64.78+11.68

% Silt/Clay 0.39+0.60 8.7147.91 14.60+11.11 15.96+8.76

Q5o (mm) 0.87+:0.59 0.17+0.07 0.13+0.05 0.14+0.05

Sedimentary type Coarse//medium sand Fine/muddy sand Fine/muddy sand Muddy sand/Sandy mud

Y% TOM 0.14+0.19 0.44+0.18 0.86+0.48 1.130.77

% Carbonates 38.47+21.92 40.49+8.66 29.50+6.48 28.15+13.7

nity” (THORSON, 1957), corresponding to

the fine sand bottoms of the center and

northern outer margin of the ría in

which the bivalve Chamelea striatula and

the gastropod Nassarius reticulatus show

their greater abundance. Both species

are also spread in shallower finer sandy

bottoms although in lower abundances.

Even though group 4 have some

species typical of the “Venus gallina com-

munity” such as Chamelea striatula and

Mactra stultorum, there is an important

presence of several other species which

show preference for muddier sediments

(Corbula gibba, Thyasira flexuosa, Nucula

nitidosa). Thus, this group could be con-

sidered as a mix between the already

mentioned community and the “Syndos-

mia (=Abra) alba community” of PE-

TERSEN (1918). This situation agrees with

the results showed by Sánchez-MATA

AND MORA (1999b) for all groups of

macrofauna in the Ares sector of the ría.

On the other hand, the “Abra alba com-

munity” has been reported along Euro-

pean coasts in different types of muddy

bottoms (REES AND WALKER, 1983; GEN-

TIL, ÍIRLINGER, ELKAIM AND PRONIEWSKI,

1986; LASTRA, MORA, SÁNCHEZ AND

TRONCOSO, 1988) as well as in Galician

Iberus, 23 (2), 2005

rías (CADÉE, 1968; OLABARRÍA, URGORRI

AND TRONCOSO, 1998; SÁNCHEZ-MATA

AND MORA, 1999b; MOREIRA ET AL.,

2005). However, multivariate analyses

have distinguished two further groups

within group 4, which can be character-

ized as two different malacological “fa-

cies”. The facies corresponding to group

4A is determined by the dominance of

Ringicula auriculata and Pandora inae-

quivalvis, and that present in group 4B is

characterized by Nassarius pygmaeus and

Dentalium novemcostatum and shows a

greater presence of species preferring

muddier sediments. Several authors

have suggested that the different pro-

portion of sand and silt/clay is a major

factor in structuring benthic communi-

ties (RHOADS AND YOUNG, 1970; GRAY,

1974). WEBB (1969) pointed out that

even a small silt/clay content affects the

sediment porosity and therefore the fau-

nal composition. In the Ría de Ares-Be-

tanzos, the variations in amount of fine

sand and silt/clay seems to condition

mollusc species abundance across these

bottoms and consequently the presence

of any given facies.

In general, mollusc distribution in

sandy sediments of the outer and

central part of the ría is similar to those

observed in Ría de Coruña (LÓPEZ-

JAMAR AND MEJUTO, 1985) and Ense-

nada de Baiona (MOREIRA ET AL., 2005).

The transition between the “Venus

gallina and the “Abra alba” communities

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ACKNOWLEDGEMENTS

The authors want to express their

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O Sociedad Española de Malacología —__—_——— lIberus, 23 (2): 39-46, 2005

Estado de la especie amenazada Patella ferruginea Gmelin,

1791 (Gastropoda: Patellidae) en la bahía de Algeciras y

Gibraltar

Status of the endangered limpet Patella ferruginea Gmelin, 1791

(Gastropoda: Patellidae) in the Algeciras bay and Gibraltar

Free ESPINOSA*, Darren A. FA** y Terence M. J. OCAÑA*+**

Recibido el 28-IV-2005. Aceptado el 1-VI11-2005

RESUMEN

El molusco marino Patella ferruginea está considerado como una de las especies en mayor

peligro de extinción del Mediterráneo. En 2004 se ha llevado a cabo un estudio para esta-

blecer la distribución, abundancia y talla de la especie en la bahía de Algeciras. Un total de

140 individuos fueron censados sobre una línea de costa de unos 20 km, presentando los

mayores valores a lo largo de la costa este de la bahía. Esto representa la mayor población

conocida de esta especie en las costas de la península Ibérica. Aunque se encontraron un

mayor número de ejemplares en construcciones artificiales, como diques, las mayores densi-

dades aparecieron en sustratos naturales. Las diferencias entre ambos tipos de sustratos no

fueron significativas, aunque esto pueda deberse al bajo número de ejemplares encontrados

en comparación con otras áreas mediterráneas del norte de Africa. La distribución normal de

tallas encontrada indica que el área muestreada podría albergar una población bien esta-

blecida y reproductora. Se requieren urgentemente nuevos estudios para establecer progra-

mas de gestión y conservación para esta especie, dado su elevado riesgo de extinción.

ABSTRACT

The marine mollusc Patella ferruginea is considered one of the species in greatest danger of

extinction in the Mediterranean. A study was carried out to establish the distribution extent,

abundance and size of this species in the Bay of Algeciras (Strait of Gibraltar) in 2004. A

total of 140 individuals were found over approximately 20km of coastline, with the highest

numbers occurring along the eastern shores of the Bay. This represents the largest known popu-

lation of this limpet species along the coast of Iberia. Although higher overall numbers were

encountered along artifical constructions such as breakwaters, highest densities occurred on

natural rocky substrates. Differences between both types of substrates were not significant, pro-

bably due to the relatively low overall numbers encountered. A normal distribution of body

sizes was found in this study, indicating that the area sampled could contain a wellestablis-

hed breeding population. New studies are urgently required to establish and implement mana-

gement programmes and conservation plans for this species due its high risk of extinction.

PALABRAS CLAVE: Patella ferruginea, conservación, bahía de Algeciras, Gibraltar, lapas.

KEYWORDS: Patella ferruginea, conservation, Algeciras bay, Gibraltar, limpets.

* Laboratorio de Biología Marina. Avda. Reina Mercedes, 6. 41012. Sevilla. Spain. e-mail: freeVus.es

** Gibraltar Museum. 18-20 Bomb House Lane. Gibraltar. United Kingdom.

*** Institute of Mediterrenean Ecology, 7F Malvasia, Vineyards, Gibraltar, United Kingdom.

50)

Iberus, 23 (2), 2005

INTRODUCCIÓN

El molusco marino Patella ferruginea

Gmelin, 1791, es una especie que se

encuentra protegida tanto por las leyes

de la Unión Europea (Directiva

92/43/CEE de 21 de mayo de 1992 en el

Anexo IV “especies animales y vegetales

de interés comunitario que requieren

una protección estricta”), así como por

la legislación española (Orden de 9 de

junio de 1999. Especie considerada en

“peligro de extinción”) y gibraltareña

(Government of Gibraltar, Nature Pro-

tection Ordinance 1991). Está conside-

rada como una de las especies marinas

mediterráneas más amenazadas de

extinción (LABOREL-DEGUEN Y LABOREL,

1991a) y su área de distribución se ha

visto reducida a unas pocas localidades

en el Mediterráneo occidental en los

últimos tiempos (FISCHER-PIETTE, 1959;

LABOREL-DEGUEN Y LABOREL, 199%la;

CRETELLA, SCILLITANI, TOSCANO,

TURELLA, PICARIELLO Y CATAUDO, 1994;

TEMPLADO, CALVO, GARVÍA, LUQUE,

MALDONADO Y MOZO, 2004).

En la península Ibérica las citas

recientes (a partir de los años 80 del

siglo XX) son escasas. La primera cita

escrita reciente de la especie en las

costas peninsulares españolas ha sido

aportada por GRANDFILS (1982) y

GRANDFILS Y VEGA (1982) que mencio-

nan 17 ejemplares en la costa de Málaga.

Después, GARCÍA-GÓMEZ (1983) la cita

como rara en la bahía de Algeciras,

mientras que MORENO (1992) encontró

dos ejemplares en Cabo de Gata

(Almería). También Fa (1990), encontró

un solo individuo en el puerto de

Gibraltar durante 1989. Más reciente-

mente, varios ejemplares fueron detecta-

dos en el puerto de Gibraltar por Tem-

plado en 1995 (comunicación personal

en RAMOS, 1998) y TEMPLADO, Fa Y

OCAÑA en 2002 (comunicación perso-

nal), así como un ejemplar aislado en

Punta Carnero (Fa, 1998). Según TEM-

PLADO Y MORENO (1997), la especie se

encuentra prácticamente extinguida de

las costas continentales europeas,

situándola al borde de la desaparición

en las costas del sur de la península,

dónde persistiría en puntos aislados. Es

de gran interés describir y cuantificar la

estructura de las poblaciones de Patella

ferruginea allí dónde aún persisten

(PARACUELLOS, NEVADO, MORENO,

GIMÉNEZ Y ALESINA, 2003). Por ello, el

presente estudio tiene como objetivo

cuantificar y describir la población pre-

sente en la bahía de Algeciras, represen-

tando el primer estudio poblacional

sobre Patella ferruginea llevado a cabo en

las costas de la península Ibérica. El

conocimiento de las poblaciones aún

existentes es fundamental para imple-

mentar medidas de conservación enca-

minadas a evitar la crítica situación de

la especie, situación que roza la extin-

ción total, si nos referimos a las costas

peninsulares.

MATERIAL Y MÉTODOS

En base a las informaciones previas,

la bahía de Algeciras podía albergar

alguna población de Patella ferruginea,

por ello se centró el estudio en todo el

arco de todo este golfo natural, desde

punta Carnero (36 04,600" N - 5? 25,460'

O) como límite occidental, hasta la zona

este del peñón de Gibraltar (36* 07,698'

N - 5% 20,477" O) como límite oriental. Se

visitaron en marzo de 2004 todas las

localidades susceptibles de albergar

poblaciones de esta lapa, éstos fueron

tanto en sustratos rocosos naturales

como diques artificiales que se encontra-

ban presentes en el área de estudio. Las

zonas de playa fueron excluídas del

muestreo. En cada localidad se realizó el

censo de todos los ejemplares observa-

dos durante la bajamar, anotando la

talla del eje anteroposterior de la concha

con un calibre y registrando la posición

de cada localidad mediante un GPS

Garmin 45XL. Para el cálculo de la den-

sidad se estimó la distancia muestreada

en cada localidad utilizando la carta

naútica 445A del Instituto Hidrográfico

de la Marina. En la Figura 1 se pueden

apreciar las localidades de muestreo,

mientras que en la Tabla 1 se detallan las

coordenadas geográficas de cada una.

Adicionalmente, durante el presente

ESPINOSA ET AL.: Patella ferruginea en la bahía de Algeciras y Gibraltar

Península

Ibérica

ALGECIRAS

Provincia de Cádiz

Acerinox Mayorga

Jl pe

San Felipe Pz

North Mole4” y-

Europort

GIBRALTAR

Mota Ta S East Side

SA)

Little Bay

San García

La Ballenera Europa Point

“3 Punta Carnero

Mar Mediterráneo

Figura 1. Mapa de la bahía de Algeciras y Gibraltar donde se indican las localidades de muestreo.

En trama punteada se denotan las zonas de playa.

Figure 1. Map of Algeciras Bay and Gibraltar, showing the sampling localities. Shaded areas indicate

beach zones.

estudio se prospectaron algunas otras

localidades en el sur peninsular.

Los datos obtenidos fueron someti-

dos a un test no paramétrico Kruskal-

Wallis de comparación de medias, ya

que no se verificó normalidad de los

datos con el test de Shapiro-Wilk. Estos

tratamientos estadísticos fueron realiza-

dos con el Biomedical Statistical

Package (BMDP) (Dixon, 1983).

RESULTADOS

El número total de ejemplares detec-

tados en el presente estudio fue de 140.

Este contingente aparecía, sin embargo,

distribuído de forma muy irregular (Fig.

2). La zona este de la bahía de Algeciras

(Gibraltar y La Línea: desde East Side hasta

San Felipe) albergaban la mayor cantidad

de ejemplares (114), mientras que en la

zona oeste (desde Puente Mayorga a San

García), sólo se encontraron 26 ejempla-

res. En las restantes localidades muestre-

adas no apareció ningún ejemplar. Aten-

diendo a la densidad obtenida, los resul-

tados son similares, presentando los

mayores valores la zona externa a la bahía

hacia el Mediterráneo (Fig. 3).

Al comparar la densidad en sustrato

natural frente a la encontrada en sustrato

artificial, se observa como la media en

sustrato natural fue de 0,14 ind./m

(+0,20), mientras que en el caso del sus-

trato artificial fue de 0,08 ind./m (+0,04),

prácticamente el doble. No obstante no

se apreciaron diferencias estadística-

mente significativas mediante el test no

paramétrico de Kruskal-Wallis entre

ambas situaciones (K=0,06; p=0,80),

debido a la elevada desviación estándar

obtenida, ya que existía una gran varia-

bilidad entre localidades (Fig. 4).

La estructura de tallas de la pobla-

ción se ajustaba a una distribución

Iberus, 23 (2), 2005

Tabla I. Coordenadas geográficas de cada una de las localidades muestreadas. Los asteriscos indican

sustrato natural.

Table I. Geographic coordinates of each sampled locality. Asteriscs indicate natural substrates.

Localidad

Eost Side*

Europa Point*

Little Bay

Rosia Bay

North Mole

Europort

Dique San Felipe

Puente Mayorga?

Acerinox

Puerto del Saladillo

Punta San García*

La Ballenera

Punta Camero*

normal (W=8848; p=0,1413), previa

transformación de los datos con la raíz

cuadrada (Fig. 5). La mayor frecuencia

aparecía en la clase 5-6 cm, siendo la

clase 4-5 cm la que presentaba a conti-

nuación una frecuencia más elevada. No

se detectaron individuos menores de 2

cm.

Por otro lado, fuera del área de la

bahía de Algeciras no se detectaron indi-

viduos de la especie ni en escolleras del

puerto de la Atunara (La Línea, ya en

zona mediterránea), ni en diferentes

escolleras en Benalmádena (Málaga), sin

embargo se encontró un ejemplar de 13

mm en roquedos naturales del faro de

Calaburras (Málaga), dónde se prospec-

taron unos 500 m de línea de costa tra-

tando de localizar ejemplares en base a

citas previas de la especie (GRANDFILS,

1982, GRANDFILS Y VEGA, 1982). Igual-

mente se detectó otro ejemplar aislado

en la escollera exterior de Puerto Banús

(Marbella), con una talla entre 2 y 3 cm.

DISCUSIÓN

La población encontrada es de gran

interés, pues supone el mayor contin-

gente de Patella ferruginea en la penín-

sula Ibérica. La especie había sido citada

Coordenadas

36" 07,698" N- 5* 20,477'0

36" 06,511'N - 5% 20,/77'0

36" 06,815" N-5* 21,025'0

36" 07,247'N -5* 21,154'0

36" 08,760" N - 5* 21,890'0

36" 08,500"N - 5 21,510" 0

36" 09,309'N - 5* 21,671" 0

36" 10,841'N - 5* 24,312'0

30 10,517 5525590750

36" 07,010'N - 5* 26,141'0

36" 06,330" N - 5* 25,850'0

36" 04,990"N - 5* 25,530" 0

36" 04,600" N - 5* 25,460" 0

en la bahía de Algeciras (localidad de

Campamento y Los Roncadillos, entre el

Club Naútico de La Línea de la Concep-

ción y la desembocadura del río Guada-

rranque) por GARCÍA-GÓMEZ (1983), sin

embargo, muchas especies han desapa-

recido en esta zona en los últimos años

debido a la elevada presión antrópica

que sufre (ver SÁNCHEZ-MOYANO,

ESTACIO, GARCÍA-ADIEGO Y GARCÍA-

Gómez, 1998). La especie también había

sido citada en las costas de Málaga y

Granada (punta Chullera, zona de Fuen-

girola-Calaburras, paseo marítimo de

Málaga y en la Punta de la Mona, en la

localidad granadina de La Herradura)

por GRANDFILS (1982), GRANDFILS Y

VEGA (1982) y LUQUE (1986). Así mismo,

Barrajón encontró en 1994 un ejemplar

vivo en los acantilados de Maro, en el

límite entre las provincias de Málaga y

Granada (com. pers. en TEMPLADO,

2001). No obstante, TEMPLADO (2001) no

encontró ejemplares vivos en la visita

realizada en 1995 a las localidades mala-

gueñas dónde había sido citada. Proba-

blemente se traten de poblaciones muy

fragmentadas, formadas por escasos

ejemplares aislados, difíciles de detectar,

puesto que la especie parece que conti-

núa presente en la zona de Málaga, de

acuerdo a los ejemplares encontrados en

ESPINOSA ET AL.: Patella ferruginea en la bahía de Algeciras y Gibraltar

S.Garcia

Saladillo

ES Acerinox

3 Pte. Mayorga

E San Felipe

O Rosia Bay

- :

Little Bay

Europa Point

East Side

USOS 202350 3 AD 4950.

2 N0 de ejemplares

E gos

: 3

Y 0

= DS

y OR

E 220

Ez 3 1

= O

a EA

A Sustrato natural Sustrato artificial 5

S.Garcia

Saladillo

Acerinox

Pte. Mayorga

San Felipe

Rosía Bay

Little Bay

Europa Point

East Side

0 0,1 0,2 0,3 0,4 0,5

3 Densidad (ind./m)

0-1 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10

Talla (cm)

Figura 2. Número de ejemplares de 2 ferruginea por localidad. Figura 3. Densidad de ejemplares

de P ferruginea por localidad. Figura 4. Número de ejemplares de P ferruginea en sustrato natural y

artificial. Figura 5. Distribución de frecuencias de talla del conjunto de la población.

Figure 2. Number of individuals of P. ferruginea per locality. Figure 3. Density of individuals of P.

ferruginea per locality. Figure 4. Number of individuals of P. ferruginea on natural and artifical subs-

trate. Figure 5. Size frequency distribution for the whole population.

el faro de Calaburras y en Puerto Banús

durante el presente estudio. Todo parece

indicar que se trata de individuos aisla-

dos (ya que se prospectó toda el área

adyacente sin detectar más ejemplares),

reductos de poblaciones existentes en

tiempos recientes o producto del asenta-

miento de alguna larva procedente de

otras localidades del área mediterránea.

En este sentido LABOREL-DEGUEN Y

LABOREL (1993) sostienen la hipótesis de

que ciertos ejemplares relictos de las

costas continentales francesas podrían

proceder de contingentes larvarios de

Córcega. Así, teniendo en cuenta la

corriente atlántica superficial que desde

el Estrecho se dirige hacia el este para-

lela a la costa de Málaga a una veloci-

dad de 20 cm/seg. (ARÉVALO Y GARCÍA,

1983), los ejemplares aislados aún pre-

sentes en esta área podrían proceder de

la población presente en la bahía de

Algeciras, alcanzando, en poco menos

de seis días, la zona del faro de Calabu-

rras, de acuerdo al modelo de poblacio-

nes donantes (donadoras de contingen-

tes larvarios) frente a poblaciones recep-

toras (no viables y mantenidas por

aportes externos) (BROWN Y KODRICK-

BROwN, 1977; HoLr, 1985; PULLIAM,

1988). En la costa oriental de Andalucía

la última cita escrita de la especie en la

península Ibérica (MORENO, 1992)

procede de Cabo de Gata (Almería),

aunque los dos ejemplares encontrados

desaparecieron posteriormente, y la

especie no ha vuelto a ser detectada en

las costas de Almería (Moreno, com.

pers.). Su ausencia en la parte atlántica

(peninsular y africana) del estrecho de

Gibraltar ha sido puesta de manifiesto

por FISCHER-PIETTE (1959), Fa (1998) y

GUERRA-GARCÍA, CORZO, ESPINOSA Y

GARCÍA-GÓMEZ (2004), lo que corrobora

Á3

Iberus, 23 (2), 2005

su área de distribución endémica del

Mediterráneo occidental (CRETELLA ET

AL., 1994). Fuera de Andalucía la especie

carece de citas recientes. HIDALGO (1917)

la cita en Cadaqués, Fornells y Mahón,

no obstante estas citas deberían tomarse

con cautela pues dicho autor recibía

material y datos de muy diversas

fuentes, en muchas ocasiones sin con-

trastar; si bien en la colección de Locard

de 1892 depositada en el Museo Nacio-

nal de Historia Natural de París existen

ejemplares procedentes de Baleares. En

conclusión, se puede afirmar que la pre-

sencia de la especie en las costas penin-

sulares es absolutamente residual, sin

presentar poblaciones bien asentadas y

reproductivas, perfilándose quizá como

única excepción, la población encon-

trada en el presente estudio, la cual

podría constituir una población repro-

ductora. El hecho de que no se hayan

detectado juveniles puede deberse al

ciclo reproductor anual, ya que la libera-

ción de gametos concluye a final de

diciembre (FRENKIEL, 1975), por lo que

los juveniles podrían no ser detectados

en un recuento hasta varios meses más

tarde. Según LABOREL-DEGUEN Y

LABOREL (1991b), en estudios llevados a

cabo en las costas de Córcega, los juve-

niles aparecían en los recuentos a finales

del verano-principios del otoño, cuando

alcanzaban una talla de varios milíme-

tros.

Si bien los ejemplares aparecieron

bastante dispersos, el área objeto de

estudio no abarcaba en total más de 20

km lineales de costa, y el mayor número

de individuos se localiza entre La Línea

(espigón de San Felipe) y el peñón de

Gibraltar, quizá debido a la mayor dis-

ponibilidad de sustrato idóneo para la

especie O a la mayor influencia atlántica

de la parte occidental de la bahía de

Algeciras, con corrientes superficiales

atlánticas de entrada (ver Fa, 1998). Esta

mayor concentración de ejemplares en

la parte oriental de la bahía de Algeciras

podría posibilitar la fecundación de los

huevos (ver TEMPLADO, 2001), permi-

tiendo la persistencia de la población.

Este hecho se ve apoyado en la distribu-

ción normal de tallas observada, exis-

tiendo individuos tanto de clases de

talla menores, como mayores. Debido a

que se trata de una especie protándrica

(inicialmente macho, a partir de unos 25

mm, y luego hembra) (FRENKIEL, 1975;

TEMPLADO ET AL., 2004), la existencia de

distintas clases de talla aseguraría la

presencia futura de hembras, por creci-

miento y cambio de sexo a partir de los

machos existentes. La presencia de una

distribución con distintas clases de talla

(n=81) es considerada por Laborel-

DEGUEN Y LABOREL (1990) como una

población vigorosa y fértil. Otro tipo de

distribuciones de tallas, como las encon-

tradas por PARACUELLOS ET AL. (2003) en

la población de la isla de Alborán, con

predominancia de individuos de gran

talla (hembras), podría suponer una

dificultad para la reproducción, por la

carencia de machos, puesto que la

mayor parte de los ejemplares serían

hembras.

Respecto al sustrato, las mayores

densidades aparecen en sustrato

natural, sin embargo, en Ceuta son los

sustratos artificiales los que albergan las

mayores densidades (GUERRA-GARCÍA

ET AL., 2004). No obstante, hay que tener

en cuenta que en el presente estudio no

aparecieron diferencias significativas, en

parte debido al bajo número de ejempla-

res, mientras que en Ceúta sí existían

diferencias significativas con un número

muy superior de ejemplares. En cual-

quier caso, esta diferencia puede

deberse a dos factores, en primer lugar

Ceuta presenta una costa más alterada

respecto a su fisonomía que Gibraltar, lo

que ha propiciado la presencia de gran

cantidad de sustratos artificiales que-

dando los sustratos naturales relegados

a zonas muy localizadas. En segundo

lugar, los sustratos naturales en Ceuta

son preferentemente superficies hori-

zontales poco heterogéneas (obs. pers.)

que reciben una gran irradiación solar

con respecto a los sustratos artificiales.

Sin embargo, en la bahía de Algeciras

los sustratos naturales presentaban una

mayor heterogeneidad espacial con más

zonas de sombra, especialmente en

Gibraltar, lo que podría favorecer a la

especie. En este sentido WILLIAMS Y

ESPINOSA ET AL.: Patella ferruginea en la bahía de Algeciras y Gibraltar

MORRITT (1995) encontraron un mayor

estrés térmico con aumento de la morta-

lidad en enclaves horizontales para el

patélido Cellana grata en las costas de

Hong Kong, y TAKADA (1999) señala una

mayor diversidad de gasterópodos

intermareales en enclaves umbríos del

litoral japonés.

La crítica situación de la especie en

las costas peninsulares, unida a la pre-

sencia de una población potencialmente

reproductora en la bahía de Algeciras,

hace urgente un plan de conservación

de los ejemplares encontrados para

evitar la total extinción de la especie en

la península Ibérica, así como la protec-

ción de los enclaves de sustrato natural

BIBLIOGRAFÍA

ARÉVALO, L. Y GARCÍA, T., 1983. Corrientes de la

costa de Málaga, Métodos y resultados. Informes

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nover rates in insular biogeography: effect of

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449.

CRETELLA, M., SCILLITANI, G., TOSCANO, F., Tu-

RELLA, P., PICARIELLO, O. Y CATAUDO, A.,

1994. Relationships between Patella ferruginea

Gmelin, 1791 and the other Tyrrhenian spe-

cies of Patella (Gastropoda: Patellidae). Jour-

nal of Molluscan Studies, 60: 9-17.

Dixon, W. J., 1983. BMDP Statistical Sofware.

University California Press, Berkeley. 214 pp.

Fa, D. A., 1990. A diversity-based comparative

study of the rocky coasts of Gibraltar. Under-

graduate thesis (unpublished). St. Mary's

College, University of Surrey, U.K. 198 pp.

Fa, D. A., 1998. The influence of pattern and scale

on the rocky-shore macrobenthic communities

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published). University of Southampton, U.K.

316 pp.

FISCHER-PIETTE, E., 1959. Contribution a l'eco-

logie intercotidale du Detroit de Gibraltar. Bu-

lletin de l'Institut Océanographique, 1145: 1-32.

FRENKIEL, L., 1975. Contribution a l'étude des

cycles de reproduction des Patellidae en Al-

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Napoli, 39: 153-189.

GARCÍA-GÓMEZ, J. C., 1983. Estudio comparado

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aún no perturbados donde se asienta P.

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Son necesarios nuevos estudios para

conocer la situación exacta de la especie

en el sur peninsular, para de esta forma

poder diseñar medidas urgentes de con-

servación y protección de su hábitat.

AGRADECIMIENTOS

El presente trabajo ha sido finan-

ciado por la Autoridad Portuaria de

Ceuta, así como por una beca de investi-

gación FPU del Ministerio de Educación

y Ciencia, concedida a F. Espinosa

(código AP-3556-2001).

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lla lungo da costa di Malaga (Spagna). La

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Y GARCÍA-GÓMEZ, J. C., 2004. Assessing ha-

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paña, Portugal y las Baleares. Trabajos del

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velles observations sur le population de Pa-

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ADIEGO, E. M. Y GARCÍA-GÓMEZ, J. C., 1998.

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O Sociedad Española de Malacología

Iberus, 23 (2): 47-52, 2005

The discovery of a radula in a Dentimargo species and its

taxonomic implications

Descubrimiento de la rádula en una especie de Dentimargo y sus

implicaciones taxonómicas

Franck BOYER*

Recibido el 28-IV-2005. Aceptado el 6-X-2005

ABSTRACT

The discovery of a comb-like radula in specimens of Dentimargo cf. aureocinctum (Stearns,

1872) is recorded. The occurrence of such a radula in a presumed non-radulate genus suggests

a very close relationship between the genus Dentimargo and the genus Volvarina Hinds, 1844.

This matter invalidates the taxonomic organisation currently accepted within the Marginellinae,

and especially that between the Tribes Marginellini and Prunini. The conchological difterenciation

of Dentimargo from the Volvarina-Dentimargo common stem is shown to have arisen before

the loss of the radula, and this loss is considered to have little taxonomic value within the mar-

ginellid gastropods. Due to its high conchological similarity with the type species D. dentifera

(Lamarck, 1803), D. cf. aureocinctum is conserved in the genus Dentimargo, which is provisionally

considered as being composed both of radulate and of non-radulate species.

RESÚMEN

Se reseña el descubrimiento de una rádula en forma de peine en ejemplares de Dentimargo

cf. aureocinctum (Stearns, 1872). La presencia de tal rádula en un género que se considera

carecer de la misma indica una relación estrecha entre el género Dentimargo y el género

Volvarina Hinds, 1844. Ello invalida la ordenación taxonómica actualmente aceptada entre

los Marginellinae y particularmente entre las Tribus Marginellini y Prunini. Se muestra que la

diferenciación conquiológica de Dentimargo a partir del estirpe común Volvarina-Denti-

margo se ha producido antes de la pérdida de la rádula y que esta pérdida es de escaso

valor taxonómico en gasterópodos marginéllidos. Considerando su gran semejanza con-

quiológica con la especie tipo D. dentifera (Lamarck, 1803), D. cf. aureocinctum se man-

tiene en el género Dentimargo, considerando en este tanto especies provistas de rádula

cómo careciendo de la misma.

PALABRAS CLAVE: Marginellidae, Dentimargo, Volvarina, radula, clasificación supraespecífica, Caribe.

KEY WORDS: Marginellidae, Dentimargo, Volvarina, radula, supraspecific organisation, Caribbean.

INTRODUCTION

The generic name Dentimargo Coss-

mann, 1899, based on Marginella dentifera

Lamarck, 1803 from the Mid-Eocene

(Lutetian) of the Paris Basin, is com-

monly attributed to a series of marginel-

lid species widely distributed in the

Recent throughout the tropical and the

subtropical zones from littoral to bathyal

* 110 Chemin du Marais du Souci, 93270 Sevran, France.

Iberus, 23 (2), 2005

levels. The generic placement in Denti-

margo is currently used for species with a

tall spired, rather biconic, tiny shell,

whitish shaded, generally not decorated

or poorly banded, and presenting more

or less developed labial denticles, the

uppermost one being the largest.

Defined on this basis, the genus has

been considered to be non-radulate

because a radula could not be found in

individual species checked respectively

by BARNARD (1969), PONDER (1970) and

COOVERT (1987). Commenting upon the

record of “9 species known or strongly

suspected to be non-radulate”, COOVERT

AND COOVERT (1995) considered the

whole genus Dentimargo as non-radu-

late, analogous with the genera Mar-

ginella Lamarck, 1799 and Glabella

Swainson, 1840, also claimed to be non-

radulate and considered to be closely

allied with Dentimargo. Marginella and

Glabella are defined on the basis of

having larger and thicker shells, gener-

ally not bearing a stronger upper labial

denticle and exhibiting a richer axial,

spiral or ocellate decoration.

COOVERT AND COOVERT (1995)

grouped together Dentimargo, Marginella,

Glabella and several Dentimargo-looking

genera in the tribe Marginellini Fleming,

1828, considered to be distinct from the

other Marginellinae tribes (Aus-

troginellini COOVERT AND COOVERT 1995

and Prunini COOVERT AND COOVERT,

1995), principally by their rather biconic

shell outline and by the claimed lack of a

radula (the exception being the genus

Hyalina Schumacher, 1817, which is said

to have lost its radula but placed in the

RESULTS

Prunini because of its light cylindrical

shell similar to that found in many

species of Volvarina Hinds, 1844).

BOYER (2001: 160) underlined that

“the presumed lack of a radula in the

genus Dentimargo was controlled only

about a restricted number of species and

the type species D. dentifera Lamarck,

1803, a fossil species apparently repre-

sented from the Eocene to the Miocene,

was naturally not checked for this char-

acter”. BOYER (2001: 160) also explained

that “numerous marginelliform species

from the European Eocene... constitute

a poorly differentiated Volvarina-Denti-

margo complex, in which the “comb-

like” radulae (typical of the radulate

Prunini species) or their derived forms

might be often represented”. BOYER

(2001: 160) observed that “the loss of the

radula in a uniserial rachiglossan group

is probably contracted easily, and this

derived character seems to have been

formed on several occasions in the radi-

ation of the Marginellidae. From that, it

is inferred that some Dentimargo lin-

eages may have conserved their radula,

like some Volvarina species may have

lost theirs”.

The present article is devoted to

reporting on the discovery of a comb-

like radula in a Caribbean species attrib-

utable to Dentimargo, and to the first

general taxonomic inferences that can be

issued from this discovery.

Abbreviations:

ERC: E. Rolán Collection.

FBC: Collection of the author.

Family MARGINELLIDAE Fleming, 1828

Subfamily MARGINELLINAE Fleming, 1828

Genus Dentimargo Cossmamn, 1899

Type species by original designation: Marginella dentifera Lamarck, 1803.

Dentimargo cf. aureocinctum (Stearns, 1872) (Figs. 1-3)

Marginella (Glabella) aureocincta Stearns, 1872: 22 [Type locality: Long Key, Florida].

BOYER: The discovery of a radula in a Dentimargo species and its taxonomic implications

10 um

Figures 1-3. Dentimargo cf. aureocinctum (Stearns, 1872). 1: shell from Puerto Morelos, Yucatan,

height 3.50 mm; 2: radula plate from a juvenile specimen of 3.20 mm of shell length; 3: position

of the radula seen by transparency (same specimen as in Figure 1).

Figuras 1-3. Dentimargo cf. aureocinctum (Stearns, 1872). 1: concha de Puerto Morelos, Yucatán,

altura 3,50 mm), 2: diente radular de un juvenil de 3,20 mm de longitud de concha; 3: posición de la

rádula vista por transparencia (mismo especimen que la Figura 1).

Type material: Holotype (live collected) in the United States National Museum. Not examined.

Other material examined: Banded form. Florida: 4 adult specimens, 2 juvenile shells, screening in

mud and grass, low tide, 2 feet, Tampa Bay (FBC); 2 adult specimens, hand dredged, 1-2 feet, St

Andrews Bay (FBC).

White form. Florida: 2 adult shells, Crawl Key (FBC): the squatter of both shells as D. cf. aureocinc-

tum. Yucatan: 3 adult (Figs. 1, 3) + 1 subadult + 1 juvenile specimens, 1 juvenile shell, Puerto Morelos

(FBC, ex-ERC, lot 57 M 1994): all as D. cf. aureocinctum.

Description: STEARNS (1872: 22).

COOVERT (1987: 35-37) provided a

good figure of the type-2 live animal of

D. aureocinctum from southwest Florida

(p. 36, fig. 3) and an extensive descrip-

tion of the animal external anatomy and

chromatism. Two specimens were

studied, with the adult and subadult

specimens shell lengths being 3.88 mm

and 4.36 mm respectively. The shell in

dorsal view is shown to have a slender,

biconic profile, with 2 dark narrow

spiral bands on the body whorl and 1

band on the spire whorls.

Radula: COOVERT (1987: 37) did not

“attempt to extract radula from this

species”, but on the basis of the lack of

radula displayed by PONDER (1970) in

Dentimargo cairoma (Brookes, 1924) and of

the apparent lack of radula in the Florid-

ian Dentimargo eburneola (Conrad, 1834)

checked by himself, COOVERT (1987: 37)

Iberus, 23 (2), 2005

considered that it was “quite likely that

D. aureocincta is also non-radulate”.

In the frame of this study, a radula

has been extracted from an adult (shell

length= 3.50 mm), a subadult (shell

length= 3.50 mm) and a juvenile (shell

length= 3.20 mm) specimens originating

from Yucatan (Puerto Morelos, FBC ex-

ERC) and preserved in alcohol.

Adult specimen (Figs. 1, 3): radular

extraction R-310, undetermined number

of comb-like radular plates bearing 20-

21 cusps. The radular ribbon is very

small and sub-translucent (see the size

of the ribbon at the tip of the black

arrow in Figure 3) and it was very diffi-

cult to find. The length of this ribbon

was 0,256 mm for a shell length of 32

mm. The ratio ribbon length/shell

length is of about */12.

Subadult specimen: radular extraction

R-609. The radula was observed by

transparency through the soft parts but

was lost during the extraction process

by low dissolving, due to its minute

size. The radula was lying within a

pouch situated at the distal tip of the

extended proboscis. This pouch is inter-

preted as being the buccal pouch.

Juvenile specimen: radular extraction R-

308, 54 comb-like radular plates of 20 um

of width and bearing 20-21 cusps (Fig. 2).

Distribution: The species is said to

range from Florida to Yucatan and the

Greater Antilles, but the real identity of

the tiny littoral Dentimargo species

recorded from the Greater Antilles remains

to be verified, due to the possible presence

of several similar species in this area.

Remarks: VOKES AND VOKES (1983)

record our species from Yucatan as

“Marginella (Dentimargo) aureocincta

immaculata Dall”, and they picture (pl.

18, fig. 7) a shell of 2.9 mm length resem-

bling our specimens closely (Fig. 1). This

shell is however thicker, with a strong

labrum bearing one produced upper

denticle and 4 smaller ones positioned

below the mid-part of the inner labrum.

Our 3 adult specimens have lighter

shells with a thinner labrum, one of

them bearing a singular, pronounced

upper labial denticle, the rest of the inner

labrum being smooth (Fig. 1), whereas

the 2 other adult shells show 2 faintly

distinct denticles below the pronounced

upper one. For all the other features, our

specimens perfectly match the one pic-

tured in VOKES AND VOKES (1983).

The shells from Florida show the same

general morphology and the same organ-

isation of the columellar plaits and of the

denticulated labrum (more commonly 3-

4 tiny denticles below the produced upper

one, occasionally only 2 tiny denticles or

only the larger upper one), but they present

a more slender outline with a more pointed

spire and a narrower aperture. Most of the

shells from Florida bear one honey-orange

narrow spiral band on the spire whorls

and 2 bands on the body whorl, on a light

honey to deep white background colour,

but some specimens or populations show

a full-white shell (form immaculata Dall,

1890). Intergrades between the “banded

form” and the “white form” are currently

found. The squat form represented in

Yucatan (the shell pictured in VOKES AND

VOKES, 1983 and our lot from Puerto

Morelos) is scarcely found off Florida (FBC:

1 white shell of “squat form” collected

together with a white shell of the “slender

form” at Crawl Key). No evident inter-

grades between the “squat form” and the

“slender form” are known to us, so the

squat-shelled populations are provision-

ally named as D. cf. aureocinctum.

The successful finding of a minute

radula in D. cf. aureocinctum has occurred

3 times out of the 3 checkings made in the

limited material at hand. Examined by

transparency within the alcohol-preserved

animal, the radula is verified to be situ-

ated at the tip of the proboscis, as well

when the proboscis is in extended posi-

tion (observation in the subadult speci-

men) than when the proboscis is in re-

tracted position (observation in an adult

specimen, Fig. 3). In 2 out of the 3 tenta-

tives, the radular extraction was per-

formed with success and the same minute

comb-like radula was documented (Fig.

2). These data allow to leave out the hy-

pothesis giving these radulae as remains

of digested preys. The data at hand con-

firm the radulate status of the species and

the belonging of this radula to the “Volva-

rina-Prunum comb-like pattern”.

BOYER: The discovery of a radula in a Dentimargo species and its taxonomic implications

DISCUSSION

The centrally depressed outline of

the plates and the smallest cusps

placed in median position are unusual

features compared to the comb-like

radulae known to us from the Volva-

rina-Prunum series (COOVERT AND

COOVERT, 1990), which have generally a

straight or faintly convex anterior

cusped edge, together with uniformly

distributed sub-equal small cusps, or

sub-equal small cusps with a larger

central cusp, or series of sub-equal

small cusps separated by isolated

larger cusps. However the number of

plates like the number and the shape of

the cusps in D. cf. aureocinctum are

similar to the pattern found in

numerous Volvarina-Prunum species. In

summary, the radula of D. cf. aure-

ocinctum is coherent with the range of

variability found in the Volvarina-

Prunum series, more than with the

radular patterns found in the Serrata

series (high number of subequal cusps)

or in the Mesoginella complex (trian-

gular anterior cusped edge, with a

large central cusp and few laterals).

Despite the minute size of the radula

found in D. cf. aureocinctum, there is no

reason, in the present state, to consider

it as vestigial. For instance, a minute

radula with single, narrow plates is also

found in the marginellid genus

Hydroginella known as ectoparasit

feeding at night on sleeping fishes

(BOUCHET, 1989; JOHNSON, JOHNSON

AND JAZWINSKI, 1995). As assumed by

JOHNSON ET AL. (1995) about the similar

case found in Colubraria, such minute

plates seem to work as cutting out the

fish's skin, as precondition of a feeding

process by suction of the fish's blood.

This point allows to infer that the comb-

like radula of D. cf. aureocinctum may as

well be functional despite its minute

size. Such a minute size of the radula

may also be considered, from an evolu-

tionnary point of view, as an intergrad-

ing stage towards the loss of the radula.

Due to its biconical shell outline, its

produced upper labial denticle and its

faint lower denticles, D. cf. aureocinctum

is demonstrated to be morphologically

very similar to the fossil type species D.

dentifera. For this reason its placement in

Dentimargo sensu stricto is conserva-

tively proposed as the most parsimo-

nious solution.

However, 1t must be underlined that

other Dentimargo-shelled species closely

matching with D. dentifera [like for

instance the New Zealand D. cairoma

(Brookes, 1924) studied by PONDER

(1970), the Floridian D. eburneola

(Conrad, 1834) checked by COOVERT

(1987) or the Mascarene D. pumila (Red-

field, 1870) checked by the author]

really do seem to be devoid of a radula.

The matter signifies that the occurrence

of the radula is represented as a hetero-

geneous character within the Dentimargo

series and that it cannot be used as a

diagnosis feature for the genus. In other

words, Dentimargo is provisionally con-

sidered as being composed both of

radulate and of non-radulate species.

As a direct consequence, the loss of

the radula is likely to be of low discrimi-

nating value in marginellid gastropods.

Besides, it must be emphasized that the

loss of the radula may have arisen inde-

pendantly in different lineages. This

point leads to a reconsideration not only

about the phyletic unity of Dentimargo,

but also about the degree of relationship

occurring between Dentimargo and the

supposed non-radulate genera Mar-

ginella and Glabella.

On the other hand, the Volvarina-pat-

terned radula found in D. cf. aureocinc-

tum proves that the conchological dis-

tinction between Dentimargo and Volva-

rina (not well marked in the Eocene, but

more clearly displayed in the Recent)

took place before the loss of the radula,

at least in one of the Dentimargo lin-

eages. Secondarily, the radula found in

D. cf. aureocinctum suggests a very close

relationship between Volvarina and Den-

timargo, without care about the order of

disbranching from a comb-like radulate

ancestor.

The discovery of a radula in the sup-

posed non-radulate group Dentimargo

and the inferred phyletic proximity

between Dentimargo and Volvarina lead

Iberus, 23 (2), 2005

to the assumption that the generic diag-

nosis and the supraspecific distinctions

currently accepted within the Marginel-

linae remain mainly non-operative, in

particular about the separation between

the Prunini and the Marginellini, and

about their respective composition.

The matter requires a general

reassessment of the organisation of the

Marginellinae, based on a new docu-

mentation provided by extensive com-

parisons concerning the morphologic

disparity occurring in fossil and Recent

shell material, and by correlative

researches about the radula. Additional

comparisons concerning internal

anatomy and DNA patterns may prove

to be more decisive for a clear reconsti-

BIBLIOGRAPHY

BARNARD, K. H., 1969. Contributions to the

Knowledge of South African marine Mol-

lusca, Part VI. Supplement. Annals of the South

African Museum, 47 (4): 595-661, pls. 1-2.

BOUCHET, P., 1989. A marginellid gastropod

parasitizes sleeping fishes. Bulletin of Marine

Science, 45 (1):76-84.

BOYER, F., 2001. Especes nouvelles de Margin-

ellidae du niveau bathyal de la Nouvelle-

Calédonie. Novapex, 2 (4): 157-169.

COOVERT, G. A., 1987. The external anatomy of

two species of Caribbean Marginellidae. Mar-

ginella Marginalia, 3 (5): 33-37.

COOVERT, G. A. and COOVERT, H. K., 1990. A

study of marginellid radulae. Part I: Type 6

radula, “Prunum / Volvarina” Type. Marginella

Marginalia, 8 (1-6) and 9 (1-6): 1-68.

COOVERT, G. A. and COOVERT, H. K., 1995. Re-

vision of the Supraspecific Classification of

Marginelliform Gastropods. The Nautilus,

109 (2-3): 43-110.

COssMANN, M., 1899. Essais de paléoconchologie

comparée, 3: 1-201, pls. 1-8. Cossmanmn, Paris.

tution of the disbranchings within the

subfamily.

ACKNOWLEDGEMENTS

I am greatly indebted to Dr Emilio

Rolán (Vigo University), for giving

material from his 1994 collecting trip to

Yucatan, extracting the radulae and pro-

viding the pictures.

Thanks are due also to Alain Robin

(Le Mesnil St Denis, France) for prepar-

ing the digital plate, to Andrew Wake-

field (Buckhurst Hill, UK) for correcting

the English form, and to Robert and

Nicole Hasselot (Jouques, France) for

typing out the manuscript.

JOHNSON, S., JOHNSON, J. AND JAZWINSKI, S.,

1995. Parasitism of sleeping fishes by gas-

tropod mollusks in the Colubrariidae and

Marginellidae at Kwajalein, Marshall Islands.

The Festivus, 27 (11): 121-126.

PONDER, W. F., 1970. Some Aspects of the Mor-

phology of Four Species of the Neogastropod

Family Marginellidae with a discussion of the

Evolution of the Toxoglossan Poison Gland.

Journal of the Malacological Society of Australia,

2 (1): 55-81.

STEARNS, R. E. C., 1872. Descriptions of new

species of marine mollusks from the coast of

Florida. Proceedings of the Boston Society of

Natural History, 15: 21-24.

VOKEs, H. E. AND VOKES E. H., 1983. Distribu-

tion of Shallow-Water Marine Mollusca, Yu-

catan Peninsula, Mexico. Mesoamerican Ecol-

ogy Institute, Monograph 1: 1-VIII, 1-183, pls.

1-50.

O Sociedad Española de Malacología —__—_——— lIberus, 23 (2): 53-67, 2005

About a sibling species of Mitrella minor (Scacchi, 1836)

Sobre una especie gemela de Mitrella minor (Scacchi, 1836)

Franck BOYER* and Emilio ROLÁN**

Recibido el 1-XI-2004. Aceptado el 6-X-2005

RESUMEN

Se revisa Columbella minor Scacchi, 1836 en el género Mitrella Risso, 1826, basándose

en la figura y descripción original, así como en el estudio de materialdel Mediterráneo y

Atlántico próximo. Se designa un neotipo de C. minor. Este taxon tiene prioridad sobre el

nombre más antiguo Mangelia vitrea Risso, 1826 siguiendo el artículo 23.9 de el Código

de Nomenclatura Zoológica. Se discuten la variabilidad en la morfología y decoración

de la concha y el cromatismo del animal.

Una especie gemela, que se extiende desde la Isla de Alborán al norte de Senegal, se

describe como Mitrella hernandezi spec. nov. Su diagnosis está ligada en primer lugar a

una protoconcha grande y pupoide en vez de la castaña y puntiaguda de M. minor, y en

un periostraco ténue pero con un aspecto enrejado, en vez del fuerte y toscamente arru-

gado presente en M. minor.

ABSTRACT

Columbella minor Scacchi, 1836 is revised in the genus Mitrella Risso, 1826, on the basis

of its original description and figure, and of the study of shell material from Mediterranean

and the nearby Atlantic. A neotype of C. minor is designated. C. minor takes priority on

the older name Mangelia vitrea Risso, 1826 on the basis of the article 23.9 of the Code

of Zoological Nomenclature. The variability of the shell morphology, of the shell decora-

tion and of the animal chromatism is discussed.

A sibling species ranging from Alboran Island to Northern Senegal is described as

Mitrella hernandezi sp. nov. lts principal diagnostic features lie in a large white pupoid

protoconch instead of a small brown pointed one in M. minor, and in a thin lattice-pat-

terned periostracum instead of a thick coarsely wrinckled one in M. minor.

KEY WORDS: Columbellidae, Mitrella, Columbellopsis, sibling species, Mediterranean, Lusitanian Province,

Northwest Africa.

PALABRAS CLAVE: Columbellidae, Mitrella, Columbellopsis, especies gemelas, Mediterráneo, Lusitanica

Provincia, Africa del noroeste.

INTRODUCTION

The species currently named in the species from the Lusitanian Province

literature as Mítrella minor (Scacchi, 1836) (LUQUE, 1986a; POPPE AND GOTO, 1991).

is known to be one of the most common The species is said to range at lower

and widespread of the columbellid infralittoral and at circalittoral levels

* 110, Chemin du Marais du Souci, 93270 Sevran, France.

** Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela, Spain.

Iberus, 23 (2), 2005

along the Mediterranean Sea (HIDALGO,

1917), the Atlantic side of the Iberian

Peninsula (NOBRE, 1940; ROLÁN, 1983)

and the Canary Islands (NORDSIECK AND

GARCIA-TALAVERA, 1979 as M. svelta

Kobelt, 1901; HERNÁNDEZ-OTERO AND

HERNÁNDEZ GARCÍA, 2003). Currently

placed in the genus Mitrella, Risso, 1826

[Type species by subsequent designa-

tion (Cox, 1927: 28) Mitrella flaminea

Risso, 1826 = Mitrella scripta (Linnaeus,

1758)], Columbella minor Scacchi is the

type species of the genus Columbellopsis

Bucquoy, Dautzenberg and Dollfus,

1882 (original designation).

LUQUE (1986a) noted that the popu-

lations of M. minor ranging off the

Canary Islands present a rich shell deco-

ration of small light dots on a dark

chestnut ground and of darker subsu-

tural marks, to be compared with the

uniform horny brown background

found in the populations from Mediter-

ranean and from the nearby Atlantic

(range extension said to lie from

Mogador to Vigo). LUQUE (1986a) also

recorded the occurrence of a pinkish

aperture with white labial denticles and

of about 10 striae on the base of the last

whorl in the populations from the

Canary Islands, to be compared with the

whitish aperture and the 12-14 striae

found in the northern populations. Due

to the very limited amount of shells

checked from the Canary Islands (1

from La Palma and 2 from Tenerife),

LUQUE (1986a) said to be unable to

confirm the specific identity of the

Canarian population.

The study of a large material from

the whole Lusitanian Province allows us

to state on the distinct specific identity

TAXONOMY

of the Canarian morph, and to describe

it as a new species.

MATERIAL AND METHODS

The shells studied come principally

from the MNAHN collections and from the

private collections of S. Gori, J. M. Hernán-

dez, P. Micali, C. Mifsud, A. Peñas and E.

Swinnen. Live animals have been studied

from Vigo, Algeciras and Gran Canaria.

The term of “sibling species” is used

in its trivial meaning of “very similar

species” (KNOWLTON, 1993).

Abbreviations:

AMNH American Museum of Natural

History, New York

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Muséum national d'Histoire

naturelle, Paris

MNHST Museo de la Naturaleza y el

Hombre, Santa Cruz de Tenerife

NHM The Natural History Museum,

London

ZSM Zoologische

Muenchen, Munich

CAL collection A. Locard (MNHN)

CAP collection A. Peñas

CCM collection C. Mifsud

CER collection E. Rolán

CFB collection F. Boyer

CFS collection F. Swinnen

CJH collection J. Hernández

CPM collection P. Micali

CSG collection S. Gori

s shell

sp specimen

stn station

Staatsammlung

Genus Mitrella Risso, 1826.

Type species by subsequent designation (Cox, 1927): Mitrella flaminea Risso, 1826 [= Mitrella

scripta (Linné, 1758)].

Mitrella minor (Scacchi, 1836) (Figs. 1-7, 13, 18-24, 29-31, 38, 42, 43)

Mangelia vitrea Risso, 1826. Hist. Nat. Eur. Mérid.: p. 222-223, no fig.

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figuras 1-3. Mangelia vitrea. 1, 2: holotipo, L = 8,6 mm (MNHN); 3: protoconcha del holotipo.

Buccinum politum Cantraine, 1835, non-(Lamarck, 1822) nec-(Basterot, 1825). Bull. Ac. Roy. Sci.

Bruxelles, 11: 17, no fig.

Columbella minor Scacchi, 1836. Catal. Conchy. regni Neapolitani.: p. 10, fig. 11.

Buccinum scacchi Calcara, 1840. Monog. dei Gen. Claus. e Bulim.: p. 51, no fig.

Type material: The original type material of Columbella minor Scacchi is lost, destroyed in Napoli

during the Second World War (CRETELLA, CROVATO, CROVATO, FASULO AND TOSCANO, 2005).

A specimen from “Punta Pagliarolo, Salerno, 35 m” (Fig. 21), from the collection J. Hernández, orig-

inating from the vicinity of Napoli and matching the original description of the species in all respects,

including the reticulated colour pattern of the shell, is designated as neotype of Mitrella minor. The

shell measurements are 9.47 x 3.68 mm. This neotype is deposited in MNHN.

Other material examined: A supposed syntype of Mangelia vitrea Risso, 1826 (MNHN), 8.6 x 3 mm

(Figs. 1-3), assumed to originate from the coasts of the French Riviera.

Mediterranean: 1 s, Ras il Wata, Malta, 80-100 m (CES); 3 sp, Ras il Raheb, Malta, 120 m (CFS); 15

sp, Gneja Bay, Malta,130 m (CCM) (Figs. 31, 42, 43); 1sp, 3 s, Siracusa, Sicily, 100 m (CJH) (Figs. 24,

30); 1 s, Siracusa, 35 m (CES); 2 s, Capo Asporano, Sicily (CES); 32 s, Villaggio Pace, Messina, 10-30

m (CPM) (Fig. 22); 2 sp, Punta Faro, Messina, 30 m (CPM); 2 sp, Scilla, Messina, 50-70 m (CPM)

(Fig. 20); 1 sp, Capo S. Alessio, 7-8 m (CES); 1 s, Sowona (CFS); 2 s, Boccodasse, (CES); 1 s, Infres-

chi, Marina di Camerota, 30 m, (CPM); 3 s, Ponza Island, South Thyrrenian, 600 m (CPM); 2 s, Capri

(CAL, MNHN); 3 s, Secca Murelle, Latium, 23-27 m (CPM); 6 sp, Vada, Leghorn, 80-120 m (CSG);

2 s, Ajaccio, Corsica (CAL, MNHN) (Fig. 23); 2 s, St Raphael (CAL, MNHN); 2 s, St Tropez (CAL,

MNHN); 2 s, Toulon (CAL, MNHN); 2 s, Bandol (CAL, MNHN); 2 s, Le Grau du Roi (CAL,

MNHN >; 1 s, Oran (CAL, MNHN), 45 s, Vilassar del Mar, 20-60 m (CAP; 1 s, Puerto de Solles, Mal-

lorca, 90 m (CES); 1 s, Calahonda (Málaga), beach (MNHN); 3 s, Málaga (CAP); 6 s, Rincón de la -

Victoria, Málaga, 20-40 m (MNHN); 6 s, Málaga, paseo marítimo, 20-40 m (MNHN); 2 sp, Tor-

reguadiaro, Sotogrande nets, circalittoral, (MNHN); 3 sp, Marbella, nets (MNHN); 3 s, Marbella,

30-40 m (CFB); 7 s, Marbella, beach (MNHN); 3 sp, Marbella, 70-80 m (CJH) (Figs. 4, 7, 13, 18, 19,

29,38); 3 sp, Algeciras, La Línea, 18-22 m (CFB); 2 sp, M'diq, Northern Morocco, nets (MNHN); 21

s, Ceuta, beach in the harbour (MNHN); 2 s, Ceuta South, Anse Almadrabe, 35? 52.5” N, 05* 10.0'

W, 35-45 m (MNHN); 1 sp, Ceuta, 35* 53 N, 05* 17'-05* 19 W, 50 m (MNHN); 4 s, Ceuta North,

Iberus, 23 (2), 2005

Playa Benitez, 35” 54.6' N, 05” 20.0” W, 15-25 m (MNHN); 1 sp, Ceuta Restinga (C]H); 6 s, Alborán

Island, 60-250 m (MNCN).

North East Atlantic: 1 s Tanger, market (UNHN); 4 s Tanger, beach (MNHN); 1 s, Cap Spartel,

North Morocco, 100 m (MNAHN); 5 sp, Algarve, Burgau, 37” 03.4” N, 08 46.30” W, 35 m (MNHN);

4 s, Tavira, Pedro do Barril, 25 m (MNHN); 2 sp, Algarve, Baia Belixe, 37” 00.0” N, 08* 58.0” W, 23

m (MNHN); 10 sp, 5juv, Algarve, between Salema and Praia de Luz, 70 m (MNHN); 1 sp, Algarve,

between Sagres and Faro, 40-50 m (MNHN); 2 sp Algarve, Porto de Sagres, 37” 00.6' N, 08* 55.6'

W, 9-15 m (MNHN); 1 juv, Algarve, Sagres, Punta Balecira, 17-23 m (MNHN); 2 s, Lagos, Portugal

(CES); 8 s, Ria de Vigo, 30-40 m (CER) (Figs. 5, 6); 1 s, Bouzas, Vigo, 20 m (CER); 32 sp, Cambados,

Ría de Arousa, Galicia (collection J. Horro); 2 sp, Dakhla, Western Sahara, 50-60 m (CFB).

Type locality: Gulf of Napoli.

Original descriptions: In Risso (1826),

as Mangelia vitrea: “584. M. vitrea (N.),

M. vitrée. M. Testa glaberrima, nitidissima,

vitrea; apertura lucida. Coq. tres lisse, fort

luisante, vitrée; á ouverture translucide.

Long. 0.010. Séj. Régions coralligenes.

App. printemps”.

No original figure. Supposed

syntype MNHN in Figures 1-3.

In ScacchHi (1836), as Columbella

minor: “Columbella... minor Nobis (12).

Testa parva, laevi, albo-flavescente, line-

olis fulvis obsolote reticula; anfractibus octo,

ultimo in medio pallidiore; in speciminibus

perfectis epidermide flavescente per lungum

striata obtecta; columella oblique striata,

ultra labrum parum porrecta; labro crassius-

culo interne denticulato. Alta lin: 4. Colum-

bellae flamine ac similis, sed minor, gracil-

tor, et columella productiore. In sinu

Neapolitano rara, et Inarimes insulae fos-

silis. Inspice fig. 11”.

Original type material lost. Neotype

MNHN in Figure 21.

Complementary description: Illustra-

tions in ROLÁN (1983), LUQUE (1986a),

POPPE AND GOTO (1991) and GIANNUZZI-

SAVELLI, PULSATERI, PALMERI AND EBREO

(2003).

The pointed brown protoconch

(Figs. 3, 7, 18, 19, 29-31, 38) does not

show an evident separation from the

teleoconch, but under magnification it is

suggested to have 1.75 whorls, and the

diameter of the nucleus is about 260-320

um. The “nucleus width/base width

ratio” of the protoconch is of about ?/-2.

A more or less depressed zone lies

below the beginning of the nucleus, as

clearly visible in Figure 31. In few cases,

the protoconch is rather bulbous (Figs.

23, 24) instead of pointed, and/or

5Ó

whitish (Fig. 23) instead of brown. Even

the protoconch displayed in Fig. 23,

assumed to result from a teratologic

event, holds a “nucleus width/base

width ratio” of !/2.

The very high spire shows generally

straight or very faintly concave sides

and more or less turriculated whorls.

The shape of the aperture is somewhat

variable, from rectangular to losangic,

often rather wide with a columellar

border faintly angled, columellar folds

from well-marked to absent, outer lip

moderately convex to moderately

angled, and siphonal canal rather short

and widening. The spiral striae at the

base of the last whorl range from 9 to 14,

more currently being 12-13. The ground

colour is creamy white to horny beige

with wide tan-brown spiral bands cov-

ering most of the whorls. A lighter

narrow spiral blank at the mid part of

the last whorl often separates 2 wide

brown zones. In few shells, mainly

coming from Central and Southern

Mediterranean, the brown spiral bands

tend to be fragmented in brown marks

making a stripped (Fig. 20), a reticulated

(Figs. 21, 22) or a flamed pattern (Figs.

23, 24).

The length of the adult shells ranges

from 8 mm up to 13 mm.

The periostracum is thick, with

coarse spaced axial wrinkles giving a

“waffle” appearance to the surface (Figs.

5, 6, 42, 43).

The colour pattern of the soft parts

shows as somewhat variable. The main

features are displayed by ROLÁN AND

OTERO-SCHMITT (1996) after an animal

from Vigo. Two animals examined from

Algeciras by the first author presented

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figures 4-7. Mitrella minor. 4: specimen from Marbella, 70-80 m, 13 mm (CJH); 5, 6: specimens

from the Ría de Vigo, 9.1 mm and 9.4 mm (CER); 7: protoconch of the specimen from Figure 4.

Figures 8-11. Mitrella hernandezí. 8: holotype, NW Gran Canaria, 11.0 mm (MNCN); 9:

paratype, 10.5 mm, (CJH); 10: specimen from off Banc d'Arguin, Mauritania, 80 m, 12.7 mm

(CJH); 11: protoconch of the holotype.

Figuras 4-7. Mitrella minor. 4: ejemplar de Marbella, 70-80 m, 13 mm (CJH); 5, 6: ejemplares de la

Ría de Vigo, 9,1 mm and 9,4 mm (CER); 7: protoconcha del ejemplar de la Figura 4. Figuras 8-11.

Mitrella hernandezi. 8: holotipo, NW Gran Canaria, 11,0 mm (MNCN); 9: paratipo, 10.5 mm,

(CJH); 10: ejemplar del Banc d'Arguin, Mauritania, 80 m, 12,7 mm (CJH); 11: protoconcha del

holotipo.

Iberus, 23 (2), 2005

the same general pattern: underside and

periphery of the sole whitish, column of

the foot shaded on its sides by large

blackish zones on a whitish grey

ground, front zone whitish grey, back

zone blackish; head, top of the neck and

tentacles black, except around the eyes,

the underside and the tip of the tenta-

cles which are whitish; siphon blackish,

all the whitish or whitish grey zones

being flecked by deep white tiny dots.

The operculum is hyalinous yellow.

The radula is summarized in LUQUE

(1986a, pl. 3, fig. f).

Distribution: The whole Mediter-

ranean and the nearby Atlantic from the

Straight of Gibraltar to Galicia. Live

specimens from 7 m to 130 m. The

occurrence along the Atlantic coasts of

the Northern Morocco is probable, but it

remains to be fully documented. The

occurrence off Western Sahara remains

dubious, as only one record is known

from this area (2 sp, Dakhla, CFB), pos-

sibly coming from accidental mixing of

shells from Mediterranean and from

Western Sahara. Not represented in the

Canary Islands.

Remarks: VAN AARTSEN, MENKHORST

AND GITTENBERGER (1984: 80) noted that

Columbellopsis has been considered later

by its authors as falling in synonymy

with Atilia H. and A. Adams, 1853, and

they claimed that the “poorly known

boundaries between species in this

group” did not allow “to place M. minor

in a subgenus of its own”.

VAN AARTSEN ET AL. (1984) did not

care with the arguments displayed by

RADWIN (1978: 331-332), who shows that

the designation of Mitrella minor Scacchi

as type species of Atilia was invalid, that

the species has been removed from

Atilia by the next reviewer and that

Columbellopsis can be used as brother

genus of Mitrella in the present state,

being represented by its own peculiar

shell features besides Mitrella as far back

as the early Eocene. These consistent

arguments will be to take in considera-

tion in a taxonomic revision of the eclec-

tic genus Mitrella, but the matter

requires a general analysis of the phy-

logeny prevailing about the different

species group currently lumpered

within Mitrella s.1. In the wait of such a

revision, we feel more appropriate to

keep conservatively Columbella minor in

the genus Mitrella.

Despite its status of ubiquitous

species, few synonyms occurred and the

specific name of C. minor prevailed in

the literature since the Scacchi's descrip-

tion. Buccinum politum Cantraine, 1835,

which is a preoccupied name, was

proved to be a nomen dubium by VAN

AARTSEN ET AL. (1984), Buccinum minus

Philippi, 1836 was a misspelling of the

Scacchi's name, and Buccinum scacchi

Calcara, 1840 was apparently the last

name proposed as referring to the

species. CALCARA (1845: 40) considered

apparently its Buccinum scacchi Calcara,

1840 as synonym of Columbella minor

Scacchi, 1836.

The elusive name of Mangelia vitrea

Risso, 1826, neglected in the literature

due both to its deceitful original generic

attribution and to its poorly descriptive

original definition, is considered herein

as an objective older synonym of C.

minor, on the ground of the supposed

syntype stored in MNHN. We must

express however some reserves about

the status of this syntype and about the

identity of the taxon Mangelia vitrea.

These reserves lie first in the fact that

ARNAUD (1977) did not recognize any

material referring to M. vitrea in MNHN

at the time of his study (type said to be

lost), and that the subsequent determi-

nation of the so-said syntype of M. vitrea

was made by the MNHN curator on the

basis of a label from the hand of A.

Risso (“Mangelia vitrea Risso”) joined to

the supposed syntype but without

further correlation between this label

and the shell specimen. Secondly, the

figure of M. vitrea pictured by P. Gény in

ARNAUD (1977: pl. 11, fig. 201) and sup-

posed to represent a shell labelled under

this name in the Risso's collection, does

not match really the shell morphology

of the supposed MNHN syntype. The

figure given by P. Gény presents a shell

with a narrower, more slender and more

acute spire, and a longer siphonal canal

with a well-marked notch, the general

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figures 12, 14-17. Mitrella hernandezí, paratype (CER). 12: detail of the aperture; 14: operculum;

15: radula; 16, 17: protoconch of the shell from Fig. 12. Figures 13, 18, 19. Mitrella minor, speci-

men from Marbella, 70-80 m, (CJH). 13: detail of the aperture; 18, 19: protoconch of the shell

from Figure 13.

Figuras 12, 14-17. Mitrella hernandezi, paratipo (CER). 12: detalle de la abertura; 14: opérculo; 15:

rádula; 16, 17: protoconcha de la concha de la Fig. 12. Figuras 13, 18, 19. Mitrella minor, ejemplar de

Marbella, 70-80 m (CJH). 13: detalle de la abertura; 18, 19: protoconcha de la concha de la Figura 13.

Iberus, 23 (Q), 2005

outline suggesting a turrid species. On

the other hand, the poor original

description of M. vitrea matches more

closely the supposed MNHN syntype

than any turrid species from Mediter-

ranean known to us. Any of these argu-

ments does not allow by itself to define

M. vitrea as a dubious species.

The junior name Columbella minor

Scacchi, 1836 is considered to have prior-

ity on the older name Mangelia vitrea

Risso, 1826 on the ground of the article

23.9 of the Code of Zoological Nomen-

clature (Precedence Inversion): M. vitrea

(probably considered as a nomen

dubium by the subsequent authors) was

apparently not used in the literature

after 1899, and C. minor was used in

more than 25 works (generally under the

genus Mitrella), published by more than

10 authors during the last 50 years along

a period of more than 10 years. Refer-

ences and the pages where the species

was mentioned are the following:

PASTEUR-HUMBERT (1962: 159), NORD-

SIECK (1968: 124), PARENZAN (1970: 171),

SCHIRO (1979: 7), SABELLI AND SPADA

(1981: 1), TERRENT (1981-35), RROLÁN

(1983: 248), VAN AARTSEN ET AL. (1984:

37), BRUSCHI, CEPPODOMO, GALLI AND

PIANI (1985: 25), LUQUE (1986a: 234),

LUQUE (1986b: 91), CASTAÑO, CIVIS AND

GONZÁLEZ DELGADO (1988: 178),

SABELLI, (GIANNUZZI-SAVELLI AND

BEDULLI (1990: 206), POPPE AND GOTO

(1991: 152), ARDUINO, LOCATELLI,

ORLANDO AND REPETTO (1995: 82),

ROLÁN AND OTERO SCHMITT (1996: 100),

GIRIBET AND PEÑAS (1997: 52), MACEDO,

MACEDO AND BORGES (1999: 204),

ARDOVINI AND COSSIGNANI (1999: 62),

BOUCHET, LE RENARD AND GOFAS (2001:

194), GIANNUZZI-SAVELLI ET AL. (2003:

256), HERNÁNDEZ-OTERO AND HERNÁN-

DEZ GARCÍA (2003: 84), OZTURK,

BUZZURO AND AVNI BENLI (2004: 58),

ARDOVINI AND COSSIGNANI (2004: 166),

POGGIANI, MATTIOLI AND MICALI (2004:

116), and CRETELLA ET AL. (2005: 120).

This solution is proposed for the

benefit of the stability of the nomencla-

ture.

Mitrella minor does not show evident

geographic forms. However, the speci-

mens from Vigo (Figs. 5, 6) have smaller

and darker shells than those from

Mediterranean, and they seem to have a

more prominent periostracum (compare

with a specimen from Malta in Figures 42

and 43). The populations from the coasts

of Portugal and Spain, also those from

Southern France and from the Italian Lig-

uria, mainly show an uniform tan-brown

shell colour pattern, sometimes dull

creamy-White, whereas the populations

from Central and Southern Mediter-

ranean (Corsica, Western Italy from

Toscana to Sicily, North Africa coasts)

present most frequently a “flamed”,

“reticulated” or “slack check-patterned”

shell decoration (from the words of SCAC-

CHI, 1836), corresponding to the original

description of M. minor. A white vitreous

to hyalinous form with browny to grey-

ish protoconchs, which matches the origi-

nal description of Mangelia vitrea (original

locality belonging probably to the French

Riviera), seems also to be more frequent

in Central Mediterranean [specimens

observed from Leghorn (CSG) and from

Siracusa (CJH)]. Full tan-brown shells

and intergrades are also found off West-

ern Italy, and full tan-brown shells are

found off Malta.

The noticeable variations in the pro-

toconch morphology and background

colour pictured herein (Figs. 3, 7, 18, 19,

29-31, 38) do not correlate with the vari-

ations observed in the morphology and

in the decoration of the shells (Figs. 1, 2,

4-6, 13, 20-24, 42). One shell from

Ajaccio, Corsica (Fig. 23) is comparable

to the following species for its whitish

bulbous protoconch, but it is provision-

ally named as M. cf. minor, due to the

fact that this protoconch seems to result

from a teratologic event rather than to

express a distinctive specific feature.

The wide aperture and the colour

pattern of broad flames occurring in the

shell pictured in Figure 23 are coherent

with the variability observed in M.

minor, but not with the variability

observed in the following species.

Further inquiries will verify if the distri-

bution of the following species reaches

Corsica and if the shell pictured in

Figure 23 really belongs to M. minor.

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figures 20-22, 24. Mitrella minor. 20: specimen from Scilla, Messina, 50-70 m, 10.8 mm (CPM);

21: neotype, specimen from Punta Pagliarolo, Salerno, 35 m, 9.47 mm (MNHN, ex-CJH); 22: shell

from Villaggio Pace, Messina, 10-30 m, 9.5 mm (CPM); 24: specimen from Siracusa, Sicilia, 100 m,

8.5 mm (CJH). Figure 23. Mitrella cf. minor, shell from off Ajaccio, Corsica, 11 mm (MNHN).

Figures 25-28. Mitrella hernandezi. 25: specimen from Alboran Island, 20 m, 9.8 mm (CJH); 26:

specimen from Alboran Island, 20 m, 10.20 mm (CJH); 27: specimen from Alboran Island, 20 m,

9.25 mm (CJH); 28: specimen from La Manchita, NW Gran Canaria, 35 m, 9.5 mm (CJH).

Figuras 20-22, 24. Mitrella minor. 20: ejemplar de Scilla, Messina, 50-70 m, 10,8 mm (CPM); 21:

neotipo, ejemplar de Punta Paglialoro, Salerno, 35 m, 9,47 mm (MNHN, ex-CJH); 22: concha de Vil-

laggio Pace, Messina, 10-30 m, 9,5 mm (CPM); 24: ejemplar de Siracusa, Sicilia, 100 m, 8,5 mm

(CJHA). Figura 23. Mitrella cf. minor, concha de Ajaccio, Corsica, 11 mm (MNAN). Figuras 25-28.

Mitrella hernandezi. 25: ejemplar de la isla de Alborán, 20 m, 9,8 mm (CJH); 26: ejemplar de la isla

de Alborán, 20 m, 10,2 mm (CJH); 27: ejemplar de la isla de Alborán, 20 m, 9,25 mm (CJH); 28:

ejemplar de La Manchita, NO Gran Canaria, 35 m, 9,5 mm (CJH).

Iberus, 23 (2), 2005

Mitrella hernandezi sp. nov. (Figs. 8-12, 14-17, 25-28, 32-37, 39-41)

Type material: Holotype (Figs. 8, 11) in MNCN (15.05 /46626); paratypes in AMNH (1), NHM (1),

MNHN (1), ZSM (1), MNHST (1), CJH (0) (Fig.9), CER (2) (Figs 12, 14-17), CFB (2), all from the

type locality.

Other material examined: Mediterranean: 5 sp, Alborán Island, 20 m (CJH) (Figs. 25-27, 32-34).

Canary Islands: 2 s, Gran Canaria (CJH); 43 s, off NW Gran Canaria, 170 m, (CJH); 24 sp, Barranco

de Guayedra, NW Gran Canaria (CJH); 2 s, NW Gran Canaria, 150 m (CJH) (Figs. 35, 36); 2 s, Gran

Canaria (CJH); 4 sp, NW Gran Canaria, 302 m (CJH); 8 s, La Manchita, Gran Canaria, 35 m (CJH)

(Fig. 28); 5 sp, NW Gran Canaria, 150-200 m (CFB); 5 s, off Las Nieves, NW Gran Canaria, 34 m

(CFB); 1 s off San Cristobal, NE Gran Canaria, 40-60 m (CJH).

Northwest Africa: 3 s, south of Cape Bojador, 25” 40” N, 15” 03'4 W, 100 m (CSG); 1 s, Western

Sahara (CES); 1 sp, Western Sahara, 30 m (CJH); 6 sp, Western Sahara, 30-40 m (CJH); 10 sp, Western

Sahara, 60 m (CJH) (Figs. 37, 39-41); 2 sp, Western Sahara, 40 m (CJH); 15 sp, 2 s, Western Sahara,

30 m (C]H); 6 s, Western Sahara, 60 m (CJH); 1 s, 22* 00' N, 17? 22” W (CJH); 2 sp, Dakhla, Western

Sahara, 50-60 m (CFB); 5 sp, Western Sahara, 30-40 m (CFB); 3 s, Western Sahara, 50-60 m (CFB); 1

s, Meteor stn 36, 21” 19.5” N, 17? 13.1” W, 58 m (ZSM); 1 sp, 9 s, off Banc d'Arguin, Mauritania, 80

m (CJH) (Fig. 10); 1 s, Lompoul, Northern Senegal, 150 m (CFB); 1 s, Meteor stn 60.77, 17? 17' N,

16% 30” W, 85 m (MNHN); 1 s, NO “N Diago” 1981 stn 119, 18? 36' N, 16” 28” W, 70 m (MNHN); 1

s, NO “N' 'Diago” 1981 stn 103, 18” 48" N, 16” 22” W, 28 m (MNHN); 1 s, NO “N Diago” 1981 stn

248, 17” 54 N, 16? 20” W, 76 m (MNHN); 1 s, NO “N'Diago” 1981 stn 232, 17? 42 N, 16* 05 W, 12

m (MNHN); 1 s, Dakar harbour, 11-12 m (MNHN); 1 s, NO “N Diago” 1981, stn 240, 17” 48' N, 16?

24” W, 38 m (MNHN); 1 s, region of Dakar, 14? 19 N, 17? 32” W, 132 m (MNHN); 1 sp, 14 s, region

of Dakar, 14” 24” N, 17” 23” W, 78 m (UNHN); 1 s, Dakar, 14" 51 N, 17* 30 W, 165-180 m (MNHN);

1 s, Dakar, 14” 52” N, 17? 30” W, 140-150 m (MNHN); 2 s, NO “N' Diago” 1981 stn 63, 20? 42' N, 17?

21” W, 43 m (MNHN); 1 s, 1 fragment, region of Dakar, 20” 58” N, 17? 37” W, 110 m (MNHN); 8 s,

South of Gorée, 48-50 m (MNHN); 1 s, South of Gorée, 95-98 m (MNHN).

Type locality: Off Northwest Gran Canaria, 50-170 m.

Etymology: The specific name is dedicated to José María Hernández, Canarian malacologist devot-

ing to the study of the molluscan fauna from the Southern Lusitanian Province.

Description: The shell (Figs. 8-10) is

solid, smooth, lanceolate with an elon-

longed internally as lirae. The outer lip

has a wide external thickening.

gate spire. The whitish protoconch (Fig.

11) is large, pupoid and smooth. No

clear separation with the teleoconch

occurs, but under magnification it

appears to be made of one whorl reach-

ing about 750 um of diameter at the base

and of a wide bulbous nucleus of about

500 um. The spire has about 5 smooth

flat whorls, an evident incised suture

without subsutural step and its sides

show a slightly concave outline. There

are about eleven spiral striae occur at

the base of the body whorl. The aperture

is small, subtriangular, comma-shaped,

and it is continued with a sinuous

siphonal canal relatively long and

narrow for the genus. The columella

bears 5 scarcely evident teeth. The col-

umellar callus is slightly extended exter-

nally, convex and moderately promi-

nent. Seven well marked labial teeth

occur inside the aperture and are pro-

The colour pattern is light honey

brown with numerous small ovoid

white dots all over the shell; a discontin-

uous suprasutural line is formed by

dark brown spots alternating with

lighter ones, a similar subsutural line is

formed by longer, larger and more

spaced out dark brown dashes.

The lattice-patterned periostracum

(Figs. 40, 41) is very thin.

Dimensions: The holotype (Fig. 8) is

11.0 x 3.8 mm. Some shells reach up to

13 mm length.

Headfoot: Some specimens have been

observed alive from Northwest Gran

Canaria, dredged at 50 m. The headfoot

shows a creamy-white background with

brown-violet dashes and stains, the

siphon showing also small yellow spots.

The distal parts are white, like are the

lateral sides of the foot and a line bor-

dering the propodium and the opercu-

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figures 29-31, 38. Mitrella minor, protoconchs. 29: specimen from Marbella, 70-80 m (CJH); 30:

specimen from Siracusa, 100 m, (CJH); 31: specimen from Gneja Bay, Malta (CCM); 38: same as

Figure 29. Figuras 32-37, 39. Mitrella hernandezí, protoconchs. 32: specimen from Alboran Island,

20 m (CJH); 33: specimen from Alboran Island, 20 m, (CJH); 34: specimen from Alboran Island,

20 m (CJH); 35: specimen from NW Gran Canaria, 150 m (CJH); 36: specimen from NW Gran

Canaria, 150 m (CJH); 37: specimen from Western Sahara, 60 m (CJH); 39: same as Figure 37.

Figuras 29-31, 38. Mitrella minor, protoconchas. 29: ejemplar de Marbella, 70-80 m (CJA); 30: ejem-

plar de Siracusa, 100 m, (CJH); 31: ejemplar de Gneja Bay, Malta (CCM); 38: el mismo que la Figura

29. Figuras 32-37, 39. Mitrella hernandezi, protoconchas. 32: ejemplar de la isla de Alborán, 20 m

(CJH); 33: ejemplar de la isla de Alborán, 20 m, (CJH); 34: ejemplar de la isla de Alborán, 20 m

(CJ); 35: ejemplar del NO de Gran Canaria, 150 m (C]H); 36: ejemplar del NO de Gran Canaria,

150 m (CJH); 37: ejemplar de Sahara Occidental, 60 m (CJH); 39: el mismo que la Figura 37.

Ó3

Iberus, 23 (2), 2005

lum area. A live animal is pictured in

HERNÁNDEZ AND BOYER (2005).

- The operculum (Fig. 14) is ovoid

with a terminal nucleus and an insertion

mark divided in two parts by a strong

wrinkle.

Radula: Typical of the genus, with a

rectangular rachidian tooth and two

stretched marginal teeth bearing one

bulbous cusp near to the base and two

hooked cusps along the distal side (Fig. 15).

Distribution: From Alborán Island to

Northern Senegal, live specimens from

20 to 300 m.

Remarks: The protoconch shows

about 1.75 whorl, and the “nucleus

width/base width ratio” is of about ?/3

(Figs. 16, 17, 32, 33, 35-37, 39), except in

the case of one specimen from Alborán

Island (Fig. 34) which has a greyish

pointed protoconch with a ratio closer to

1/2. However, all the other shell features

of this specimen reach the common fea-

tures of Mitrella hernandezi sp. nov. and

its protoconch is assumed to result from

a teratologic event (low stepped apex

differing noticeably from the pointed

apex of M. minor). A depressed zone un-

der the nucleus is sometimes evident in

the protoconch of M. hernandezi (Figs. 32,

33), but it is often faint or lacking (Figs.

34-37). The striae at the base of the last

whorl count 9 to 16, being more often 10-

11. The columellar teeth count 5 to 7. For

the other morphologic features of its

shell, like for its colour pattern, M. her-

nandezi is very constant all along its wide

range of distribution.

Some populations of M. hernandezi

from Western Sahara show a very light

and attenuated shell colour decoration.

In any case they show the fine suprasu-

tural and subsutural interrupted lines

mentioned in the description. Few shells

from Gran Canaria may show a pupoid

light brown protoconch instead of

whitish as usual (CFB).

NORDSIECK AND GARCIA-TALAVERA

(1979) mentioned and illustrated our

new species M. hernandezi as “Mitrella

suelta (Monterosato) Kobelt, 1901” from

the Canary Islands (La Palma) and from

North Africa. In CHIARELLI, MICALI AND

QUADRI (2003), M. svelta Kobelt, 1889 (ex

Monterosato ms) is considered to be an

error pro M. spelta (Kobelt, 1893), which

is placed in synonymy with Mitrella

lanceolata (Locard, 1886), a species very

similar to the common Mediterranean

Mitrella scripta (Linnaeus, 1758).

M. hernandezi must be considered as

a sibling species of M. minor (“pseudo-

sibling species” according to KNOWLTON,

1993), being distinguishable principally

on the basis of its strong pupoid (gener-

ally white) protoconch with a “nuclear

width /base width ratio” of about ?/3 in-

stead of a smaller pointed (generally

brown) protoconch in M. minor with a

ratio of about */2, and on the basis of its

very thin lattice-patterned periostracum

instead of a thick coarsely wrinckled one

in M. minor. The shell aperture in M. her-

nandezi is more triangular, often smaller

and narrower than in M. minor, the

siphonal canal is generally longer, the

outer lip more angular and the labial

denticles stronger. The incised suture

and the concave outline of the spire sides

seem also to be specific features of M.

hernandezi, despite the occurrence of

some intergrading cases in M. minor. The

usual shell decoration of M. hernandezi is

characterized by a pattern of small

packed white dots, which has not a real

equivalent in M. minor (Figs. 21, 22), who

presents a much more variable shell dec-

oration.

Albeit the average number of spiral

striae at the base of the body whorl is

10-11 in M. hernandezi and 12-13 in M.

minor, higher numbers of striae may

occur in M. hernandezi and lower

numbers may occur in M. minor. The

colour design of the animal is similar in

both species.

Despite the lack of data about the

forms ranging off the western coasts of

Morocco between Cape Spartel and

Cape Juby, the record of specimens of

M. hernandezi collected by J. M. Hernán-

dez himself off Alborán Island (CJH, 20

m) and the record from the same place

of shells of M. minor (MNCN, 60-250 m)

allow to state about the sympatry of

both species at least off Alborán Island,

and probably all along the western

coasts of Morocco.

BOYER AND ROLÁN: About a sibling species of Mitrella minor (Scacchi, 1836)

Figures 40, 41. Mitrella hernandezi, periostracum. 40: specimen from Western Sahara, 60 m, 10

mm (CJH); 41: same as Figure 40, detail. Figures 42, 43. Mitrella minor, periostracum. 42: speci-

men from Gneja Bay, Malta, 130 m, 9.7 mm (CCM); 43: same as Figure 42, detail.

Figuras 40, 41. Mitrella hernandezi, periostraco. 40: ejemplar de Sahara Occidental, 60 m, 10 mm

(CJH); 41: el mismo que la Figura 40, detalle. Figures 42, 43. Mitrella minor, periostraco. 42: ejem-

plar de Gneja Bay, Malta, 130 m, 9,7 mm (CCM); 43: el mismo que la Figura 42, detalle.

Iberus, 23 (2), 2005

ACKNOWLEDGEMENTS

-_ We are indebted to José María

Hernández (Gáldar, Gran Canaria,

Spain) for the loan of material, for the

gift of the type specimens and for field

observations. The material from Gran

Canaria and from Western Sahara was

collected mostly by Arcadio Benitez

Galván (Agaete, Gran Canaria, Spain).

Important lots of material from

Mediterranean were loaned by Sandro

Gori (Livorno, Italy), Pasquale Micali

(Fano, Italy), Constantine Mifsud (Ra-

bat, Malta), Anselmo Peñas (Vilanova 1

la Geltrú, Spain), Juan Horro (Vigo,

Spain) and Frank Swinnen (Lommel,

Belgium).

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O Sociedad Española de Malacología —__—_——— Iberus, 23 (2): 69-93, 2005

Notes on the columbellid fauna from the infralittoral and

circalittoral levels of the Canary Islands

Notas sobre los columbelidos del infralitoral y circalitoral de Canarias

José María HERNÁNDEZ* and Franck BOYER**

Recibido el 1-XI-2004. Aceptado el 6-X-2005

ABSTRACT

The columbellid species found in the infralittoral and the circalittoral off the Canary Islands

and attributed to the genera Columbella, Mitrella, Anachis, Parvanachis, Zafra and Nassa-

rina are discussed. Taxonomy, phenetic variability and range of distribution of the species

are commented.

Mitrella turbita (Duclos, 1840) is confirmed to be found off Gran Canaria, and it is

recorded for the first time from Fuerteventura. Nitidella ocellina Nordsieck, 1975 and

Pusionella scripta Nordsieck, 1975 are considered as junior synonyms of Mitrella broderipi

(Sowerby, 1844). A slender “deeper form” of Mitrella broderipi is recorded from the

Canary Islands and it is showed to belong to the morphologic variability of the species. The

overall morphologic similarity with Mitrella broderipi and the presence of the same array of

chromatic variation leads to make the hypothesis that Anachis avaroides Nordsieck, 1975

might be a ribbed variation of M. broderipi. The occurrence of Mitrella bruggeni van Aart-

sen, Menkhorst and Gittenberger, 1984 is confirmed in the Canary Islands, with a stout

shelled “shallow form”, similar to the populations found in Mediterranean, and a slender

“deeper form” restricted to the Canary Islands. Buccinum canariense d'Orbigny, 1839 is

stated to be a junior synonym of Mitrella ocellata (Gmelin, 1791).

The Caribbean species Parvanachis obesa (C. B. Adams, 1845) is recorded from the har-

bour of Santa Cruz de Tenerife. This occurrence is interpreted as resulting from an acciden-

tal human introduction, but the maintaining of the discovered population remains to be con-

firmed. Conversely, the Indo Pacific species Zafra exilis (Philippi, 1849) is confirmed to oc-

cur all around Gran Canaria and possibly to be settling in Tenerife, as a case of successful

introduction by the naval traffic. The documentation at hand leads to consider that failed in-

troductions may be frequent, as resulting directly from the contemporary maritime economy.

Nassarina rietae Segers and Swinnen, 2004 is considered as a possible endemic from the

Canary Islands.

The presence of axial ribs is showed to be very variable within a species like M. turbita or

within a species-group like the M. broderipi / M. avaroides one. As a result, the separa-

tion between the Mitrella group and the Anachis group on the basis of the lack or of the

presence of axial ribs is appreciated as being artificial. This point is proposed as an argu-

ment for a reviewing of the supraspecific classification of the Columbellidae.

RESUMEN

Se discuten las especies de la familia Columbellidae presentes en el infralitoral y el circali-

toral de Canarias, atribuidas a los géneros Columbella, Mitrella, Anachis, Parvanachis,

* Capitán Quesada 41, 35460 Gáldar, Gran Canaria, Spain.

** 110 Chemin du Marais du Souci, 93270 Sevran, France.

Iberus, 23 (2), 2005

Zafra y Nassarina, poniendo especial énfasis en su taxonomía, variabilidad fenética y

distribución de las especies.

Se confirma la presencia de Mitrella turbita (Duclos, 1840) en Gran Canaria, y se cita por

primera vez en Fuerteventura. Se considera Nitidella ocellina Nordsieck, 1975 and Pusio-

nella scripta Nordsieck, 1975 como sinónimos posterior de Mitrella broderipi (Sowerby,

1844). Una forma alargada, de profundidad, de Mitrella broderipi se cita de Gran Cana-

ria y se demuestra su pertenencia a la variabilidad morfológica de la especie. Asimismos,

la semejanza de sus características morfológicas y la presencia en ambas del mismo rango

de variaciones cromáticas nos hace pensar que Anachis avaroides Nordsieck, 1975 podría

ser una variedad con costillas de Mitrella broderipi. Se confirma la presencia de Mitrella

bruggeni van Aartsen, Menkhorst and Gittenberger, 1984 en Canarias, con dos formas,

una forma rechoncha, de aguas someras, similar a las poblaciones del Mediterráneo, y una

forma esbelta, de aguas profundas, restringida a Canarias. Se considera Buccinum cana-

riense d'Orbigny, 1839 como sinónimo posterior de Mitrella ocellata (Gmelin, 1791).

La especie del Caribe Parvanachis obesa (C.B.Adams, 1845) se cita en aguas someras de

Santa Cruz de Tenerife. Esta presencia es interpretada como resultado de una introducción

- humana accidental, pero el mantenimiento de la población descubierta, queda pendiente

de confirmar. Se confirma la presencia de la especie indo-pacífica Zafra exilis (Philippi,

1849) en toda Gran Canaria y su posible establecimiento en Tenerife, como un caso de

introducción exitosa, debido a la actividad industrial. La documentación disponible, nos

conduce a pensar que introducciones fallidas de especies son probablemente frecuentes,

como resultado directo de la economía marítima contemporánea. Se considera Nassarina

rietae Segers and Swinnen, 2004 como una posible especie endemica de las Canarias.

Se muestra que la presencia de costillas axiales es muy variable dentro de especies como

M. turbita o en el complejo M. broderipi / M. avaroides. Como resultado, la separación

entre el grupo Mitrella y Anachis en base a la ausencia o presencia de costillas axiales,

parece ser artificial. Esto se propone como argumento para un replanteamiento de la cla-

sificación supraespecífica de los Columbellidae.

KEY WORDS: Columbellidae, taxonomy, phenetic variability, distribution, deep forms, sibling species, intro-

duced species, Canary Islands.

PALABRAS CLAVE: Columbellidae, taxonomía, variabilidad fenética, distribución, especies gemelas, especies

introducidas, Islas Canarias.

INTRODUCTION

Whereas the Columbellidae from the

Mediterranean have been the subject of

recent works of revision (VAN AARTSEN,

MENKHORST AND GITTENBERGER, 1984;

LuQuE, 1986; CHIARELLI, MICALI AND

QUADRI, 2003), so much attention has

not be given to the species from North-

east Atlantic.

The illustrated catalogue of NORDSIECK

AND GARCÍA-TALAVERA (1979) on the

species from the Canary Islands is the only

attempt to present a general view of a local

columbellid fauna within this area,

through the picturing and the comment

of 14 morphospecies. The recent list of

Columbellidae published by HERNÁNDEZ

OTERO, GARCÍA-TALAVERA AND HERNÁN-

DEZ GARCÍA (2003) in the frame of the Biota

project (Inventory of the Canarian marine

fauna) gives 11 taxa, of whose only 6 are

quoted by NORDSIECK AND GARCÍA-TALAV-

ERA (1979). This simple fact shows how

much controversial remains the taxonomy

of the local columbellids.

The limited scope of this article is to

summarize the present knowledge on

the columbellid fauna from the infralit-

toral and upper circalittoral levels of the

Canary Islands (about 0-100 m), with a

special point on the taxonomy of species

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary ls.

and genera, on their phenetic variability

and on their range of distribution. The

genera Amphissa H. and A. Adams, 1853

and Astyris H. and A. Adams, 1853 are

both recorded from the Canary Islands

by NORDSIECK AND TALAVERA (1979)

through 3 specific taxa which are con-

sidered by RADWIN (1978 b) as corre-

sponding to 2 controversial amphiat-

lantic species from deep waters of

Northern Atlantic. Due to their status of

bathyal species, the study of these

Canarian items are out of the scope of

the present article.

Despite the opinion of RADWIN (1978

a) about the limited conception of the

genus Anachis H. and A. Adams, 1853,

implied by Tate“s selection of the type

species Columbella scalarina, the use of

this taxon is preferred here to the use of

the more recent genus Costoanachis

Sacco, 1890 which does not resolve the

issue Of the wide morphologic disparity

at work in the complex of axially ribbed

spindle-shaped columbellids. The other

genera are used following the RADWIN'S

position (1977, 1978 a, 1978 b).

This study is based principally on the

observations and on the collection of both

authors, on the M. Bermejo collection

SYSTEMATIC PART

deposited in Museo Canario (Las Palmas,

Gran Canaria), and on the private collec-

tions of W. Engl and of E. Swinnen.

The term of “sibling species” is used

in the trivial sense of “species sharing

very similar features” (KNOWLTON, 1993).

Abbreviations:

sh: shells.

many sh: > 20 sh.

CVI: Cape Verde Islands.

Fu: Fuerteventura

GC: Gran Canaria;

Go: La Gomera

Hi: El Hierro

La: Lanzarote

Pa: La Palma

Ma: Madeira

Te: Tenerife

WS: Western Sahara

MNHN: Muséum National d'Histoire

Naturelle, Paris.

SMF: Senckenberg Museum, Frankfurt.

FBC: F. Boyer Collection.

FSC: FE. Swinnen Collection.

JHC: J. Hernandez Collection.

MBC: M. Bermejo Collection, Museo Ca-

nario, Las Palmas.

WEC: W. Engl Collection.

Family COLUMBELLIDAE Swainson, 1840

Genus Columbella Lamarck, 1799

Type species by monotypy: Voluta mercatoria Linnaeus, 1758.

Columbella adansoni Menke, 1853 (Fig. 1)

Material examined: Ma: 1 sh, 9-12 m, JHC. Hi: 1 sh, 5-12 m, JHC. GC: many sh, 0-90 m, JHC (Fig.1);

many sh, 0-3 m, FBC. La: many sh, 0-3 m, FBC. Fu: 12 sh, 0-60 m, JHC.

Taxonomy: Attributed for a long time

to the non-planktotrophic species

Columbella rustica (Linnaeus, 1758) and

more specially to the morph C-. striata

Duclos, 1835 (for instance in NORDSIECK

AND GARCÍA-TALAVERA, 1979), the

Columbella species distributed in the

Canary Islands has been recently

demonstrated to belong to the plank-

totrophic sibling species C. adansoni

Menke, 1853, described from the Cape

Verde Islands (MOOLENBEEK AND

HOENSELAAR, 1991). C. adansoni was pic-

tured in NORDSIECK AND GARCÍA-TALAV-

ERA (1979) as “C. rustica striata Duclos,

1835” and also as “Columbella spec.” for a

tall-spired form (subadult shell).

The belonging of C. adansoni to the

genus Columbella is not suspicious, as its

shell morphology is very close to that of

Iberus, 23 (2), 2005

the type species C. mercatoria (Linnaeus,

1758).

Distribution: C. adansoni was said to be

restricted to the Macaronesian Islands by

MOOLENBEEK AND HOENSELAAR (1991),

whereas C. rustica was said to be restricted

from Mediterranean to Senegal. OLIVERIO

(1995) enlarged the distribution of C. adan-

soni to the remainder of the West African

Province, this distribution being con-

firmed from Sierra Leone to Central

Angola by ROLÁN AND RYALL (1999).

The distribution of C. adansoni in

Canary Islands is general, from low tide

level to 90 m. The species is especially

abundant under boulders in shallow water.

Remarks: he complex C. rustica / C.

adansoni has been cited (for instance in

THORSON, 1949) as a classic example of

poecilogony (intraspecific variation in

the mode of larval development) in mol-

luscan gastropods.

MOOLENBEEK AND HOENSELAAR

(1991) stated the presence, in the Mac-

aronesian C. adansoni, of a multispiral

protoconch indicating a planktotrophic

development and, in the Mediterranean

and North West African C. rustica, of a

paucispiral protoconch indicating a

“direct development” (more exactly it is

intracapsular metamorphosis).

The electrophoresis analysis per-

formed by OLIVERIO (1995) confirmed the

separation of both species at the genetic

level, the initial divergence being esti-

mated from about 2 millions years. OLIv-

ERIO (1995) emphasizes that “this time can

be correlated to the onset of glaciations,

and especially with their extension to

southern regions”. This could explain the

present distribution of C. rustica, which

may have reached its full intracapsular

development during a glacial isolation

stage within Mediterranean, before to

extend to the North West African coasts,

while C. adansoni remained protected from

the cold Canary current in the offshore

Macaronesia Islands.

It must be noted that the sibling

species C. rustica and C. adansont are

presently separated only on the basis of

their respective protoconch and of their

genetic distance, but they remain to be

fully studied in other ways, specially con-

cerning the variability of the shell mor-

phology, the external features and the

anatomy of the soft parts, the ontologic

development at the juvenile stage and the

general behaviour at the adult stage.

A superficial examination of the ani-

mals of C. adansoni in Canary Islands

(milky white to creamy white ground,

with zones flecked of deep white dots,

large golden brown to amber patches,

small rounded yellowish operculum

with black axis, scalloped by a deep yel-

low-orange line in its anterior part and

by a black line in its posterior part) did

not allow to recognize any significant

difference with regard to the animals of

C. rustica examined by the authors from

Mediterranean and from Senegal.

Genus Mitrella Risso, 1826

Type species by subsequent designation (Cox, 1927:28): Mitrella flaminea Risso, 1826 [= Mitrella

scripta (Linnaeus, 1758).

Mitrella cf. minor (Scacchi, 1836) (Figs. 3, 57)

Columbella minor Scacchi 1836

Material examined: Siracusa: 2 sh, 100 m, JHC. Malaga: 12 sh, 80 m, JHC. Marbella: 3 sh, 30-40 m,

FBC. Algeciras: 3sh, 18-22 m, FBC. Ma: 1 sh, 80 m, FSC. Pa: 4 sh, 80 m, WEC; 4 sh, 60-100 m, FSC.

GC: many sh, 12-520 m, JHC (Figs. 3, 57); 10 sh, 34-200 m, FBC. La: 1 sh, FSC. WS: many sh, 30-83

m, JHC; 12 sh, 30-60 m, FBC.

Taxonomy: In the recent literature, the

attribution of Columbella minor Scacchi,

1836 to the genus Mitrella Risso, 1826 is

generally preferred to the use of the

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary ls.

genus Columbellopsis Bucquoy and

Dautzenberg, 1882, specially created for

giving a distinct status to C. minor. In

fact, C. minor presents original morpho-

logic features in the anterior part of its

shell, with a narrow-: and sinuous

siphonal canal, a small triangular aper-

ture and a very concave left side of the

base. These features are clearly diver-

gent from the ones found in the other

Mitrella species ranging in the Lusitan-

ian Province, especially from the

Mediterranean M. scripta (Linnaeus,

1758), type species of Mitrella. The genus

Mitrella being applied to a vast array of

shell morphologies and being. still

waiting for a general revision, it seems

that the conservative way is more

appropriate in the present case and we

propose to keep the generic taxon

Mitrella for the placement of Columbella

minor.

LuQuE (1986) reports some differ-

ences between the shells from Canary

Islands attributed to “Mitrella minor”

and those from Mediterranean, western

Iberian Peninsula and northwest

Morocco. The shells from Canary

Islands are said to show a somewhat

different colour pattern and a lower

number of spiral striae at the base of the

last whorl. On this ground, LUQUE

remains reserved on the specific attribu-

tion of the Canarian population, which

is described as a new taxon in a com-

panion paper by BOYER AND ROLÁN

(2005). M. cf. minor from the Canary

Islands is reported and pictured as

“Mitrella svelta (Mtrs) Kobelt 1901” in

NORDSIECK AND GARCÍA-TALAVERA

(1979). M. svelta is a misspelling for M.

spelta (Kobelt, 1893), considered to be a

dubious species by VAN AARTSEN ET AL.

(1984), possibly matching the shallow

Mediterranean morph M. lanceolata

(Locard, 1886) belonging to the M.

scripta complex.

Distribution: Mitrella minor sensu

lato is distributed in Mediterranean and

from Vigo to northern Senegal. It is

widely distributed in Canary Islands

from 30 to about 500 m, apparently on

soft and detritic bottoms.

Remarks: The animals observed from

the Canary Islands are mottled of brown

and flecked of deep white dots on a

whitish ground (Fig. 57). This colour

pattern is very similar to the one

observed in specimens from Algeciras.

The oval operculum is light yellowish.

Mitrella pallaryi (Dautzenberg, 1927) (Figs. 2, 58)

Pyrene pallaryi Dautzenberg, 1927

Material examined: Marbella: 4 sh, 70-80 m, FBC. Alboran Island: 1 sh, 20 m, JHC. Pa: 15 sh, 150-

250 m, FBC. Go: 1 sh, JHC. Te: 2 sh, 60-100 m, JHC. GC: many sh, 60-520 m, JHC (Figs. 2, 58). La: 1

sh, 46-50 m, WEC. WS: 1 sh, 58 m, JHC.

Taxonomy: Pyrene pallaryi Dautzen-

berg, 1927 is placed in Mitrella by all the

recent authors. The use of the taxon

Pyrene is certainly unappropriate in the

present case, as the type species Pyrene

punctata (Bruguiere, 1789) has an ovate

outline, a somewhat turbinate top, a

narrow accent-shaped aperture and

very strong basal cords.

P. pallaryi shares most of the classic

shell features of Mitrella except for its

large size and for its turriculated spire.

However, we propose to keep the

species in Mitrella, following the last

reviewers (LUQUE, 1986; ROLÁN AND

TRIGO, 2000) and in the wait of a general

re-assessment of this group.

M. pallaryi is pictured under its right

specific name by NORDSIECK AND

GARCÍA-TALAVERA (1979), but associated

to the unusual subgenus Paratilia.

Distribution: The species is known to

range in circalittoral and upper bathyal

from Galicia and Mediterranean to

northern Angola, comprising the

Acores, Madeira and the Canary

Islands, but it may be a discontinuous

distribution, records being lacking south

Iberus, 23 (2), 2005

from Senegal to Congo (ROLÁN AND

TRIGO, 2000).

The species is widely distributed in

Canary Islands from 50 to about 500 m,

apparently on soft and detritic bottoms.

Remarks: The animals observed from

the Canary Islands are whitish, mottled

of reddish brown spots and patches

(Fig. 58). The sole is clearer. The opercu-

lum is subtranslucent, faintly square,

with a brownish violet patch, “Y”

shaped at its center.

LUQUE (1986) and ROLÁN AND TRIGO

(2000) gave details about the shell, the

protoconch, the operculum and the

radula in populations from continental

Spain and from Angola. The multispiral

protoconch has 3 to 3.5 smooth whorls,

and the species is considered to have a

planktotrophic development (ROLÁN

AND TRIGO, 2000). As expected in such a

case, M. pallaryi looks as being very con-

stant in its whole range of distribution.

Mitrella turbita (Duclos, 1840) (Figs. 4-6)

Columbella (Seminella) rac Dautzenberg, 1891

Material examined: GC: 2 sh, 10-15 m, WEC (Figs. 4-6). Fu: 3 sh, 0 m, MBC.

Taxonomy: Mitrella rac (Dautzenberg,

1891) was named from a material col-

lected in Dakar, Senegal, after a non-

binomial name given by Adanson. The

species named by DAUTZENBERG (1891)

is accurately described and pictured,

and it corresponds to one of the most

abundant and distinctive species of

Mitrella found about the Peninsula of

Cap Vert. However PELORCE AND BOYER

(2005) have shown that the name

Mitrella turbita (Duclos, 1840) has prece-

dence and must be used as the valid

name for this taxon.

The few shells found in Canary

Islands (Figs. 4-6) match perfectly the

material studied from Dakar. It must be

noted that most of the shells wear

strong sinuous axial ribs at the center of

the last whorl and strong spiral cords at

the base of the last whorl. As such, M.

turbita might be interpreted as an inter-

grade between the genera Mitrella and

Anachis. M. turbita is not recorded

neither pictured by NORDSIECK AND

GARCÍA-TALAVERA (1979).

Distribution: The species is well-

known from the Peninsula of Cap Vert

(Senegal), as restricted to different kinds

of hard bottoms from O to 40 m. It was

observed in Cape Blanco (northern

Mauritania) by E. Rolán (pers. comm.),

and it was recorded from Gran Canaria

through one sampling in ALONSO AND

JIMÉNEZ MILLÁN (1979). Its presence in

Gran Canaria and in Fuerteventura is

confirmed here by 2 new findings. This

thermophilic species is probably

restricted to the tepid shallow water

ranging off these two islands and it

(Right page) Figure 1. Columbella adansoní, 16 mm, 12 m, Sardina, Gran Canaria, JHC. Figure 2.

Mitrella pallaryi, 15 mm, 232 m, off Tasarte, Gran Canaria, JHC. Figure 3. Mitrella cf. minor, 11

mm, 150 m, off North West Gran Canaria, JHC. Figures 4-6. Mitrella turbita. 4, 5: 8.4 mm, 10-15

m, Gran Canaria, WEC; 6: 9.1 mm, 10-15 m, Gran Canaria, WEC. Figures 7-9. Mitrella bruggenz.

7: 12 mm, low tide, Orzola, Lanzarote, WEC; 8, 9: 9 mm, 1 m, Isla de Lobos, Fuerteventura, JHC.

(Página derecha) Figura 1. Columbella adansoni, 16 mm, 12 m, Sardina, Gran Canaria, JHC.

Figura 2. Mitrella pallaryi, 15 mm, 232 m, off Tasarte, Gran Canaria, JHC. Figura 3. Mitrella cf.

minor, 11 mm, 150 m, off North West Gran Canaria, JHC. Figuras 4-6. Mitrella turbita. 4 5: 8.4

mm, 10-15 m, Gran Canaria, WEC; 6: 9.1 mm, 10-15 m, Gran Canaria, WEC. Figuras 7-9. Mitre-

lla bruggeni. 7: 12 mm, marea baja, Orzola, Lanzarote, WEC; 8, 9: 9 mm, 1 m, Isla de Lobos, Fuer-

teventura, JHC.

-a

¡un

Ittora

Infrali

HERNÁNDEZ AND BOYER

Iberus, 23 (2), 2005

seems to reach there the northern limit

of its distribution.

Despite the fact that M. turbita has

not been recorded from Western Sahara

until now, it must be noted that this area

remains very poorly sampled as far as

hard bottoms are concerned, and there

is no concrete reasons, in the present

state, to believe that the scarce popula-

tions of M. turbita in the Canary Islands

are only relics of an older expansion of

the species during a past warmer pe-

riod, or even resulting from an acciden-

tal introduction coming from the human

industry.

Remarks: Only dead shells have been

collected in Canary Islands, so the animal

in these populations was not compared

with the animals from Senegal docu-

mented by the authors. However the few

shells studied from Canary Islands match

perfectly the most common shell mor-

phology and colour pattern found in M.

turbita off the Peninsula of Cap Vert.

Mitrella broderipi (Sowerby, 1844) (Figs. 10-18, 28-30, 40, 59)

Columbella broderipi Sowerby, 1844

Material examined: “Shallow form”: Málaga: 8 sh, 10 m, JHC. Estepona: 4 sh, 1-2 m, FBC. Algeci-

ras: many sh, 1-3 m, FBC. Cádiz: 5 sh, 0 m, JHC. Ceuta: 3 sh, 0 m, JHC. Alborán Island: 18 sh, 20 m,

JHC. Ma: 7 sh, 15-30 m, FSC. Selvagen Grande: many sh, FSC. Pa: 2 sh, FSC. GC: many sh, 0-135 m,

JHC (Figs 12, 29, 30, 59); many sh, 0-3 m, FBC (Figs. 10, 11, 13, 14); many sh, 0-15 m, FSC; many sh,

MEC. La: 19 sh, 0-2 m, JAC; many sh, 1-3 m, FBC; many sh, MBC; many sh, FSC. Fu: many sh, 0-2

m, JHC; 1 sh, 0 m, FBC (Fig. 15); many sh, MBC.

“Deep form”: Hi: many sh, 30-60 m, WEC (Figs. 16-18, 28). Go: many sh, 12 m, WEC. GC: 1 sh, 90-

96 m, JHC (Fig. 40). La: 9 sh, 46-80 m, WEC; many sh, 30-50 m, FSC.

Taxonomy: M. broderipi (Sowerby,

1844) was revised by VAN AARTSEN ET

AL. (1984) and by LUQUE (1986), with a

distribution limited to the Alboran Sea,

the Ibero-Moroccan Gulf and one

finding at a great depth in Acores

(DAUTZENBERG, 1927: 87). This last

record is dubious and seems better to be

a misidentification of Astyris profundi

(Dall, 1889) from the lower circalittoral

and the bathyal of Northern Atlantic

(ABBOTT, 1974). The phena M. broderipi

is in fact common in the Canary Islands,

but known under the junior name of

Nitidella ocellina Nordsieck, 1975, gener-

ally placed in Mitrella. In the same time

than M. ocellina, was also described

Pusionella scripta Nordsieck, 1975, which

belongs with evidence to the range of

variability of the same species. NORD-

SIECK AND GARCÍA-TALAVERA (1979) did

not use longer the taxon “P. scripta”, and

they pictured 3 shells of M. broderipi

under the name of “N. ocellina”. Both

taxa N. ocellina Nordsieck, 1975 and P.

scripta Nordsieck, 1975 are proposed

here as junior synonyms of M. broderipi.

Distribution: The species is con-

firmed to range on hard bottoms from

the Alboran Sea and the Ibero-Moroccan

(Right page) Figures 10-18. Mitrella broderipi. 10: 6.4 mm,1-2 m, Arinaga, Gran Canaria, EBC;

62 mm. 12m: Arinaga, Gran Canaria, FBC; 12: 7 mm, 12 m, Gando, Gran Canaria, JHC;

13: 6,2 mm, 1-2 m, Arinaga, Gran Canaria, FBC; 14: 6 mm, 1-2 m, Arinaga, Gran Canaria, FBC;

15: 6.9 mm, low tide, Granillo, Fuerteventura, FBC; 16: 5 mm, 30-55 m, Hierro, WEC; 17: 5.2

mm, 30-55 m, Hierro, WEC; 18: 5.1 mm, 30-55 m, Hierro, WEC.

(Página derecha) Figuras 10-18. Mitrella broderipi. 10: 6.4 mm, 1-2 m, Arinaga, Gran Canaria,

FBC; 11: 6.2 mm, 1-2 m, Arinaga, Gran Canaria, FBC; 12: 7 mm, 12 m, Gando, Gran Canaria,

JHC; 13: 6,2 mm, 1-2 m, Arinaga, Gran Canaria, FBC; 14: 6 mm, 1-2 m, Arinaga, Gran Canaria,

FBGC; 15: 6.9 mm, marea baja, Granillo, Fuerteventura, FBC; 16: 5 mm, 30-55 m, Hierro, WEC; 17:

5.2 mm, 30-55 m, Hierro, WEC; 18: 5.1 mm, 30-55 m, Hierro, WEC.

7Ó

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary Is.

Iberus, 23 (2), 2005

Gulf to Madeira and the Canary Islands.

In this last place, shallow water popula-

tions (0-15 m) and deeper water popula-

tions (12-60 m) show distinct shell mor-

phologies.

Remarks: The populations from

shallow water inhabiting the Alboran

Sea and the Canary Islands have been

compared in live conditions by the

authors: they perfectly match in all fea-

tures of the shells, of the soft parts and

of the operculum. They show the same

range of variability for the shell mor-

phology and colour pattern, and for the

chromatism of the soft parts. The

animals from the Canary Islands are

brownish to jet black (with blue shades

in this case) with whitish tips. Limited

zones are flecked of deep white dots

(Fig. 59). The sole is whitish to jet black.

The operculum is subtranslucent, faintly

square or more tear-shaped, with a dark

patch at its center.

The populations from the Canary

Islands ranging in deeper water (Figs.

16-18, 28) show generally a smaller,

lighter and more slender shell with a

dull chromatism, a higher spire with

more convex whorls and a thinner

labrum than in shallow water popula-

tions (Figs. 10-15, 29, 30). However,

some intergrades can be found (Fig. 40),

mostly from mid-infralittoral level

(shallow form, deep form, and inter-

grades are found in the lot from La

Gomera, WEC, collected in 12 m), with

similar protoconch and shell morphol-

ogy. As a matter of fact, the shell mater-

ial found in shallow water (0-3 m) and

the one from deeper water (30-60 m)

present a real unity. The animals from

deeper water were not examined and

the chromatism of their soft parts

remains unknown.

It must be noted that the shell mater-

ial collected under 10 m. in Alboran

Island (10-20 m) and in Madeira (15-30

m) does not differ from the “shallow

form” found everywhere, whereas the

shell material from the Canary Islands

found under 10 m represents mostly the

“deeper form” (12 m in La Gomera) or is

exclusively composed by it (30-60 m in

Hierro and Lanzarote).

On the ground of the elements at

hand, there is no reason to state on a

distinct taxonomic status of the “deeper

form” of M. broderipi from the Canary

Islands, and the transformation of the

shell morphology with the depth can be

interpreted as a phenetic adaptation to

different abiotic constraints (or a simple

variation randomly selected), geneti-

cally fixed but submitted to casual

reproductive mixing with “shallow

water” populations. This point deals

with the important question of the drift

of the genetic pool in such condition

and of the degree of genetic exchanges

between shallow and deeper popula-

tions. That refers to the topic of the

“deep forms” status in marine gas-

tropods, which waits for further investi-

gations.

Mitrella bruggeni van Aartsen, Menkhorst and Gittenberger, 1984 (Figs. 7-9,

37-39, 41-45, 55, 56)

Material examined: “Shallow form”: Malaga: 5 sh, 10 m, JHC. Algeciras: 2 sh, 1-3 m, JHC (Fig. 55);

16 sh, 1-3 m, FBC. Cadiz: 5 sh, 0 m, JHC. Ceuta: 1 sh, 0 m, JHC. Alboran Island: 5 sh, 20 m, JHC.

Ma: many sh, FSC. Selvagen Grande: 1 sh, 0 m, MNHN. Te: 2 sh, 1-2 m, FSC; 3 sh, 0 m, MNHN.

GC: 3 sh, 0-2 m, JHC; 1 fragment, 0 m, FBC. La: 19 sh, 0-2 m, FBC; 3 sh, 0-2 m, WEC; many sh, FSC.

Fu: 8 sh, 0-2 m, JHC (Figs. 8, 9). 1 sh, MBC (Fig. 7).

“Deep form”: Hi: 27 sh, 30-55 m, WEC (Figs. 37-39); GC: 5 sh, 15-90 m, JHC (Figs. 43-45, 56) La: 8

sh, 8-30 m, FBC; 15 sh, 46-50 m, WEC (Figs. 41, 42); 5sh, FSC. Fu: 1 sh, 0 m, JHC; 1 sh, BMC.

Taxonomy: Despite the statement of

LUQUE (1986), Mitrella bruggeni van

Aartsen, Menkhorst and Gittenberger,

1984 has priority over the name M. mal-

donadoi Luque, 1984, issued in an

abstract (LUQUE, 1984) which does not

match the requirements of the Code of

Nomenclature.

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary Is.

The shallow form of the species is

easily distinguished from its relatives in

Alboran Sea as well as in the Canary

Islands, due to its ventricose body

whorl, its regularly arched outer lip

with subequal labial denticles extended

on the inner wall, its slender pyramidal

spire, and its bulbous stepped proto-

conch (Figs. 55-58). M. bruggeni differs

from M. turbita by its stepped uni-

coloured browny to whitish protoconch

instead of domed whitish protoconch

with a light purple tip, the 6 whorls of

its teleoconch instead of 5, its wide oval

aperture instead or longer, narrower,

rather rectangular and slightly comma-

shaped, its 4 to 6 plaits on a very convex

columellar callus instead of 2 to 4 plaits

on a poorly convex callus, its subequal

labial teeth instead of a much stronger

tooth just below a small upper one, its

poorly marked spiral cords at the base

of the shell instead of strongly marked,

and its less incised siphonal canal. Even

if most of the shells of M. turbita show

sinuous axial ribs at the mid-part of the

last whorl, some specimens do not hold

and they are similar to M. bruggeni from

this point of view. Very few shells of M.

bruggent from Canary Islands and some

more from Mediterranean present a

colour pattern of white ocelles and axial

stripes on a reddish-brown ground com-

parable to the common “reticulated

pattern” found in M. turbita.

M. bruggent might be a possible junior

synonym of M. coccinea (Philippi, 1836).

The topic was tackled but not resolved by

PALMERI (1987) and CHIARELLI, MICALI

AND QUADRI (2003), and it is under study

by the second author.

M. bruggeni is pictured by NORD-

SIECK AND GARCÍA-TALAVERA (1979)

under the names of “Mitrella decollata

(Brusina, 1865)” and of “M. hidalgoi

Monterosato, 1889”.

Distribution: The species ranges on

hard bottoms in shallow water (0-3 m)

from the Alboran Sea to the Ibero-

Moroccan Gulf and from Madeira to the

Canary Islands. In this last place, popu-

lations from deeper levels (8-90 m)

present a smaller, lighter, and more

slender shell.

Remarks: The populations from shal-

low water inhabiting the Alboran Sea

and the Canary Islands (Figs. 7-9, 55, 56)

have been compared in live conditions:

they perfectly match together in all fea-

tures of the shells, on the soft parts and

of the operculum, and they show the

same range of variability for the mor-

phology and for the colour pattern of

the shells as well as for the chromatism

of the soft parts. The animal is light yel-

lowish to light beige with sparse light

brown patches on the foot and on the

siphon. The sole is light yellowish. The

head and tentacles are whitish. The fore-

head and the sides of the head have lon-

gitudinal light brown marks; the axis of

the tentacles is light brown. The sole, the

siphon and the tentacles are flecked of

deep white dots. The oval operculum is

light yellow amber.

The populations from the Canary

Islands ranging in deeper water (Figs.

37-39, 41-45, 57, 58) have a small lanceo-

late and subtranslucent shell, showing a

higher spire with more convex whorls

and a thinner labrum than in shallow

water populations. The deeper water

form seems to have been confused until

now by collectors with the sympatric M.

broderipi (Fig. 40). However, some inter-

grades between the shallow and the

deeper water forms of M. bruggeni can

be found (Fig. 41), mostly from the

middle levels (as most of the 8 shells

from southeast Lanzarote, 8-30 m, FBC),

and the general morphology of the

shells and of the protoconchs are

similar. The animals from deeper water

were not examined and the chromatism

of their soft parts remains unknown.

It must be noted that the shell mater-

ial collected below 10 m in Alboran

Island (10-20 m) does not differ from the

“shallow form” (0-3 m) found every-

where, whereas the shell material from

the Canary Islands found below 8 m ( 8-

90 m) represents only the deeper form

or intergrading morphs ( found in 8-30

m as well as in 46-50 m). Few shells of

the “deeper form” can be collected as

beached material in Fuerteventura,

together with shells of the “shallow

form”, without evident intergrades.

Iberus, 23 (2), 2005

These elements do not allow to infer

that two sibling species are represented

here. From a general point of view, the

situation is similar to the one found in M.

broderipi, with a shallow form and a

deeper form which seem to range accord-

ing to a bathymetric cline, and to present

a somewhat homogeneous morphology

at the population level. The casual

finding of shells of the “deeper form” at

the shore level may come from accidental

transports of live larvae or dead shells

due to local hydrodynamic conditions, or

to rejects of artisanal fisheries.

Mitrella ocellata (Gmelin, 1791) (Figs. 46-48)

Voluta ocellata Gmelin, 1791

Material examined: Ma: 4 sh, 0-17 m, FSC. Te: many sh, 0 m, FBC (Fig. 46). GC: many sh, 0-8 m,

JHC; 8 sh, 0-1 m, FBC (Fig. 47). La: 3 sh, 0 m, FBC (Fig. 48). Fu: many sh, 0 m, MBC.

Taxonomy: Depending on the

authors, M. ocellata (Gmelin, 1791) is

accepted as an amphiatlantic species or

as a pantropical one, but the matter is

still awaiting for further demonstration.

The M. ocellata complex is discussed by

RADWIN (1978 b), who cites several

sibling forms described from various

places in the Indo-Pacific and Panamic

Provinces. At least one of these sibling

forms, ranging in the Galapagos Archi-

pelago, can be separated at the specific

level on the ground of the characters of

its lateral radular tooth. One of the orig-

inal features of the shells, besides their

original subrectangular aperture with

strong labial denticles and their ocel-

lated colour pattern, is the usual lack of

the apex in the adult stage.

M. ocellata was described without

original locality. RADWIN (1978 b) gave

the Bahama Islands as subsequent type

locality of the species. The sibling morph

M. cribaria (Lamarck, 1822), frequently

used in the literature for the populations

ranging in Eastern Atlantic, was

described from the Java Seas. The name

M. canariensis (d'Orbigny, 1839), based on

a shell from Tenerife belonging to the

same complex M. ocellata, has not been

used in the literature for the West African

populations and rarely for the Canarian

populations. The shell pictured in D'ORr-

BIGNY (1839, pl. 6, figs. 35-37) as Buccinum

canariense shows a morphology similar to

that of M. ocellata, as far as the slender

oval outline of the body whorl, the long

pointed spire with flat to concave sides,

the very acute apex, and the long narrow

aperture are concerned. The dull shell

decoration matches the ocellated pattern

found in M. ocellata, as well as the alter-

nate subequal white and dark subsutural

square marks, and the dark spiral bands

at the mid-part of the body whorl on a

light chestnut brown ground. Despite the

presence of the apex in the shell pictured

in D'ORBIGNY (1839), which is generally

removed in adult shells of M. ocellata, and

despite the fact that the type material of

(Right page) Figures 19-27. Anachis avaroides. 19: SME syntype (as “holotype”), 7 mm, Gran

Canaria, SMF; 20: 4.6 mm, Sao Miguel, Acores, WEC; 21: 4.9 mm, 20 m, Funchal, Madeira,

WECGC; 22: 5 mm, 20 m, La Palma, WEC; 23: 5.4 mm, 20-30 m, San Sebastián, La Gomera,

WEC; 24: 6 mm, 30-55 m, El Hierro, WEC; 25: 5.9 mm, 30-55 m, El Hierro, WEC; 26: 5.8

mm, 30-55 m, El Hierro, WEC; 27: 5 mm, 30-55 m, El Hierro, WEC.

(Página derecha) Figuras 19-27. Anachis avaroides. 19: syntype SMF (como “holotipo”), 7 mm, Gran

Canaria, SMF; 20: 4.6 mm, Sao Miguel, Agores, WEC; 21: 4.9 mm, 20 m, Funchal, Madeira, WEC;

22: 5 mm, 20 m, La Palma, WEC: 23: 5.4 mm, 20-30 m, San Sebastián, La Gomera, WEC; 24: 6

mm, 30-55 m, El Hierro, WEC: 25: 5.9 mm, 30-55 m, El Hierro, WEC; 26: 5.8 mm, 30-55 m, El

Hierro, WEC; 27: 5 mm, 30-55 m, El Hierro, WEC.

“Y

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50)

Ittora

Infrali

HERNÁNDEZ AND BOYER

Iberus, 23 (2), 2005

B. canariense was not examined, there is

no serious reasons to doubt about the

identity of this taxon, considered here as

junior synonym of M. ocellata. 1t must be

noted that some adult shells of M. ocellata

from Tenerife (G. Hervillard Collection)

and from Dakar (FBC) were observed to

have kept their apex. The fact that the

type material of B. canariense was found

“in roots of gorgonas, fished off the

harbour of Orotava” (D'ORBIGNY, 1839)

does not signify necessarily the occur-

rence of a deep water species. The very

steep slopes encountered along the coasts

of Tenerife cause currently the fall of live

mollusca or of shells from shallow water

to deeper levels.

NORDSIECK AND GARCÍA-TALAVERA

(1979) pictured as “Nitidella ocellata

(Gmelin, 1889)” a shell matching the

common form of M. ocellata found in

Canary Islands (Figs. 46-48), itself per-

fectly similar, as far as the shell mor-

phology and colour pattern are con-

cerned, to the populations ranging in

Cape Verde Islands, Senegal and

Caribbean. On the same plate, NORD-

SIECK AND GARCÍA-TALAVERA (1979) pic-

tured as “Nitidella canariensis (d'Or-

bigny, 1839)” a reddish shell with an

intact apex and with dark spiral bands

under the suture and at the mid-part of

the last whorl. Even if uncommon, this

form must be accepted within the

natural variability of M. ocellata.

We propose to use provisionally the

name M. ocellata for the whole Eastern

Atlantic populations, thus accepting the

possible occurrence of a united amphiat-

lantic species, and we propose to consider

provisionally B. canariense d'Orbigny, 1839

as junior synonym of M. ocellata. However,

the hypothesis of a sibling species in

Eastern Atlantic waters, genetically and /or

reproductively distinct from the Caribbean

population, constitutes a possible alter-

native, due to the presence of a rather

short protoconch (1.5 to 2.0 whorls with a

coiled bulging top) supposed to be non-

planktotrophic, that means having an

intracapsular metamorphosis or a very

short free-swimming larval stage

(lecitotrophic non-feeding mode). Such a

protoconch leads normally to a limited

ability of dispersion, to a strong capacity

of reproductive isolation and to the for-

mation of distinct species at local or at

regional scale. That is clearly the case in

most of the Mitrella species known to us,

which present both a short paucispiral

protoconch and a limited distribution.

Distribution: The Eastern Atlantic pop-

ulations of M. ocellata range on hard

bottoms in very shallow water (intertidal

to 3 m) from Acores to Madeira and the

Cape Verde Islands, and from Western

Sahara to Senegal. The species is distrib-

uted in the whole Canary Archipelago. It

was not found in Agadir nor in Gambia

(pers. obs.) or in Ghana (P. Ryall pers.

comm.) and in the rest of the Gulf of

Guinea (in the literature). The record from

Santa Helena must be confirmed to deal

really with the same species.

Remarks: The soft parts of the animals

from the Canary Islands are dark to-

bacco brown, the whitish tip of the tenta-

cles and few whitish zones on the foot or

the head being flecked of deep white

dots. The sole is whitish. The same chro-

matism of the animal was observed in

the populations from Senegal.

(Right page) Figures 28-30. Mitrella broderipi. 28: 5.5 mm, 55-60 m, El Hierro, WEC; 29: 24: 6.5

mm, low tide, Caleta de Abajo, Gran Canaria, JHC; 30: 7 mm, 12 m, Gando, Gran Canaria,

JHC. Figures 31-36. Anachis avaroides. 31: 6.1 mm, 55-60 m, El Hierro, WEC; 32: 5.5 mm, 55-

60 m, El Hierro, WEC; 33: 5.6 mm, 55-60 m, El Hierro, WEC; 34: 6.7 mm, 55-60 m, El Hierro,

WEG; 35: 6.1 mm, 55-60 m, El Hierro, WEC; 36: 5.4 mm, 55-60 m, El Hierro, WEC.

(Página derecha) Figuras 28-30. Mitrella broderipi. 28: 5.5 mm, 55-60 m, El Hierro, WEC; 29: 24:

6.5 mm, marea baja, Caleta de Abajo, Gran Canaria, JHC; 30: 7 mm, 12 m, Gando, Gran Canaria,

JAC. Figuras 31-36. Anachis avaroides. 31: 6.1 mm, 55-60 m, El Hierro, WEC; 32: 5.5 mm, 55-60

m, El Hierro, WEC; 33: 5.6 mm, 55-60 m, El Hierro, WEC: 34: 6.7 mm, 55-60 m, El Hierro, WEC:

35: 6.1 mm, 55-60 m, El Hierro, WEC: 36: 5.4 mm, 55-60 m, El Hierro, WEC.

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Iberus, 23 (2), 2005

Genus Anachis H. and A. Adams, 1853

Type species by subsequent designation (Tate, 1868:13): Columbella scalarina Sowerby, 1832.

Anachis avaroides Nordsieck, 1975 (Figs. 19-27, 31-36)

Material examined: Acores: many sh, 0-15 m, FBC; 1 sh, 6 m, WEC (Fig. 20); 9 sh, 8 m, FSC. Ma: 2

sh, 9-12 m, JHC; 4 sh, 14-21 m, WEC (Fig. 21); many sh, FSC. Pa: 1 sh, 6 m, JHC (Fig. 22); 6 sh, 20-

40 m, WEC; 2 sh, 40 m, FSC. Hi: 1 sh, 5-12 m, JAC; many sh, 30-60 m, WEC (Gigs 24-27, 31-36);

many sh, FSC. Go: 4sh, 20-30 m, WEC (Fig. 33). Te: 2 sh WEC. GC: 1 sh, SMF syntype, stored as

“holotype” (Fig. 19); 3 sh, 9-12 m, JHC. La: 1 sh, 46-50 m, FSC.

Taxonomy: Anachis avaroides Nord-

sieck, 1975 was described on the basis of

a dark “grey brown” subadult shell of

6.5 x 2.5 mm (NORDSIECK, 1975: 6, fig.

29), said to come from Las Palmas (Gran

Canaria), and explicitely designated as

holotype in the original description

(referred as collection number Nr 73.35).

The original description does not deal

with any paratype and does not suggest

the study of further shells. The shell

stored as “holotype” in SMF (Fig. 19),

labelled as coming from Gran Canaria

with no register or collection number,

measures 7.0 x 2.75 mm and presents a

light orange colour ground. This shell

has the same slender stepped spire than

the type-figure, the same subadult outer

lip and a similar macro sculpture of

axial ribs and spiral cords at the base of

the last whorl. Its colour pattern shows

however a spiral decoration of white

marks on the shoulder and on the spiral

cords at the base of the last whorl,

whereas the type-figure shows only a

spiral row of white marks at the middle

of the last whorl. So it can be stated that

the so-said SME “holotype” is not the

holotype originally designated by

NORDSIECK (1975). As any revision or

any new type designation did not occur

about this topic, and in the wait of the

possible rediscovery of the authentic

holotype, it is felt to be more appropri-

ate to consider the so-said SME “holo-

type” as a simple syntype subsequently

joined to the type lot by F. Nordsieck.

The original attribution of the species

to the genus Anachis was clearly founded

on the presence of strong spiral ribs. A.

avaroides was compared by NORDSIECK

(1975) to the Caribbean A. avara (Say,

1822), which shows however a more

spindle-shaped outline, a more pointed

apex, less numerous axial ribs, a more

slender aperture and a more vertical

outer lip (RADWIN, 1978 a: fig.3). NORD-

SIECK AND GARCÍA-TALAVERA (1979) pic-

ture as “Anachis avaroides F. Nordsieck,

1975”, a shell very similar to our Fig. 25.

They also picture as “Anachis atomella

(Duclos, 1840)” a shell which looks like a

subadult of A. avaroides. The type of A.

atomella, examined in MNHN, is a very

different species belonging to the Indo

Pacific Province.

Curiously, NORDSIECK AND GARCÍA-

TALAVERA (1979) do not give Gran

Figures 37-39. Mitrella bruggeni. 37: 6.6 mm, 30-55 m, El Hierro, WEC; 38: 7 mm, 30-55 m, El

Hierro, WEC; 39: 6.6 mm, 30-55 m, El Hierro, WEC. Figure 40. Mitrella broderipi, 9 mm, 90-96

m, Arinaga, Gran Canaria, JHC. Figures 41-45. Mitrella bruggeni. 41: 7.9 mm, 46-50 m, Puerto del

Carmen, Lanzarote, WEC; 42: 8.2 mm, 46-50 m, Puerto del Carmen, Lanzarote, WEC; 43, 44: 9

mm, 40-60 m, Puerto de La Luz, Gran Canaria, JHC; 45: 9 mm, 40-60 m, Puerto de La Luz, Gran

Canaria, JHC.

Figuras 37-39. Mitrella bruggeni. 37: 6.6 mm, 30-55 m, El Hierro, WEC; 38: 7 mm, 30-55 m, El

Hierro, WEC; 39: 6.6 mm, 30-55 m, El Hierro, WEC. Figura 40. Mitrella broderipi, 9 mm, 90-96 m,

Arinaga, Gran Canaria, JHC. Figuras 41-45. Mitrella bruggeni. 41: 7.9 mm, 46-50 m, Puerto del Carmen,

Lanzarote, WEC; 42: 8.2 mm, 46-50 m, Puerto del Carmen, Lanzarote, WEC: 43, 44: 9 mm, 40-60

m, Puerto de La Luz, Gran Canaria, JHC: 45: 9 mm, 40-60 m, Puerto de La Luz, Gran Canaria, JHC.

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HERNÁNDEZ AND BOYER

Iberus, 23 (2), 2005

Canaria in the distribution of A.

avaroides, but only La Palma, Selvagen

and Porto Santo (Madeira). It is sug-

gested here that the type material of A.

avaroides may well come from La Palma,

where the species is abundant in moder-

ate depths, better than from the type

locality of “Las Palmas” (Gran Canaria),

where the species looks as being very

scarce and as ranging at deeper levels.

Distribution: The species is known

from the Acores, Madeira, Selvagen

Islands and the Canary Islands, but it is

not recorded from the continental shelf.

Off the Acores and Madeira, shells are

commonly found in the beach drift or in

moderate depths (0-20 m), whereas the

species is generally found at deeper

level in the Canary Islands (20-60 m),

rarely in shallower water. It seems that

the species is somewhat common in

hard bottom environments off the

western Canary Islands (lower infralit-

toral and upper circalittoral) but very

uncommon in the central and eastern

Canary Islands.

Remarks: A. avaroides shows as a

rather variable phena. Most of the shells

(Figs. 20-24, 26, 31, 34-36) are stout and

thick, with strong axial ribs and a some-

what stepped outline. However, few

shells show a more slender outline (Figs.

19, 25), sometimes with very faint axial

ribs (Fig. 27) or just limited to the 2 or 3

first whorls (Figs. 32, 33). All the inter-

grades exist, even with M. broderipi

which presents a similar range of varia-

tion of the shell outline and of the aper-

ture (Figs. 10-18, 28-30, 40), and the same

diversified patterns of the shell chroma-

tism (Figs. 28-36). The protoconch is the

same in A. avaroides and in M. broderipi.

Both phenae “A. avaroides” and “M.

broderipi” have been collected together

in several places, and each one looks as

representing a tip of the morphologic

cline of one single species. The matter

remains however to be accurately veri-

fied. One of the most contradictory

point lies in the fact that where A.

avaroides is abundant (for instance in the

lower infralittoral from Hierro), it is

mixed with the slender “deeper form”

of M. broderipi (Figs. 16-18) and the pos-

sible morphologic intergrades (Fig. 27)

are very scarce and unclear. It must be

noted that the slender “deeper form” of

M. broderipi never suggests a tendency

to axial ribbing, neither to the formation

of fine spiral striae at the top of the

whorls as it often occur in A. avaroides,

even in poorly ribbed shells (Figs. 32,

33). That means that, in case where A.

avaroides would be a “deep form” of M.

broderipi, it would range in apparent

syntopy (at least in Canary Islands) with

another “deep form” of the same species

without evident intergrade.

If we consider the phenetic complex

“M. broderipi / A. avaroides” as a whole,

it presents a much larger variability of

the shell morphology and of the colour

pattern in Madeira and in the Canary

Islands (the highest variability being

recorded from the western Canary

Islands), whereas the populations from

Alboran Sea (with only the M. broderipi

shallow morph) and from the Acores

(Right page) Figures 46-48. Mitrella ocellata. 46: 9.2 mm, low tide, Tenerife, EBC; 47: 10.5 mm,

low tide, Tarajalillo, Gran Canaria, FBC; 48: 10.2 mm, low tide, Tarajalillo, Gran Canaria, FBC.

Figures 49-51. Zafra exilis. 49: 3.5 mm, 40 m, San Cristobal, Gran Canaria, J. Ferreiro Coll.; 50: 3,2

mm, 2-3 m, Pasito Blanco, Gran Canaria, FBC; 51: 3.5 mm, 40 m, San Cristobal, Gran Canaria,

JHC. Figures 52-54. Parvanachis obesa. 52: 4.9 mm, 9 m, Santa Cruz de Tenerife, WEC; 53: 5.2

mm, 9 m, Santa Cruz de Tenerife, ESC; 54: 4.4 mm, 9 m, Santa Cruz de Tenerife, FSC.

(Página derecha) Figuras 46-48. Mitrella ocellata. 46: 9.2 mm, marea baja, Tenerife, EBC; 47: 10.5

mm, marea baja, Tarajalillo, Gran Canaria, FBC; 48: 10.2 mm, marea baja, Tarajalillo, Gran

Canaria, EBC. Figuras 49-51. Zafra exilis. 49: 3.5 mm, 40 m, San Cristobal, Gran Canaria, J.

Ferreiro Coll.; 50: 3,2 mm, 2-3 m, Pasito Blanco, Gran Canaria, FBC: 51: 3.5 mm, 40 m, San Cristo-

bal, Gran Canaria, JHC. Figuras 52-54. Parvanachis obesa. 52: 4.9 mm, 9 m, Santa Cruz de Tenerife,

WEC; 53: 5.2 mm, 9 m, Santa Cruz de Tenerife, FSC; 54: 4.4 mm, 9 m, Santa Cruz de Tenerife, FSC.

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HERNÁNDEZ AND BOYER

Iberus, 23 (2), 2005

(with only the A. avaroides morph) show

as much less variable. If the specific

unity of this complex would be con-

firmed in the future, the reduced vari-

ability occurring in Alboran Sea and in

the Acores may result from a “founder

effect”. Further inquiries are required

about this topic, and in the present state

we feel more appropriate to consider A.

avaroides as a possible sibling species of

M. broderipi.

The phena A. avaroides has appar-

ently never been collected in live condi-

tions but only as shells.

Genus Parvanachis Radwin, 1968

Type species by original designation: Buccinum obesum C.B.Adams, 1845.

Parvanachis obesa (C.B.Adams, 1845) ( Figs. 52-54)

Buccinum obesum C.B.Adams, 1845

Material examined: Te: 5 sh, 9 m, WEC (Fig. 52); 5 sh, 9 m, FSC (Figs. 53, 54).

Taxonomy: Parvanachis obesa (C. B.

Adams, 1845) was revised by RADWIN

(1978 a) who characterizes the genus Par-

vanachis as gathering “the stout, promi-

nently ribbed columbellids with inflated

body whorl and heavily thickened,

flaring apertural lips”, all features well

represented in P. obesa. Radwin stresses

on the diagnostic feature represented by

“the strongly down-hooked proximal

cusp of the lateral radular tooth ”, con-

sidered as typical of Parvanachis. In fact,

P. obesa shows as very distinct from the

Anachis species found in Eastern

Atlantic, by its inflated body whorl, its

lattice patterned sculpture, the rounded

shape of the thick outer lip and the

strong upper denticle.

The few shells found in the harbour of

Santa Cruz de Tenerife do not differ from

the shells found in Lesser Antilles. The

presence of P. obesa in Eastern Atlantic was

never recorded in the literature.

Distribution: The species was

described from Jamaica and it is known

to have a large range in Western

Atlantic, from Maryland to central

Uruguay, that means well beyond the

limits of the Caribbean Province. In

Eastern Atlantic, the species is only

known from Tenerife where few shells

were collected by diving in the harbour

of the main town of Santa Cruz, at a

depth of 9 m (1982-1983).

Remarks: This record of P. obesa in

Tenerife clearly corresponds to an acci-

dental introduction by shipping. Several

of the shells being in very fresh condi-

tion, some with the dry animal inside,

the presence of a live population is

Figures 55, 56. Mitrella bruggeni, protoconchs. 55: 1-3 m, Algeciras, Andalucía, JHC; 56: 40-60

m, Puerto de la Luz, Gran Canaria, JHC. Figure 57. Mitrella cf. minor, animal, shell length 10

mm, 150 m, off northwest Gran Canaria, JHC. Figure 58. Mitrella pallaryi, animal, shell length

14 mm, 200 m, off northwest Gran Canaria, JHC. Figure 59. Mitrella broderipi, animal, shell

length 7 mm, 30 m, San Cristobal, off northeast Gran Canaria, JHC. Figure 60. Zafra exilis,

animal, shell length 3 mm, 30 m, San Cristobal, off northeast Gran Canaria, JHC.

Figuras 55, 56. Mitrella bruggeni, protoconchas. 55: 1-3 m, Algeciras, Andalucía, JHC; 56: 40-60 m,

Puerto de la Luz, Gran Canaria, JHC. Figura 57. Mitrella cf. minor, animal, largo de concha 10 mm,

150 m, noroeste de Gran Canaria, JHC. Figura 58. Mitrella pallaryi, animal, largo de concha 14 mm,

200 m, noroeste de Gran Canaria, JHC. Figura 59. Mitrella broderipi, animal, largo de concha 7

mm, 30 m, San Cristobal, noreste de Gran Canaria, JHC. Figura 60. Zafra exilis, animal, largo de

concha 3 mm, 30 m, San Cristobal, noreste de Gran Canaria, JHC.

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary Is.

Iberus, 23 (2), 2005

assumed. However, any new sampling

was not recorded along the last twenty

years, and it does not seem that the

species has spread out in Tenerife or

even survived in the harbour of Santa

Cruz.

Genus Zafra A. Adams, 1860

Type species by monotypy: Zafra mitriformis A. Adams, 1860.

Zafra exilis (Philippi, 1849) (Figs. 49-51, 60)

Columbella exilis Philippi, 1849

Material examined: Te: 2 sh, 3-8 m, JHC. GC: 7 sh, 2-200 m, JHC (Figs. 51, 60); many sh, 2-3 m, FBC

(Fig. 50); 1 sh, 40-60 m, J. Ferreiro Coll (Fig. 49).

Taxonomy: The species was cited and

redescribed from the Red Sea by DRIVAS

AND JaY (1997). Despite a somewhat

variable decoration, the shell shows a

very homogeneous morphology and

cannot be confused with any of the

other Indo Pacific Zafra.

The genus Zafra was revised by

DRIVAS AND Jay (1990), but it remains a

poorly characterized group, not clearly

distinguished for instance from the

genus Seminella Pease, 1868 and from

the genus Ascalista Drivas and Jay, 1990.

Zafra exilis is recorded from Gran

Canaria by SEGERS AND SWINNEN (2003)

as the first mention of this Indo Pacific

species in the Atlantic waters.

Distribution: Z. exilis, described from

Aden, is endemic to the Red Sea and to the

Gulf of Aden. First discovered as one shell

in 1993 by P. Segers on the South East coast

of Gran Canaria, and as live specimens in

2001 by A. M. Garcia at 5 m off Santa Cruz

de Tenerife, the species is overall well

settled in Gran Canaria, where live spec-

imens were collected by both authors, by

E. Swinnen and by J. Ferreiro, all around

the island in shallow to deep waters.

Remarks: SEGERS AND SWINNEN (2003)

explained that the perfect correspon-

dence of the specimens collected in the

Canary Islands with the material exam-

ined from the Red Sea leads to consider

that the population from the Canary Is-

lands comes from a human introduction.

We can add that this introduction is

recent and that we are witnessing to the

progressive settling of a new species in the

Canary Islands. As a matter of fact, the

species reaches shallow waters in its Indo

Pacific distribution like in Canary Islands,

where it constitutes currently dense set-

tlements (for instance, at 2-3 m in algae on

rocks in the small bay of Pasito Blanco and

in the harbour of Arguineguin, southern

Gran Canaria, FBC). So it is very unlikely

that the settling of a population in Gran

Canaria, where active collectors are sam-

pling the shallow fauna regularly since

the seventies, might remain undiscovered

for a long. NORDSIECK AND GARCÍA-TALAV-

ERA (1979) did not record the species and

any of the assiduous collectors in the place

(except P. Segers with one shell in 1993)

did not find any trace of the species before

the years 2000.

Because the international harbour of

Las Palmas is the most evident place for

an accidental introduction of such an exotic

species (for example by cleaning the ballast

tanks of trade ships), it can be assumed

that the species has spread out from Las

Palmas towards the northwestern and the

southern tips of the Island. As the first dis-

covery occurred in 1993 about 55 km south

from Las Palmas, and considering the time

required for the dispersion of a species

supposed to have an intracapsular meta-

morphosis, it can be assumed that the

introduction of the species dates about

from the beginning of the eighties.

The animals from Gran Canaria are

whitish with the nape and the sides of

the foot light to dark brown (Fig. 60).

The chromatism of the animals from

Indo Pacific waters is unknown.

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary ls.

Genus Nassarina Dall, 1889

Type species by original designation: Nassarina bushii Dall, 1889.

Nassarina rietae Segers and Swinnen, 2004

Material examined: Pa: 1 sh, 42 m (paratype FSC).

Taxonomy: Nassarina rietae was

described from 3 shells collected at 42 m

off La Palma, Canary Islands. Any other

record was not made about this species

in the literature, and it was not found in

the whole material studied in public

and private collections.

N. rietae differs from the other Nassa-

rina species from Western Atlantic

(RADWIN, 1978 a) mainly in its rather

wide subrectangular aperture with short

and wide siphonal canal instead of

small oval aperture with longer and

narrow siphonal canal in Western

Atlantic species. N. rietae differs from

the Nassarina species from Senegal and

Guinea Bissau (PELORCE AND BOYER,

2005) mainly in its stouter outline with

CONCLUSIONS

The columbellid fauna from the

Canary Islands is made of an assem-

blage of species belonging to different

biogeographic sets.

A first group comprises species

restricted to the Lusitanian Province:

Mitrella- broderipi ranges principally

from the Alboran Sea to the Canary

Islands, being rare off Madeira and

lacking in the Acores, whereas the

closely related morph Anachis avaroides

is restricted to the northern Macarone-

sian Islands (from the Acores to the

Canary Islands), and Mitrella bruggeni

ranges from the Canary Islands, north-

west Morocco and Madeira to Alboran

Sea, southern Italy and Tunisia (as M.

coccinea in CHIARELLI, MICALI AND

QUADRI, 2003).

A second group comprises species

ranging from the latitudes of southern

Morocco to a limited part of the West

African province: the morph Mitrella cf.

much inflated whorls and in its few and

very strong axial ribs with wide inter-

vals.

Distribution: Only known from La

Palma, type locality.

Remarks: N. rietae does not seem to

belong to the Caribbean fauna, neither

to the fauna from the West African

Province. The single record from the iso-

lated place of La Palma (SEGERS AND

SWINNEN, 2004) suggests a local

endemism better than the introduction

of an Indo Pacific species. The issue

remains however to be fully checked, as

the hard bottoms from upper circalit-

toral were mainly uncollected off

Canary Islands like off western Morocco

and Western Sahara.

minor found off the Canary Islands

extends to northern Senegal, and

Mitrella turbita ranges from southern

Canary Islands to central Senegal.

A third group comprises plank-

totrophic species presenting a wide but

possibly fragmented distribution from

the Lusitanian Province to the equator-

ial latitudes: Columbella adansoni ranging

in the whole Macaronesian Archipela-

gos and from Sierra Leone to northern

Angola, and Mitrella pallaryi ranging

from Galicia and Mediterranean to

Senegal and being found also in north-

ern Angola. The case of Mitrella ocellata,

ranging from Madeira to Senegal and

possibly also in Santa Helena, is some-

what different, as the present distribu-

tion of this supposed amphiatlantic

species may result from several different

ways of spreading.

A fourth group is composed of sup-

posed introduced species from

Iberus, 23 (2), 2005

Caribbean or Indo Pacific origin, among

which only Zafra exilis seems to have

settled successfully. The positive record

of two introduced columbellids species,

the well-established Zafra exilis and the

elusive Parvanachis obesa, tends to

demonstrate that a high potential of

introduction of tropical and subtropical

species of Columbellidae occurs in the

Canary Islands.

Two shells attributable to the

Caribbean Steironepion monilifera

(Sowerby, 1844) were observed in the F.

Swinnen Collection, labelled as

“Canary Islands, from fishermen”. The

poor precision of this datum and the

fact that the species is not cited in the

literature neither observed in other

molluscan collection from the Canary

Islands lead to consider this reference

as not fully reliable. However, such an

occurrence can be appreciated as per-

fectly plausible. It is possible that a

local introduction of S. monilifera failed

after few generations, like it seems to be

the case for P. obesa. Such a situation of

failed introductions can be expected as

a current process, and the successful

introductions, like observed with Z.

exilis, are probably the less common

result. The high frequency of accidental

introductions of marine molluscs is

probably under-estimated, but it can be

reasonably considered as a direct by-

product of the contemporary maritime

economy.

The single case of possible columbel-

lid endemism in the Canary Islands may

be that of Nassarina rietae Segers and

Swinnen, 2004, only known through 3

shells from La Palma. This finding sug-

gests that lower infralittoral or upper

circalittoral new species of Columbelli-

dae may remain to discover off the

Canary Islands, especially on hard

bottoms in the most superficially

explored areas, such as Fuerteventura or

the lesser western Islands.

The cases of M.bruggeni and of the

complex M. broderipi / A. avaroides

require a clarification of the biologic

status of the “deeper forms”. The irreg-

ular distribution of our considered

“deeper forms” (apparently absent, for

instance, in the Alboran Sea) suggests

that the influence of environmental

factors such as the bathymetric pres-

sure are not fully explicative of the

morphologic differences at work. A

relative genetic autonomy between

shallow and deep populations seems to

occur and would explain the relative

homogeneity observed in each “bathy-

metric form” as well as the irregular

bathymetric and geographic distribu-

tions of the morphs.

The case of M. turbita and of the

complex M. broderipi / A. avaroides show

that the genera Mitrella and Anachis are

not separated by significant differences.

The diagnostic value of the axial ribs is

contested by its irregular presence in M.

turbita and by the continuous morpho-

logic cline represented by this feature in

the complex M. broderipi / A. avaroides.

The presence of spiral cords (irregularly

represented in A. avaroides) and of spiral

grooves (irregularly represented in M.

bruggeni) seems to follow the same

pattern. The poor reliability of these

morphologic features as diagnostic cri-

teria at the specific level leads to con-

sider them as not reliable diagnostic cri-

teria at the generic level.

This point must be considered like a

complementary argument for a review-

ing of the supraspecific classification of

the Columbellidae, and like a guideline

for the reinterpretation of the discrimi-

nating criteria within this family.

ACKNOWLEDGEMENTS

We are greatly indebted to Winfrid

Engl (Dússeldorf, Germany) and to

Frank Swinnen (Lommel, Belgium) for

the loan of their columbellids collection

from the Macaronesian Islands. A

special thanks is due to Rudo von Cosel

(Muséum national d'Histoire naturelle,

Paris), José Ferreiro (Las Palmas, Gran

Canaria) and Francisco Sicilia (Arrecife,

Lanzarote) for the various kinds of help

dedicated to us during the course of this

study.

HERNÁNDEZ AND BOYER: Infralittoral and circalittoral Columbellids of the Canary ls.

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luscos dei Norte de Gran Canaria (Es-

paña).Cuadernos de Ciencias Biológicas, 2 (29):

105-115.

BOYER, F. AND ROLAN, E., 2005. About a sibling

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Basteria, supplt. n* 2: 1-135.

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La famille Columbellidae (Gastropoda: Muricoidea) dans

Pinfralittoral de la Péninsule du Cap Vert (Sénégal)

The family Columbellidae (Gastropoda: Muricoidea) in the infralit-

toral of the Peninsula of Cap Vert (Senegal)

Jacques PELORCE* et Franck BOYER**

Recibido el 15-X11-2004. Aceptado el 6-X-2005

RÉSUMÉ

Les especes de Columbellidae représentées dans l'infralittoral de la Péninsule du Cap Vert

[Sénégal) sont étudiées sur la base de la morphologie de leur coquille et du chromatisme

de leurs parties molles. Neuf espéces anciennement décrites, parfois sous des noms tom-

bés en désuétude, sont reconnues: Columbella rustica Linné, 1758; Anachis cuspidata

Marrat, 1877 avec A. emergens Fischer-Piette et Nicklés, 1946 comme synonyme plus

récent; A. freytagi V. Maltzan, 1884 avec A. bubakensis Lamy, 1923 comme synonyme

plus récent; Parvanachis aurantia (Lamarck, 1822) avec C. cancellata Gaskoin, 1851

comme synonyme plus récent; Mitrella denticulata ([Duclos, 1840) avec C. triangulifera V.

Maltzan, 1884 comme synonyme plus récent; M. melvilli (Knudsen, 1956); M. ocellata

(Gmelin, 1791); M. psilla (Duclos, 1846) avec C. japix Duclos, 1850 comme synonyme

plus récent; M. turbita (Duclos, 1840) avec C. phylina Duclos, 1846 et C. rac Dautzen-

berg, 1891 comme synonymes plus récents. l'holotype de A. emergens Fischer-Piette et

Nickles, 1946 est désigné comme néotype de A. cuspidata Marrat, 1877. Des lectotypes

sont désignés pour A. freytagi, M. denticulata, M. psilla et M. turbita. La Péninsule du Cap

Vert est désignée comme localité type pour A. cuspidata, P. aurantia, M. denticulata, M.

psilla et M. turbita. Deux espéces de Mitrella, M. inflata sp. nov. et M. fimbriata sp. nov.,

et deux espéces de Nassarina, N. procera sp. nov. et ÑN. rolani sp. nov., sont décrites

comme nouvelles.

ABSTRACT

The species of Columbellidae represented in the infralittoral of the Peninsula of Cap Vert

(Senegal) are studied on the basis of their shell morphology and of the chromatism of their

soft parts. Nine species formerly described, sometimes under names fallen into disuse, are

recognized: Columbella rustica Linné, 1758; Anachis cuspidata Marrat, 1877 with A.

emergens Fischer-Piette and Nicklées, 1946 as junior synonym; A. freytagi V. Maltzan,

1884 with A. bubakensis Lamy, 1923 as junior synonym; Parvanachis aurantia (Lamarck,

1822) with C. cancellata Gaskoin, 1851 as junior synonym; Mitrella denticulata (Duclos,

1840) with C. triangulifera V. Maltzan, 1884 as junior synonym; M. melvilli (Knudsen,

1956); M. ocellata (Gmelin, 1791); M. psilla (Duclos, 1846) with C. japix Duclos, 1850

as junior synonym; M. turbita (Duclos, 1840) with C. phylina Duclos, 1846 and C. rac

Dautzenberg, 1891 as junior synonyms. The holotype of A. emergens Fischer-Piette and

Nickles, 1946 is designated as neotype of A. cuspidata Marrat, 1877. Lectotypes are

designated for A. freytagi, M. denticulata, M. psilla and M. turbita. The Peninsula of Cap

* 289 voie Les Magnolias, 30240 Le Grau du Roi, France, e-mail: pelorceCfree.fr

** 110 chemin du Marais du Souci, 93270 Sevran, France, e-mail: franck.boyer6Gwanadoo.fr

Iberus, 23 (2), 2005

Vert (central Senegal) is designated as type locality for A. cuspidata, P. aurantia, M. den-

ticulata, M. psilla and M. turbita. Two species of Mitrella, M. inflata sp. nov. and M. fim-

briata sp. nov., and two species of Nassarina, N. procera sp. nov. and N. rolani sp. nov.,

are described as new.

RESUMEN

Se estudian las especies de Columbellidae representadas en el infralitoral de la Península

de Cap Vert (Senegal) en base a la morfología de las conchas y el cromatismo de partes

blandas. Se reconocen como válidas nueve especies previamente descritas, algunas de

ellas con nombres caídos en desuso: Columbella rustica Linné, 1758; Anachis cuspidata

Marrat, 1877 con A. emergens Fischer-Piette y Nicklés, 1946 como sinonímia; A. freytagi

V. Maltzan, 1884 con A. bubakensis Lamy, 1923 como sinonímia; Parvanachis aurantia

(Lamarck, 1822) con C. cancellata Gaskoin, 1851 como sinonimia; Mitrella denticulata

(Duclos, 1840) con C. triangulifera V. Maltzan, 1884 como sinonímia; M. melvilli (Knud-

sen, 1956); M. ocellata (Gmelin, 1791); M. psilla (Duclos, 1846) con C. japix Duclos,

1850 como sinonimia; M. turbita (Duclos, 1840) con C. phylina Duclos, 1846 y C. rac

Dautzenberg, 1891 como sinonímias. Se designa el holotipo de A. emergens Fischer-

Piette y Nicklés, 1946 como neotipo de A. cuspidata Marrat, 1877. Se designan lectoti-

pos para A. freytagi, M. denticulata, M. psilla y M. turbita. La Peninsula de Cap Vert (cen-

tro de la costa Senegalesa) se designa como localidad tipo para A. cuspidata, P.

aurantia, M. denticulata, M. psilla y M. turbita. Dos especies de Mitrella, M. inflata sp.

nov. y M. fimbriata sp. nov., así como dos especies de Nassarina, N. procera sp. nov. y

N. rolani sp. nov., se describen como nuevas.

MOTS CLE: Columbellidae, Columbella, Mitrella, Anachis, Parvanachis, Nassarina, espéce nouvelle, variabili-

té, diversité, Cap Vert, Sénégal.

KEY WORDS: Columbellidae, Columbella, Mitrella, Anachis, Parvanachis, Nassarina, new species, variability,

diversity, Cap Vert, Senegal.

PALABRAS CLAVE: Columbellidae, Columbella, Mitrella, Anachis, Parvanachis, Nassarina, especie nueva,

variabilidad, diversidad, Cap Vert, Senegal.

INTRODUCTION

L'exploration de l'infralittoral de la

Péninsule du Cap Vert a permis aux

auteurs de découvrir localement, dans

une zone située a la rencontre des

influences lusitaniennes et guinéennes,

une faune de Columbellidae d'une

richesse inédite pour 1'Afrique de

l'Ouest. Treize morphospecies ont pu

étre en effet séparées la oú NICKLES

(1950) n'en reconnaissait que cinq.

La grande variabilité de la plupart de

ces especes, la confusion entretenue dans

la littérature autour de l'identité de plu-

sieurs d'entre elles et la disponibilité de

plusieurs noms anciens tombés en désué-

tude mais référables aux formes exami-

nées ont conduit les auteurs á mener une

étude systématique de la variabilité des

formes représentées et a procéder a la révi-

sion de l'ensemble des taxons considérés.

Le présent article expose les résultats

de ce travail, qui conduit a reconnaítre

comme valides dans l'infralittoral de la

Péninsule du Cap Vert 9 especes ancien-

nement décrites et á décrire comme

nouvelles 4 especes distribuées dans la

méme zone.

MATÉRIEL ET MÉTHODES

Plus de 2.000 sujets de Columbelli-

dae ont été récoltés par les auteurs au

cours de 11 séjours á Dakar de 1995 a

2002 (Fig. 1). Description des stations

comme suit.

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

1438'10"N

1723'45"W

Figure 1. Stations de récoltes.

Figure 1. Sampling stations.

Sta. 1 Au large de lle de N'Gor: 20-

25 m, plateformes et tombants basal-

tiques.

Sta. 2 Plage devant lle de N'Gor: 0-

3 m, brossage et lavage de petits blocs.

Sta. 3 Pointe des Almadies: 0-2 m, zone

des marées, tamisage et ramassage á vue,

sable et rochers, lessivage d'algues cal-

caires et de colonies de Palytoa.

Sta. 4 Fann-Plage: 0-1 m, zone des

marées, tamisage et ramassage á vue,

sable et rochers.

Sta. 5 Faillis: 23 m, éperon de roches

basaltiques avec poches de sable.

Sta 6 Les Madeleines: 7-20 m,

tombant rocheux avec poches et sable

grossier en lisiere.

Sta 7 Anse Bernard: 0-2 m, zone des

marées, petits et gros rochers sur sable.

Sta. 8 Les Blockaus: 10-17 m, amon-

cellement artificiel de rochers d'endi-

guement.

<=. 0

18 19 20

Sta. 9 Le Tacoma: 5-13 m, épave du

début des années 40, sable coquiller en

périphérie.

Sta. 10 Bel Air-Port: 10-12 m, fonds

détritiques et coquilles de Pinna.

Sta. 11 Banc de Bel Air: 9-15 m,

dalles calcaires et rochers bas avec

algues et sable, poches importantes de

coquiller détritique.

Sta. 12 Mbao, cimetiere de bateaux:

0-10 m, épaves récentes sur sable

vaseux.

Sta. 13 Kunakhe: 30-34 m, fond

rocheux avec poches de sable,.bouquets

de vers chaetopteres.

Sta. 14 Gouye Teni M'Both: 25-30 m,

banc rocheux plat avec bouquets de

vers.

Sta. 15 Vézo: 34-39 m, épave de plus

de 50 ans sur sable coquiller.

Sta. 16 Tiwa: 30-37 m, épave de plus

de 50 ans sur sable coquiller.

Iberus, 23 (2), 2005

Sta. 17 Orstom: 32-33 m, épave

récente sur sable coquiller.

Sta. 18 Séminole: 27 m, banc

rocheux, bouquets de vers.

Sta. 19 Epopal: 33 m, banc de roche

avec bouquets de vers et gorgones.

Sta. 20 Charbomnier: 29-34 m, épave

de plus de 50 ans sur sable coquiller.

Sta. 21 Grand Thiouriba: 37-40 m,

banc rocheux avec gros blocs et poches

de sable, bouquets de vers.

Sta. 22 Petit Thiouriba: 30-35 m, banc

rocheux avec gros blocs et poches de

sable, bouquets de vers.

Sta. 23 Kunk Diabar: 30-45 m, haut-

fond rocheux isolé avec poches et sable

en lisiere.

Sta. 24 Bargny: 5-10 m, sable vaseux

avec algues.

Sta. 25 Dakar: provenance indéter-

minée, autour de la presqu'ile du Cap

Vert entre 0 et 40 m de profondeur.

Sta. 26 Guédiawaye: 37 m, fond

rocheux avec poches de sable, bouquets

de vers.

Sta...2/ Epave) Ds ol epave

ancienne sur sable coquillier.

Sta. 28 Ker Diop: 46 m, fond rocheux

avec poches de sable.

Sta. 29 Sud-est de Gorée: 15-20 m,

dragage sur sable et débris coquilliers.

Une dizaine de récoltes a vue ont été

effectuées á pied et en apnée dans les

petits fonds a l'extréme nord-ouest

(Pointe des Almadies) et a l'extréme

sud-ouest (Fann et Anse Bernard) de la

Péninsule du Cap Vert. Une centaine de

récoltes ont été effectuées en scaphandre

autonome par brossage sur fonds durs

entre 3 et 40 metres, la plupart dans le

sud de la Péninsule, une au large de l'le

de N'Gor dans le nord-ouest de la Pé-

ninsule et une autre au large de la Petite

Cóte, Centre Sénégal (station Kunk Dia-

bar). Une quinzaine de demi-journées

ont été consacrées a des opérations de

dragage dans la Baie de Gorée, dans le

sud /sud-est de la Péninsule, sur sable et

débris coquilliers entre 7 et 20 m. Une

vingtaine de récoltes effectuées en sca-

phandre autonome dans le sud de la Pé-

ninsule (10-40 m) par Haidar El-Ali (Da-

kar), Patrice Petit de Voize (Quimper) et

Patrick Boyer (Saint-Martin de Brome)

dans la période 1998-2000 ont été mises

a la disposition des auteurs. Quelques

lots de coquilles provenant de la Baie de

Gorée et des Almadies ont été offerts

par Marcel Pin (Dakar).

Une fraction importante du matériel

récolté a été étudiée et a fait 1'objet de

comparaisons a l'état vivant. Cette frac-

tion du matériel a été conservée dans

l'alcool (CFB), le reste du matériel (CFB

et CJP) ayant été conservé a l'état sec.

Dans la plupart des formes examinées, á

l'exception de deux especes spéciale-

ment élusives cantonnées dans l'infralit-

toral inférieur, llanimal vivant a été

dessiné, photographié ou décrit, et la

variation du chromatisme des parties

molles a été corrélée avec la variation de

la morphologie et de la décoration des

coquilles. Les observations, les croquis

et les notes réalisés par Emilio Rolán

(Vigo) a loccasion de deux séjours á

Dakar (2002 et 2003) ont été versés a la

documentation du présent travail.

Les types de Columbellidae déposés

au Muséum de Paris et référables aux

provinces lusitanienne et ouest-africaine

ont été étudiés, ainsi que le matériel type

des Columbellidae de von Maltzan confié

par le Muséum de Berlin, celui de

Knudsen par le Muséum de Copen-

hague, et les types de Buccinum aurantium

Lamarck par le Muséum de Geneve.

Acronymes et abréviations:

IRSNB Institut Royal des Sciences Natu-

relles de Belgique, Bruxelles.

MHNG Muséum d'Histoire Naturelle

de Geneve.

MNCN Museo Nacional de Ciencias

Naturales, Madrid.

MNHN Muséum National d'Histoire

Naturelle, Paris.

ZMK Zoologisk Museum, Kobenhavn.

ZMB Zoologisk Museum, Berlin.

CER Collection privée d'Emilio Rolán.

CFB Collection privée de Franck Boyer.

CJP Collection privée de Jacques

Pelorce.

spm: spécimen, sujet récolté vivant.

coq: coquille, sujet récolté mort.

Sta.: station de récolte; coll: collection.

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

SYSTEMATIQUE

Famille COLUMBELLIDAE Swainson, 1840

Sous-famille COLUMBELLINAE Swainson, 1840

Genre Columbella Lamarck, 1799

Espece type par monotypie: Voluta mercatoria Linné, 1758

Columbella rustica (Linné, 1758) (Fig. 11)

Purpura 1. Le Siger: Adanson, 1757 (pré-Linnéen, non-disponible).

Voluta rustica Linné, 1758: 731.

Voluta punctata Allan, 1818.

Colombella guifordia Risso, 1826.

Colombella gualteriana Risso, 1826.

Voluta tringa sensu Costa O.G., 1829 non Lamarck, 1811.

Colombella elongata Philippi, 1836.

Colombella rustica var striata Duclos, 1846.

Colombella spongiarum Duclos, 1846.

Columbella rustica var apiculata Pallary, 1900.

Columbella rustica var cuneatiformis Pallary, 1900.

Matériel type: Non examiné.

Autre matériel examiné: CJP: Sta. 1: 1 coq; Sta. 2: 70 spm; Sta. 7: 3 spm, 1 coq. CFB: Sta. 2: 20 spm,

1 cog; Sta. 3: 1 spm, 3 cog; Sta. 4: 12 spm; Sta. 7: 14 spm, 15 coq.

Localité type: Mer Méditerranée.

Description des sujets du Centre

Sénégal: Coquille de taille moyenne, 10 a

14 mm, ovale, dernier tour représentant

un peu plus de la moitié de la hauteur

totale de la coquille, périostracum fin et

pratiquement transparent, stries longi-

tudinales sur la totalité de la.coquille,

protoconque paucispirée de 15 a 2

tours, spire haute á 5 tours convexes;

décoration relativement constante, com-

posée de taches claires ou marron clair

sur fond marron foncé; ouverture

étroite, labre épaissi extérieurement et

renflé au milieu du bord intérieur, 6 a 9

dents labiales blanches aux intervalles

teintés de marron, bord columellaire

occupé par un léger cal transparent

formant 3 plis.

Animal blanc laiteux á blanc créeme

décoré de larges plages en écharpe

ambre a brunátre. Opercule variable, de

petit, blanchátre et arrondi a grand, jau-

nátre et fortement échancré.

Distribution: Méditerranée et cótes

continentales de l'Est-Atlantique du

Golfe Ibéro-Marocain jusqu'au Sénégal,

fonds durs de la zone littorale.

Remarques: Les spécimens de Dakar

(Fig. 11) correspondent parfaitement a la

variété striata de DUCLOS (1846). Autour

de la Péninsule du Cap-Vert comme dans

le reste de sa distribution, C. rustica est

principalement représentée dans la zone

de balancement des marées, et tres rare-

ment en-dessous de la zone des 2-3 m.

Au plan phénotypique, C. rustica ne se

distingue de sa jumelle C. adansoni

Menke, 1853 que par sa protoconque

paucispirée suggérant un développe-

ment larvaire intracapsulaire sans stade

planctotrophe (MOOLENBEEK ET HOENSE-

LAAR, 1991). L'important polymorphisme

représenté chez C. rustica recouvre au

moins partiellement la variabilité rencon-

trée dans la morphologie et la décoration

de la coquille de C. adansoni. C. rustica ne

se distingue apparemment pas non plus

de C. adansoni par son habitat ni par la

structure de ses populations adultes. C.

adansoni semble n'étre distribuée que

dans les archipels macaronésiens et le

long de la cóte ouest-africaine de la

région guinéenne jusqu'a 1'Angola, sans

recouvrement avec la distribution de C.

Iberus, 23 (2), 2005

rustica. Un effort supplémentaire d'obser-

vation et de récolte portant sur les popu-

lations du Sénégal reste nécessaire pour

vérifier ce point définitivement, ainsi que

pour apprécier si la forme striata Duclos

peut étre considérée, par comparaison

avec la variabilité exprimée par l'espece

en Méditerranée, comme rendant compte

d'une forme géographique affirmée déte-

nant un statut de sous-espece.

Sousfamille PYRENINAE Suter, 1909

Genre Anachis H. et A. Adams, 1853

Espece type par désignation subséquente (Tate, 1868): Columbella scalarina Sowerby, 1832

Anachis cuspidata Marrat, 1877 (Figs. 10, 30-38)

Columbella (Anachis) cuspidata Marrat, 1877. Quarterly Journal of Conchology, 1: 42.

.Pyrene (Anachis) emergens Fischer-Piette et Nickles, 1946. Journal de Conchyliologie, 87: 61-62.

Matériel type: Le matériel type de Anachis cuspidata est perdu (Mc MILLAN, 1985: appendix). Le

matériel type de Anachis emergens (MNHN) est constitué d'un holotype de 9 mm provenant de la

Collection Adanson (Fig. 30) et de 22 paratypes d'un autre lot mesurant de 7 a 8.5 mm (Figs. 31,

32). L'holotype de A. emergens est désigné ici comme néotype de A. cuspidata.

Autre matériel examiné: CJP: Sta. 1: 1 spm, Sta. 2: 92 spm, 1 cog; Sta. 5: 1spm, 1 cog; Sta. 6: 1 spm;

Sta. 9: 4 coq; Sta. 14: 1 spm; Sta. 15: 6 spm, 32 coq; Sta. 16: 7 spm, 9 cog; Sta. 23: 5 spm, 52 coq. CFB:

Sta. 2: 19 spm; Sta. 3: 9 spm;, Sta. 6: 1 spm;, Sta. 7: 6 spm, 1 cog; Sta. 13: 6 spm; Sta. 14: 5 spm, 1 cog;

Sta. 15: 1 spm; Sta. 16: 3 spm; Sta. 18: 27 spm, 1 cog; Sta. 21: 6 spm; Sta. 22: 42 spm, 7 cog; Sta. 26: 37

spm, 16 cog; Sta. 27: 3 spm; Sta. 28: 1 spm.

Localité type: « Afrique de l'Ouest ». La Péninsule du Cap Vert, Centre Sénégal, est ici proposée

comme localité type plus pertinente.

Description originale: “C. testa elon-

gato-fusiformi, utrinque attenuata, spira

cuspidata; anfractibus longitudinaliter

costatis et transversim sulcatis, costis

subgranulatis; fulva fusco maculata vel

pallide cinerea, epidermide leucophoea;

columella arcuata, labio cum callo cir-

cumscripto tecto; apertura angusta;

labro intus lirato.

Hab. West Africa. Captain Davis”.

Description complémentaire: Coquille

de petite taille, 7 a 8 mm, de forme

allongée, légerement ventrue, environ

1.5 tour de protoconque et 5 tours de

téléoconque; 20 a 22 cótes axiales sur le

dernier tour, intervalles légérement plus

étroits que les cótes, minces cordons

spiraux dans les intervalles chevauchant

plus ou moins les cótes en conférant

souvent a celles-ci un aspect granuleux

vers la base et prés de la suture, cótes

axiales et cordons spiraux s'estompant

au niveau de l'ouverture, 10 sillons

spiraux á la base; décoration variable,

composée de taches marron plus ou

100

moins foncées sur fond blanchátre, pro-

toconque blanc creme avec petites

taches marrons, périostracum transpa-

rent; ouverture étroite, bord columel-

laire présentant un cal bien défini et

étroit portant 4 ou 5 plis, bord interne

du labre épaissi dans son milieu et

portant 5 ou 6 dents, canal siphonal

court et légerement vrillé.

L'animal présente une couleur de

fond blanchátre á jaunátre pále avec une

décoration variable de marques brun

léger á noirátre un peu étirées dans le

sens de la longueur, souvent plus mor-

celées, marbrées ou arrondies sur les

bords du pied, liste dorsale et nuque

plus foncées, taches parfois mordorées

sur le siphon avec anneau sombre vers

l'extrémité, moucheté de blanc sur ten-

tacules, extrémité du siphon et sole, qui

peut étre faiblement marbrée de noir ou

de brun sur le bord latéral. Opercule

jaunátre faiblement moucheté de noir.

Certains spécimens de profondeur

(Fig. 38) possedent une coquille légere-

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

Figure 2. Colombella turbita, planche 2 de DucLOs (1840), figure type. Figure 3. Colombella

phylina, planche 15 de DucLos (1846), figure type. Figure 4. Colombella denticulata, planche 9 de

DucLos (1840), figure type. Figure 5. Buccinum aurantium, planche 25 de KIENER (1834). Figure

6. Colombella psilla, planche 15 de DucLos (1846), figure type. Figure 7. Colombella japix,

planche 22 de DucLos (1850), figure type. Figure 8. Clathurella polignaci Lamy, 1923, figure

type. Figure 9. Columbella (Anachis) bubakensis Lamy, 1923, figure type. Figure 10. “Le Rac non

publié” in FISCHER-PIETTE (1942), figure type de Pyrene (Anachis) emergens Fischer-Piette et

Nicklés, 1946.

Figure 2. Colombella turbita, plate 2 in DUCLOS (1840), type figure. Figure 3. Colombella phylina,

plate 15 in DUCLOS (1846), type figure. Figure 4. Colombella denticulata, plate 9 in DUCLOS

(1840), type figure. Figure 5. Buccinum aurantium, planche 25 in KIENER (1834). Figure 6. Colom-

bella psilla, plate 15 in DUCLOS (1846), type figure. Figure 7. Colombella japix, plate 22 in DUCLOS

(1850), type figure. Figure 8. Clathurella polignaci Lamy, 1923, type figure. Figure 9. Columbella

(Anachis) bubakensis Lamy, 1923, type figure. Figure 10. “Le Rac non publié” in FISCHER-PIETTE

(1942), type figure of Pyrene (Anachis) emergens Fischer-Piette and Nickles, 1946.

101

Iberus, 23 (2), 2005

ment translucide de couleur miel uni-

forme. Les caracteres conchyliologiques

de ces spécimens et le chromatisme de

lVanimal sont néanmoins strictement

identiques á ceux des spécimens litto-

raux, et ces différentes formes sont

considérées comme conspécifiques. La

variabilité des coquilles dans les popu-

lations de l'infralittoral supérieur porte

moins sur la morphologie de la coquille

que sur la décoration de celle-ci (Figs.

33, 38). Les granulosités représentées

sur la coquille de certaines populations

littorales sont bien apparentes chez les

paratypes de A. emergens (Figs. 31, 32).

Distribution: L'espece est unique-

ment connue de la Péninsule du Cap

Vert, Centre Sénégal, 0-40 m. Elle n'a pas

été reconnue de la région de Nouadhi-

bou, dans le nord mauritanien (comm.

pers. d'E. Rolán), ni de la Gambie (obs.

pers. du second auteur).

Habitat: Fonds durs, sous pierres

avec tapis d'algues courtes et frondes

d'algues calcaires sur plages fossiles ou

sur blocs basaltiques.

Remarques: A partir d'un lot de

coquilles ramenées par le Capitaine

Davis d'un voyage effectué de Madere

jusqu'au Golfe de Guinée, MARRAT

(1877) décrit Columbella (Anachis) cuspi-

data comme espece nouvelle de

l'Afrique de l'Ouest. La description ori-

ginale, reproduite ci-dessus, n'est

accompagnée d'aucune illustration et

d'aucun commentaire. L'espece sera

citée par PACE (1902) sans tentative de

révision.

FISCHER-PIETTE (1942) retrouvera

dans la collection Adanson une coquille

collée sur un carton-cuvette portant

l'inscription “2,482, Buccin 4 Rac, hist.

Nat. Du Seneg.”. Cette coquille est citée

par Fischer-Piette comme “le Rac non

publié” (Fig. 10), par opposition avec

« Le Rac décrit et figuré », et elle est rap-

prochée de l'espéce strenella Duclos de

la région indo-pacifique. Fischer-Piette

écarte C. cuspidata de la comparaison

avec la coquille d'Adanson, considérant

ce taxon comme correspondant á une

“espece ouest-africaine, décrite trop

sommairement et non figurée”.

FIsCHER—PIETTE ET NICKLES (1946), se

fondant sur l'étude complémentaire

d'un lot de 22 coquilles obtenues plus

récemment de Dakar, créeront le taxon

Pyrene (Anachis) emergens pour désigner

l'espece relatée plus tót par

FISCHER—PIETTE (1942) comme “le Rac

non publié”, le spécimen de la collection

Adanson étant désigné comme holotype

d'A. emergens. Cette nouvelle espece est

comparée aux taxa spécifiques strenella

Duclos, sparsa Reeve, suffusa Sowerby et

cuspidata Marrat, considérés comme

synonymes possibles d'emergens.

Discussion: La description de A. cus-

pidata, attribuée par MARRAT (1877) a

l'Afrique de 1'Ouest, méme si elle est

relativement breve et non illustrée, cor-

respond sans ambiguité a l'espece

décrite ultérieurement du Sénégal

comme A. emergens par FIsCHER—PIETTE

ET NICKLES (1946). Bien que le nom de

A. cuspidata soit tombé en désuétude, la .

regle d'inversion de la précédence ne

s'applique pas ici, le taxon emergens

ayant été employé dans moins de 25

travaux publiés par moins de 10 auteurs

dans les 50 ans qui viennent de s'écouler

(Art. 23.9.1 du Code de Nomenclature

Zoologique). Par conséquent le nom cus-

pidata Marrat, 1877 doit étre considéré

comme nom valide de l'espéce, avec

emergens Fischer-Piette et Nickles, 1946

comme synonyme plus récent. Le type

de cuspidata étant perdu, nous dési-

gnons l'holotype de C. emergens

(MNHN, Collection Adanson) comme

néotype de A. cuspidata Marrat. Les taxa

spécifiques strenella Duclos, suffusa

Sowerby et sparsa Reeve correspondent

a des especes indo-pacifiques et n'ap-

partiennent pas a la synonymie de A.

cuspidata.

Anachis freytagi von Maltzan, 1884 (Figs. 9, 21-29)

Columbella (Anachis) freytagi von Maltzan, 1884. Diagnosen neuer Senegambischer Gastropoden,

Nahrichtsblatt der deutschen Malakozoologischen Gesellschaft, 5: 72.

102

PELORCE ET BOYER: Columbellidae dans lP'infralittoral de la Péninsule du Cap Vert

Figure 11. Columbella rustica, plage de N'Gor, 13.9 mm (CJP). Figure 12. Mitrella ocellata, plage de

N'Gor, 11.4 mm (CJP). Figure 13. Mitrella melvillz, Vézo, 7.2 mm (CJP). Figures 14-17. Mitrella

denticulata. 14: lectotype 6.8 mm (MNHN); 15: syntype de C. triangulifera, 5.9 mm (ZMB); 16,

17: Cap Vert, 6 et 7.5 mm (CJP). Figures 18-20. Parvanachis aurantia. 18: syntype probable, 8.7

mm, Coll. Lamarck (MHNG); 19: 7.6 mm, Coll. Duclos (MNAHN); 20: Vézo, 7.7 mm (CJP).

Figure 11. Columbella rustica, N'Gor beach, 13.9 mm (CJP). Figure 12. Mitrella ocellata, N'Gor

beach, 11.4 mm (CJP). Figure 13. Mitrella melvilli, Vézo, 7.2 mm (CJP). Figures 14-17. Mitrella

denticulata. 14: lectotype 6.8 mm (MNHN); 15: syntype of C. triangulifera, 5.9 mm (ZMB); 16, 17:

Cap Vert, 6 and 7.5 mm (CJP). Figures 18-20. Parvanachis aurantia. 18: probable syntype, 8.7 mm,

Coll. Lamarck (MHNG); 19: 7.6 mm, Coll. Duclos (MNHN); 20: Vézo, 7.7 mm (CJP).

Columbella (Anachis) bubakensis Lamy, 1923. Campagne du Sylvana, Mission du Comte Jean de

Polignac et de Mr Louis Gain Mollusques Testacés. Compte rendu du Congrés des Sociétés

savantes en 1922:. 13-14.

Matériel type: 5 syntypes (4 adultes et un juvénile) ZMB, Moll. 37022: le mieux conservé d'entre

eux (7 mm) est sélectionné ici comme lectotype (Fig. 21), les 4 paralectotypes (Fig. 22) mesurent de

5.9 a 7 mm. Holotype de C. bubakensis (7.2 mm) au MNHN (Fig. 23).

Autre matériel examiné: CJP: Sta. 1: 2 spm, 1 cog; Sta. 6: 11 spm, Sta. 9: 169 spm, 47 cog; Sta. 14:

1spm, Sta. 15: 1spm, 27 cog; Sta. 16: 1 spm, 8 coqg; Sta. 21: 1 spm, Sta. 23: 13 spm, 4 coq. CFB: Sta. 6:

20 spm, Sta. 9: 10 spm, 46 cog; Sta. 10: 45 spm, Sta. 13: 3 spm; Sta. 17: 2 spm;, Sta. 21: 1 spm; Sta. 26:

5 cog; Sta. 29: 6 coq.

Localité type: Gorée, Sénégal.

Description: Coquille de petite taille, représentant 45 % de la hauteur totale,

6 a 8 mm, allongée, spire pointue, proto- suture bien marquée, tours pratique-

conque a 1.5 tour, blanche avec bande ment plats; 13 a 22 cótes verticales lége-

marron, dernier tour de la coquille rement sigmoides atténuées vers la base

103

Iberus, 23 (2), 2005

des tours, minces cordons spiraux dans

les intervalles, parfois absents; perios-

tracum fin et transparent, couleur de

fond variable jaune á marron violacé, le

plus souvent brun tabac, avec ou sans

taches blanches, réguliéres ou irrégu-

liéres; ouverture étroite, sinus paléal et 6

dents sur un labre épais avec renflement

extérieur, cal columellaire bien marqué

et étroit portant 5 a 7 plis, canal siphonal

court et légerement vrillé.

L'animal blanchátre est décoré de

marbrures noires, plus denses et plus

foncées sur le corps que sur le dessus du

pied, le front entre les pédoncules ocu-

laires est noir, comme la partie interne

des tentacules et les bords latéraux

supérieurs de la sole. Les parties noires

peuvent étre fragmentées en segments

courts et minces accotés pour former

des bandes ou des taches organisées

dans le sens longitudinal. Siphon décoré

de taches noires réguliéres et d'un

anneau terminal de la méme couleur, le

fond blanchátre tendant á devenir noi-

rátre dans sa partie distale. Moucheté de

blanc sur les cótés et sur les bords du

pied ainsi que vers l'extrémité des tenta-

cules. Dessous de la sole blanchátre a

grisátre avec taches noires sur les bords.

Opercule jaunátre, ovale arrondi, parfois

étiré dans sa moitié arriére ou dentelé

dans sa partie avant, zone d'attache-

ment noire en forme de selle dans la

moitié antérieure.

Distribution: De la Péninsule du Cap

Vert, Centre Sénégal, jusqu'aux lles Bis-

sagos, Guinée-Bissau.

Habitat: Abondante sur tous les

fonds durs et intermédiaires de 10 a 40

metres.

Remarques: Anachis freytagi montre

une grande variabilité concernant le

nombre et le relief des cótes axiales et des

cordons spiraux (les cótes pouvant étre

tres atténuées et les cordons absents chez

certains sujets) comme pour la décoration

de la coquille. Le type de C. bubakensis

(Figs. 9, 23) appartient clairement a la

variabilité de A. freytagí (Figs. 24-29).

Genre Parvanachis Radwin, 1968

Espece type par désignation originale: Buccinum obesum C.B. Adams, 1845

Parvanachis aurantia (Lamarck, 1822) comb. nouv. (Figs. 5, 18-20)

Buccinum 3. Le Nisot: Adanson, 1757 (pré-Linnéen, non-disponible).

Buccinum aurantium Lamarck, 1822. Histoire Naturelle des animaux sans vertebres, 7: 275.

Buccinum aurantium Lam.: Kiener, 1834. Spécies Général et Iconographie des Coquilles Vivantes,

Famille des Purpuriferes, Deuxieme Partie, 1834-1841: 50.

Colombella aurantiía Lam.: Duclos, 1840. Histoire Naturelle Générale et Particuliere de tous les genres de

Coquille Univalves Marines. Genre Colombella. Planche 7.

Columbella cancellata Gaskoin, 1851. Description of twenty species of Columbellae, and one

species of Cypraea. Proc. Zool. Soc. London, 19: 6.

Matériel type: 14 syntypes probables MHNG 1452/95 (7 a 8.7 mm) (Fig. 18).

Autre matériel examiné: Collection Duclos (MNHN): 10 coq. CJP: Sta. 1: 2 cog; Sta. 2: 3 spm, Sta.

6: 2 coq; Sta. 8: 3 coq; Sta. 9: 7 spm, 6 cog; Sta. 12: 1 coq; Sta. 14: 3 spm, 1 coq; Sta. 15: 1 cog; Sta. 16:

1 juv; Stra. 22: 2 juv; Sta. 23: 5 spm, 7 coq. CFB: Sta. 2: 2 spm, Sta. 3: 2 spm, 3 cog; Sta. 6: 3 spm; Sta.

7:25 spm; Sta. 9: 4 spm, 20 cog; Sta. 13: 9 spm; Sta. 14: 9 cog; Sta. 18: 13 spm, Sta. 21: 1 spm, Sta. 22:

15 spm, 9 cog; Sta. 26: 11 cog; Sta. 28: 1 spm.

Localité type: «Mon cabinet». Selon Kiener, l'espece «habite les cótes de l'ile de Java». La Pénin-

sule du Cap Vert, Centre Sénégal, est désignée ici comme nouvelle localité type de Buccinum auran-

tium Lamarck.

Description originale: «Buccinum

aurantium. B. testa minima, ovato-

acuata, longitudinaliter es tenuissime

plicata, obsoleta decussata, luteo-auran-

tia, apica rubra, anfractibus convexo-

planis; apertura angustiuscula».

104

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

Figures 21-29. Anachis freytagíi. 21: lectotype, 7 mm (ZMB); 22: paralectotype, 6.45 mm (ZMB);

23: holotype de A. bubakensis, 7.2 mm (MNHN); 24, 25: Le Tacoma, 7.7 et 7.3 mm (CJP); 26:

Gouye Teni M'Both, 7 mm (CJP); 27-29: Le Tacoma, 8.2, 7.4 et 7 mm (CJP). Figures 30-38.

Anachis cuspidata. 30: néotype de A. cuspidata, holotype de A. emergens, 9 mm (MNHN); 31, 32:

paratypes de A. emergens, 8 et 7 mm (MNHN); 33: N'Gor, 8 mm (CJP); 34: Les Madeleines, 7.5

mm (CJP); 35, 36: N'Gor, 7.5 et 7.5 mm (CJP); 37, 38: Tiwa, 7.2 et 7 mm (CJP).

Figures 21-29. Anachis freytagi. 21: lectotype, 7 mm (ZMB); 22: paralectotype, 6.45 mm (ZMB); 23:

holotype of A. bubakensis, 7.2 mm (MNHN); 24, 25: Le Tacoma, 7.7 and 7.3 mm (CJP); 26: Gouye

Teni M'Both, 7 mm (CJP); 27-29: Le Tacoma, 8.2, 7.4 and 7 mm (CJP). Figures 30-38. Anachis

cuspidata. 30: neotype of A. cuspidata, holotype of A. emergens, 9 mm (MNAN); 31, 32: paratypes of

A. emergens, 8 and 7 mm (MNAHN); 33: N'Gor, 8 mm (CJP); 34: Les Madeleines, 7.5 mm (CJP);

35, 36: N'Gor, 7.5 and 7.5 mm (CJP); 37, 38: Tiwa, 7.2 and 7 mm (CJP).

105

Iberus, 23 (2), 2005

Description complémentaire: Selon

KIENER (1834): “Coquille tres petite,

ovale, oblongue, atténuée aux extrémi-

tés, colorée de jaune orangé; spire

pointue, composée de sept tours sub-

convexes, chargés sur toute leur surface

de plis longitudinaux nombreux, entre-

croisés de stries fines, transverses et rap-

prochées. Les stries du dernier tour un

peu plus fortement prononcées vers la

base. Les sutures ornées, prés du bord,

d'une rangée de petites granulations

séparées par un sillon transverse.

Ouverture blanchátre, ovale, étroite,

rétrécie á la base; levre droite dentelée”.

Il convient d'ajouter que la taille de

la coquille varie de 7 a 9 mm, que la

couleur de la téléoconque varie du beige

orangé a l'orange soutenu et que la pro-

toconque vitreuse a deux tours est tou-

jours rosée a rose vif.

Animal orange vif moucheté de

blanc cru, pied plus étroit que dans les

autres especes. Opercule jaune moyen a

corne pále.

Distribution: Espece uniquement

connue par des populations peu abon-

dantes distribuées autour de la Péninsule

du Cap Vert, sur fonds durs de 0 a 40 m.

Remarques: Buccinum aurantium a été

décrit par Lamarck en 1822 sans illustra-

tion et sans localité, mais en référence

aux figures 1188 et 1189 de Martini, 1780.

Ces figures sont tres petites et confuses

et ne permettent pas l'identification. KIE-

NER (1834) redécrit Buccinum aurantium

avec deux figures explicites (Fig. 5), sur

la base principalement du matériel de

Lamarck. Un lot de 14 coquilles homo-

genes (Fig. 18) de la collection Lamarck-

Massena-Delessert (MHNG) correspon-

dant a la description et aux figures de

Kiener et compatibles avec la description

de Lamarck, doit étre considéré comme

un lot de syntypes probables de B. auran-

tium, méme si ce lot pourrait comporter

quelques sujets ajoutés ultérieurement

au matériel original de Lamarck. Les co-

quilles de la collection Duclos (Fig. 19)

identifiées et figurées par celui-ci comme

C. aurantia et la description originale du

taxon Columbella cancellata Gaskoin, 1851

correspondent aussi a notre espece séné-

galaise.

Genre Mitrella Risso, 1826

Espece type par désignation subséquente (Cox, 1927): Mitrella flaminea Risso, 1826 [= Mitrella

scripta (Linné, 1758)]

Mitrella denticulata (Duclos, 1840) (Figs. 4, 14-17)

Buccinum 2. Le Jol: Adanson, 1757 (pré-Linnéen, non-disponible).

Colombella denticulata Duclos, 1840.

Columbella triangulifera V. Maltzan, 1884.

Matériel type: 9 syntypes de C. denticulata Duclos aa MNHN: l'un des mieux préservés d'entre eux

(6.8 mm) est désigné ici comme lectotype (Fig. 14), les paralectotypes mesurent 6 a 7 mm. 2 syn-

types de C. triangulifera V. Maltzan (5.8 et 5.9 mm) au ZMB, Moll. 37058 (Fig. 15).

Autre matériel examiné: CJP: Sta. 1: 1 spm, 2 coqg; Sta. 5: 1 spm, Sta. 6: 6 spm, 7 coq; Sta. 7: 3 spm,

2 coq, Sta. 8: 7 coq; Sta. 9: 75 spm, 11 coq; Sta. 23: 6 spm; Sta. 24: 1 cog; Sta. 25: 9 spm. CFB: Sta. 2: 2

spm; Sta. 3: 3 spm, 6 cog; Sta. 6: 57 spm; Sta. 7: 200 spm; Sta. 9: 81 spm, 17 cog; Sta. 18: 23 spm, 1

coq; Sta. 22: 1 coq.

Localité type: Inconnue. La Péninsule du Cap Vert, Centre Sénégal, est désignée ici comme loca-

lité type de C. denticulata Duclos.

Description: Coquille de petite taille, 6

a 7.5 mm, ventrue, dernier tour représen-

tant environ 65% de la hauteur totale de

la coquille, tours légerement arrondis vers

106

le bas, suture nette; sculpture ne compor-

tant que 7 a 8 sillons sur la partie inférieure

du dernier tour; ouverture étroite, labre

épaissi extérieurement portant 5 a 7 dents,

PELORCE ET BOYER: Columbellidae dans Pinfralittoral de la Péninsule du Cap Vert

cal columellaire transparent, ne dépassant

pas l'ouverture, portant quelques plis tres

peu marqués; décoration variant du

marron au violet intense en passant par le

rose clair, présence de points blancs orga-

nisé sous forme de bandes au milieu du

dernier tour et sur la partie inférieure au

niveau des sillons, groupés en masses tri-

angulaires sous la suture, conférant a celle-

ci une allure festonnée; protoconque a 1.5

tour, brune avec sommet blanc.

Animal blanchátre presque entiere-

ment recouvert de nappes marron clair

a brun noirátre, avec quelques lacunes

sur l'avant, l'arriere ou les cótés du pied,

sole supérieure blanchátre avec zone

mauve pále sur le bord latéral avant,

dessous de la sole blanc, tour de l'ceil et

pointe des tentacules blancs, zones

blanches mouchetées de blanc cru, sauf

autour de l'ceil. Opercule ovale arrondi

a étiré, jaune soutenu avec longue zone

d'attachement brune formant un crois-

sant périphérique.

Distribution: Cette espece est repré-

sentée par des populations localement

abondantes autour de la Péninsule du

Cap Vert, sur fonds durs entre O et 40 m.

Remarques: FISCHER-PIETTE (1942) iden-

tifie C. denticulata comme espece valide

du Sénégal, place C. triangulifera Maltzan,

1884 en synonymie et attribue l'espece au

«sous-genre» Mitrella. La variabilité réduite

de l'espece est vérifiée aussi bien dans le

matériel type (Fig. 14) et dans les figures

types (Fig. 3) que chez les syntypes de C.

triangulifera (Fig. 15) et dans l'abondant

matériel collecté a Dakar (Figs. 16, 17).

Mitrella melvilli (Knudsen, 1956) comb. nouv. (Fig. 13)

Pyrene melvilli Knudsen, 1956 Marine Prosobranchs of Tropical West Africa (Stenoglossa). Atlan-

tide Report, 4: 33-34.

Matériel type: Holotype ZMK (8.9 x 3.4 mm).

Autre matériel examiné: CJP: Sta. 15: 1 coq., CFB: Nord Sénégal: 1 coq.

Localité type: Guinée Francaise.

Description originale: «The shell has

about 9 */4 whorls, a rather high and

pointed apex. Apart from 10 spiral ridges

on the basal part of the body whorl and

closely set and very faint growth lines,

the shell is entitely devoid of sculpture,

but is somewhat shining. There does not

appear to be any demarcation between

the protoconch and the adult shell. The

suture is rather deeply incised. Aperture

small. Columellar side with a reflected

callus and 5 indistinct denticles. Outer lip

straight and almost parallel to the colu-

mella, with a somewhat thickened varix

and a distinct but slightly developed

sinus near the suture. Interior with 6 dis-

tinct denticles. Siphon broad and short.

Colouration: upper part of the shell

brown with indistinct brown and whitish

spots near the upper suture».

Animal et opercule non observés.

Distribution: Connue du Nord

Sénégal (CFB) jusqu'au Sud Angola

(comm. pers. d'E. Rolán).

Remarques: Cette espece est replacée

dans le genre Mitrella, car elle ne

possede pas la coquille biconique ni le

fort cordon subsutural qui caractérisent

le genre Pyrene, et montre en revanche

toutes les caractéristiques morpholo-

giques de M. scripta (Linné, 1758),

espece type de Mitrella.

Une seule coquille correctement

conservée de cette espece (Fig. 13), sans

protoconque, a été trouvée dans nos

récoltes, provenant d'un niveau profond

(35 m). Une coquille subfossile attri-

buable a M. melvilli a été draguée au

large de Lompoul (150 m), Nord

senégal parten 1991 (CEbB) Ce

dernier signalement constitue la localité

la plus septentrionale connue pour l'es-

pece. L'holotype provenant lui-méme de

32 m, il semble s'agir d'une espece a dis-

tribution bathymétrique assez profonde,

tout au moins dans la région d'influence

guinéenne (infralittoral inférieur et cir-

calittoral supérieur).

107

Iberus, 23 (2), 2005

Mitrella ocellata (Gmelin, 1791) (Fig. 12)

Buccinum 1. Le Barnet: Adanson, 1757 (pré-Linnéen, non-disponible).

Voluta ocellata Gmelin, 1791.

Buccinum cribrarium Lamarck, 1822.

Columbella guttata Sowerby, 1832.

Buccinum canariense Orbigny, 1839.

Matériel type: Non examiné.

Autre matériel examiné: CJP: Sta. 2: 6 spm, 2 coq; Sta. 7: 1 spm, 2 coq. CFB: Sta. 4: 3 cog; Sta. 6: 1

spm;, Sta. 7: 5 coq.

Localité type: Non définie par Gmelin. Définition subséquente: Nassau, New Providence Island,

Bahama Islands (RADWIN, 1978b).

Description: Coquille de taille

modérée, 11-13 mm, fusiforme, proto-

conque á environ 2 tours, lisse; spire

représentant les ?/5 de la hauteur totale

de la coquille, aigue quand elle est entiére

(la coquille est habituellement ététée),

profil des tours plat, suture profonde;

corps de la coquille cylindrique, ouver-

ture modérément large, levre externe

légerement épaissie, dentelée sur sa face

interne, columelle droite, lisse, canal

siphonal trés court, canal anal peu

marqué; absence de sculptures; décora-

tion de points blancs sur fond noirátre a

brun-tabac foncé. Certaines formes pos-

sedent une coquille entierement jaune-

blanc, d'autres une couleur de fond jaune

ou brun pále avec des points plus foncés;

protoconque blanche a deux tours.

[animal blanchátre décoré de lignes

croisillonées brun mordoré a rouge-brun

sur l'ensemble du pied et de la téte,

fondues en masse foncée sur le front et

fragmentées en taches irrégulieres sur

l'avant du pied et sur le siphon, dont la

partie distale porte un large anneau

marron clair. Tour de 1'ceil et extrémité des

tentacules blancs, base des tentacules brun-

rouge, partie centrale des tentacules et sole

brun léger. Extrémité du siphon faible-

ment moucheté de blanc cru. Opercule

losangique, brunátre subhyalin, portant

des stries transversales sur une longueur

et un petit éperon sur la longueur opposée.

Distribution: Zones tropicales et sub-

tropicales de 1'Atlantique, formes

jumelles dans l'Indo-Pacifique.

Remarques: M. ocellata n'est représen-

tée autour de la Péninsule du Cap Vert que

dans la zone littorale, sous blocs a tres

faible profondeur et généralement par des

spécimens isolés ou en nombre tres res-

treint. Les premiers tours sont le plus

souvent absents chez les sujets adultes.

Mitrella psilla (Duclos, 1846) comb. nouv. (Figs. 6, 7, 46-55)

Columbella psilla Duclos, 1846.

Colombella japix Duclos, 1850.

Matériel type: 1 syntype de C. psilla (4.8 mm) au MNHN, désigné ici comme lectotype (Fig. 46). 1

syntype de C. japix (5.2 mm) (Fig. 47) au MNHN.

Autre matériel examiné: CJP: Sta. 2: 3 spm;, Sta. 5: 9 spm, 7 coq; Sta. 6: 2 spm, 12 cog; Sta. 7: 1 spm; Sta.

8: 1spm, Sta. 9: 18 spm, 2 cog; Sta. 14: 9 spm, Sta. 15: 2 spm, 17 coq; Sta. 16: 2 spm, 1coq; Sta. 23: 33 spm,

1 coq; Sta. 25: 4 spm. CFB: Sta. 3: 5 cog; Sta. 4: 7 coq; Sta. 6: 2 spm, 1 cog; Sta. 7: 3 coq; Sta. 9: 1 spm, 4 cog;

Sta. 14: 1 spm, 6 cog; Sta. 16: 3 spm, Sta. 17: 13 spmy Sta. 18: 2 coq; Sta. 22: 21 spm; Sta. 29: 11 coq.

Localité type: Inconnue. La Péninsule du Cap Vert, Centre Sénégal, est désignée ici comme loca-

lité type de M. psilla Duclos.

Description: Coquille de petite taille

(de 4.8 á 6.2 mm), largeur égale ou lége-

rement inférieure á la moitié de la hau-

108

teur, dernier tour occupant environ 2/3

de la hauteur totale, suture bien mar-

quée, tours sensiblement convexes et ré-

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

Figures 39-45. Mitrella turbita. 39: lectotype, 8.8 mm (MNHN); 40: paralectotype, 9.4 mm

(MNHN); 41: syntype de C. phylina, 10 mm (MNHN); 42: syntype de C. broderipi var lutea,

8.75 mm (ZMB); 43-45: Petite Corniche, 8.3, 9.5 et 8 mm (CJP). Figures 46-55. Mitrella psilla.

46: lectotype, 4.8 mm (MNHN); 47: syntype de C. japíx, 5.2 mm (MNHN); 48: Faillis, 4.8 mm

(CJP); 49: Tiwa, 6.2 mm (CJP); 50: N'Gor, 5.9 mm (CJP); 51: Faillis, 5.0 mm (CJP); 52: Kunk

Diabar, 4.9 mm (CJP); 53, 54: Faillis, 5.2 mm et 5.3 mm (CJP); 55: Le Tacoma, 5.7 mm (CJP).

Figures 39-45. Mitrella turbita. 39: lectotype, 8.8 mm (MNHAN); 40: paralectotype, 9.4 mm

(MNHN); 41: syntype of. C. phylina, 10 mm (MNAN); 42: syntype of C. broderipi var lutea, 8.75

mm (ZMB); 43-45: Petite Corniche, 8.3, 9.5 and 8 mm (CJP). Figures 46-55. Mitrella psilla. 46:

lectotype, 4.8 mm (MNHN); 47: syntype of'C. japix, 5.2 mm (MNHN); 48: Faillis, 4.8 mm (CJP);

49: Tiwa, 6.2 mm (CJP); 50: N'Gor, 5.9 mm (CJP); 51: Faillis, 5.0 mm (CJP); 52: Kunk Diabar,

4.9 mm (CJP); 53, 54: Faillis, 5.2 mm and 5.3 mm (CJ]P); 55: Le Tacoma, 5.7 mm (CJP).

109

Iberus, 23 (2), 2005

gulierement étagés; protoconque á envi-

ron 1 tour, bulbeuse, lisse, blanche avec

quelquefois une bande de couleur mé-

diane; sculpture constituée de faibles

stries d'accroissement verticales visibles

sous un fort grossissement, ainsi que

d'une forte varice longitudinale posi-

tionnée sur le dernier tour, en retrait du

bord labial; la columelle porte une di-

zaine de stries longitudinales; ouverture

ovale, labre court et arqué, épaissi exté-

rieurement et portant de 5 a 7 dents, cal

columellaire transparent ne débordant

pas de l'ouverture, portant quelques plis

tres peu marqués; décoration tres va-

riable, généralement constituée de

taches rondes blanchátres sur fond brun

á marron plus ou moins foncé, parfois

des alignements de taches se fondent en

bandes blanches horizontales ou verti-

cales, certaines coquilles possedent une

couleur paille a marron clair uniforme

ou mouchetée de blanc (Figs. 48-55).

Animal blanchátre tacheté de noi-

rátre sur l'ensemble du pied, le dessus

de la sole et les bords latéraux sur le

dessous. Les taches noirátres peuvent

devenir mauve ou lie-de-vin dans les

zones périphériques ou inversement. Un

large anneau noirátre a mauve sur la

partie centrale des tentacules, qui sont

sensiblement spatulés a ce niveau, et un

étroit anneau noirátre a mauve vers l'ex-

trémité du siphon. Moucheté de blanc

cru a la pointe des tentacules, dans la

partie antérieure du siphon, sur l'extré-

mité avant du pied et de la sole ainsi

que sur la pointe arriere de la sole supé-

rieure. Opercule jaune hyalin tres léger,

centre grisátre flanqué d'un croissant

latéral brun noirátre clair á mi-distance

du bord de l'opercule.

Distribution: L'espece semble étre dis-

tribuée du nord mauritanien au Sénégal

et dans le sud angolais, sans solution de

continuité dans le Golfe de Guinée. II

n'est pas exclu que les populations simi-

laires d'Angola, dont les individus sont

généralement plus grands que ceux du

Sénégal, constituent une espece jumelle

de M. psilla, séparée de longue date des

populations sénégalaises.

Remarques: Mitrella psilla constitue

une espece commune dans la région de

Dakar, présente sur fonds durs dans la

plupart des stations et a toutes les pro-

fondeurs de l'infralittoral. Les propor-

tions et la décoration de la coquille

varient de maniere importante (Figs. 46-

55), mais plusieurs traits morpholo-

giques originaux et systématiquement

corrélés (les tours convexes, le labre

court et arqué et la varice située sur le

dos en retrait du labre) permettent l'at-

tribution spécifique avec certitude. Les

sujets assimilables a M. psilla de Mauri-

tanie et d'Angola présentent une homo-

généité beaucoup plus grande quant a la

morphologie et a la décoration de la

coquille.

Mitrella turbita (Duclos, 1840) comb. nouv. (Figs. 2, 3, 39-45)

Buccinum 4. Le Rac: Adanson, 1757 (pré-Linnéen, non-disponible).

Colombella turbita Duclos, 1840.

Colombella phylina Duclos, 1846.

Columbella broderipi var. lutea V. Maltzan, 1884.

Columbella rac Dautzenberg, 1891.

Matériel type: 3 syntypes de C. turbita Duclos au MNHN: le mieux conservé d'entre eux (8.8 mm)

est désigné ici comme lectotype (Fig. 39), les deux paralectotypes mesurent respectivement 9.4 mm

(Fig. 40) et 10 mm. 2 syntypes de C. phylina Duclos (9 et 10 mm) (Fig. 41) au MNHN. 3 syntypes de

C. broderipi var lutea V. Maltzan (2 adultes et 1 juvénile: 8.2 a 8.7 mm) au ZMN, Moll. 33327 (Fig.

42). 22 syntypes de C.rac Dautzenberg dans la Collection Adanson (MNHN). Les syntypes de C.

rac (IRSNB, Collection Dautzenberg, ex-Chevreux) sur lesquels Dautzenberg a fondé sa descrip-

tion n“ont pas été examinés.

Autre matériel examiné: MNHN (Collection Adanson): 22 coq accompagnées du label: « 2,483,

Buccin autre Rak... ». CJP: Sta. 2: 5 spm;, Sta. 5: 1 coq; Sta. 6: 8 spm, 17 coq; Sta. 7: 3 spm, 1 cog; Sta.

110

PELORCE ET BOYER: Columbellidae dans lP'infralittoral de la Péninsule du Cap Vert

Figures 56-60. Mitrella fombriata sp. nov. 56-59: holotype, 5.1 mm, Le Tacoma (MNHN); 60: Le

Tacoma, 5.8 mm (CJP). Figures 61-65. Mitrella inflata sp. nov. 61-64: holotype, 6.4 mm, Kunk

Diabar (MNHN); 65: paratype, Kunk Diabar, 6.4 mm (CJP).

Figures 56-60. Mitrella fimbriata sp. nov. 56-59: holotype, 5.1 mm, Le Tacoma (MNHN); 60: Le

Tacoma, 5.8 mm (C]P). Figures 61-65. Mitrella inflata sp. nov. 61-64: holotype, 6.4 mm, Kunk

Diabar (MNHN); 65: paratype, Kunk Diabar, 6.4 mm (CJP).

8: 13 spm, 29 cog; Sta. 14: 46 spm, 4 cog; Sta. 15: 2 spm; Sta. 16: 2 spm; Sta. 25: 3 coq. CFB: Sta. 3: 2

cog; Sta. 4: 7 cog; Sta. 6: 2 spm, 43 coqg; Sta. 7: 29 spm; Sta. 9: 2 spm, 9 coq; Sta. 13: 2 spm; Sta. 14: 390

spm; Sta. 18: 75 spm, 4 cog; Sta. 19: 1 spm, Sta. 22: 57 spm, Sta. 26: 6 coq.

Localité type: Inconnue. La Péninsule du Cap Vert, Centre Sénégal, est désignée ici comme loca-

lité type de M. turbita Duclos.

Description: Coquille de taille occupant environ ?/3 de la hauteur totale,

moyenne, 8 a 10 mm, largeur égale a suture bien marquée, tours pratiquement

environ 42 % de la hauteur, dernier tour plans et régulierement étagés; proto-

111

Iberus, 23 (2), 2005

conque présentant environ 1.5 tour, bul-

beuse, lisse, blanchátre avec le sommet

souvent violet; sculpture constituée de

fortes cótes axiales, espacées, générale-

ment sinueuses et centrées sur la zone

médiane du dernier tour; 8 stries trans-

verses á la base du dernier tour; ouverture

ovale, 4 a 5 dents labiales, sinus anal supé-

rieur assez marqué, cal columellaire

opaque bien délimité, ne dépassant pas

l'ouverture et portant 6 plis; décoration

variable, constituée de taches blanches

irrégulieres sur fond marron (Figs. 43-45).

Animal blanchátre a beige décoré de

flammules longitudinales brun-orange

léger a noirátre, souvent fragmentées en

- petites taches régulieres sur l'ensemble

du pied, parfois plus foncées vers la téte

et sur le siphon. Tentacules épais, portant

un anneau ou un groupe de lignes longi-

tudinales brun ou noirátre dans leur

partie intermédiaire, parfois mouchetés

de blanc cru a leur extrémité. Siphon

portant un anneau foncé assez étroit dans

sa partie distale et moucheté de blanc cru

sur toute sa moitié antérieure. Dessus de

la sole portant de larges plages noirátres,

dessous de la sole blanc. Opercule sub-

triangulaire, faiblement arrondi, jaune

moyen, avec zone d'attachement centrale

large et noirátre aux contours irréguliers

et avec une ligne noirátre longeant le

bord postérieur.

Distribution: Connue des Canaries

orientales jusqu'au Centre Sénégal

(HERNÁNDEZ ET BOYER, 2005).

Remarques: Malgré les caracteres pas-

sablement ambigus des figures type de

Ductos (1840, 1846) (Figs. 2, 3), le maté-

riel type de C. turbita (Figs. 39, 40) et celui

de C. phylina (Fig. 41) ne laissent aucun

doute quant a la prévalence de ces taxa

sur la Columbella rac de DAUTZENBERG

(1891), nom plus récent et toujours utilisé

depuis pour désigner l'espece. L'utilisa-

tion du taxon C. rac dans la littérature n/a

pas été assez fréquemment réitérée pour

autoriser une inversion de la précédence

(article 23.9.1 du Code de Nomenclature

Zoologique) et le nom plus ancien de C.

turbita Duclos, 1840 doit prévaloir pour

désigner l'espece. Le matériel type de

Dautzenberg, constitué par un lot confié

par Mr Chevreux et déposé a 'IRSNB, n/a

pu étre examiné, mais la description et la

figure type de Columbella rac (DAUTZEN-

BERG, 1891: 38-40, figs. 2 a-c) ne laissent

aucun doute sur la synonymie de ce taxon

avec C. turbita. Le lot de 22 coquilles éti-

queté « 2,483 Buccin autre Rak... » dans

la Collection Adanson et reconnu par

FIsCHER=+PIETTE (1942) est conspécifique

de C. turbita et confirme l'interprétation

faite par DAUTZENBERG (1891) du Rac

d'Adanson á partir « d'une figuration

médiocre et d'une diagnose trop courte ».

C. turbita, représentée sur fonds durs

á tous les niveaux de l'infralittoral,

apparait comme le plus commun des

Columbellidae de la Péninsule du Cap

Vert. Malgré la variabilité importante de

la morphologie, de la taille et de la déco-

ration de sa coquille (Figs. 39-45), l'es-

pece se distingue aisément des autres

especes de Columbellidae représentées

dans l'Ouest Africain.

Mitrella fimbriata sp. nov. (Figs. 56-60)

Matériel type: Holotype (5.1 x 2.0 mm) MNHN (Figs. 56-59), 17 paratypes déposés comme suit: 1

paratype (spm) MNHN, MHNG, MNCN, ZMK, ZMB, CER), 5 paratypes (1 spm, 2 coq et 2 juvé-

niles) CFB, 6 paratypes (1 spm, 3 coq et 2 juvéniles) CJP (Fig. 60), tous de la localité type.

Autre matériel examiné: CJP: Sta. 6: 5 spm, 4 cog; Sta. 7: 1 cog; Sta. 8: 9 spm, 2 coq; Sta. 9: 3 spm, 3

cog; Sta. 15: 9 coq; Sta. 16: 1 spm, Sta. 21: 2 coq; Sta. 23: 2 spm, 5 coq. CFB: Sta. 9: 5 spm, 4 coqg; Sta.

14: 1 spm, 4 cog; Sta. 26: 6 cog; Sta. 28: 1 spm; Sta. 29: 4 spm.

Localité type: Le Tacoma, 5-13 m, a l'est de lle de Gorée, Centre Sénégal.

Etymologie: Du latin fimbriatus (féminin: -1), orné d'une frange, en référence á la décoration sub-

suturale de la coquille.

Description: Coquille de petite taille (L

= 4.8 a 5.8 mm), largeur égale ou légere-

112

ment inférieure a 40 % de la hauteur,

dernier tour occupant environ ?/3 de la

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

Figures 66-69. Nassarina procera sp. nov. 66, 67: holotype, 4.3 mm, Les Madeleines, Dakar

(MNHN); 68, 69: paratype, environs de Dakar, 4.1 mm (CJP). Figures 70-75. Nassarina rolani sp.

nov. 70, 71: holotype, 4.8 mm, Grand Thiouriba, Dakar (MNHN); 72-75: paratypes, 4.2, 4.2,

4.5 et 4.3 mm, Grand Thiouriba, Dakar (CJP). Figure 76. Clathurella polignaci, holotype, 4 mm,

iles Bissagos (MNHN).

Figures 66-69. Nassarina procera sp. nov. 66, 67: holotype, 4.3 mm, Les Madeleines, Dakar

(MNAN); 68, 69: paratype, near Dakar, 4.1 mm (CJP). Figures 70-75. Nassarina rolani sp. nov. 70,

71: holotype, 4.8 mm, Grand Thiouriba, Dakar (MNHN); 72-75: paratypes, 4.2, 4.2, 45 and 4.3

mm, Grand Thiouriba, Dakar (CJ]P). Figure 76. Clathurella polignaci, holotype, 4 mm, iles Bissagos

(MNHN).

1848

Iberus, 23 (2), 2005

hauteur totale, suture incisée bien

marquée, tours légéerement convexes et

régulierement étagés; protoconque a

environ 0.75 tour, bulbeuse, lisse, blanche;

sculpture constituée de faibles stries d'ac-

croissement visibles sous fort grossisse-

ment, cordons spiraux régulierement

espacés, plus prononcés a la base du

dernier tour, moins marqués dans la zone

subsuturale et tres effacés vers la base des

tours de spire et vers le milieu du dernier

tour; ouverture ovale, canal anal tres lége-

rement marqué, canal siphonal court et

profond, sur les spécimens matures le

labre épaissi extérieurement porte 9 dents,

cal columellaire transparent ne dépassant

pas l'ouverture et portant 2 plis tres peu

marqués; décoration peu variable de

lignes axiales sinusoides marron sur fond

beige clair se rejoignant par 2 ou 3 sous la

suture ainsi que de part et d'autre d'une

lacune spirale au milieu du dernier tour

pour former un systeme de franges;

coquille translucide et fausse suture net-

tement visible par transparence.

Animal blanchátre a jaune-creme

décoré de taches vert-olive á brun foncé,

de densité et de forme variables, parfois

étroites et allongées. Opercule non-

observé.

Distribution: Connue uniquement de

la Péninsule du Cap Vert, Centre Sénégal.

Remarques: Mitrella fimbriata sp. nov.,

assez commune localement, est bien repré-

sentée entre 5 et 40 m dans la zone pros-

pectée. Les caracteres morphologiques et

la décoration de la coquille sont tout a fait

originaux pour 1'Afrique de 1'Ouest et

pour les eaux atlantiques. On attribue a

sa moindre densité et a sa distribution plus

irréguliere, ainsi qu'a son absence dans

les petits fonds de la zone littorale, le fait

qu'une espece aussi originale n'ait pas été

découverte et décrite préalablement d'un

secteur particulierement étudié de la pro-

vince Ouest Africaine.

Mitrella inflata sp. nov. (Figs. 61-65)

Matériel type: Holotype (6.4 x 2.7mm) MNHN (Figs. 61-64), 5 paratypes déposés comme suit: 1

MNCN, 1 CFB, 1 CER, 2 CJP (Fig. 65), tous de la localité type.

Autre matériel examiné: CJP: Sta. 9: 1 spm, Sta. 15: 8 coq; Sta. 21: 13 spm. CFB: Sta. 6: 2 spm, Sta.

14: 2 spm; Sta. 18: 5 spm; Sta. 19: 14 spm; Sta. 21: 16 spm, Sta. 22: 23 spm.

Localité type: Haut fond du Kunk Diabar, 30-45 m, au large de la Petite Cóte, Centre Sénégal.

Etymologie: Du latin inflatus (féminin: -a), enflé, gonflé, en référence a la forme dilatée de la coquille.

Description: Coquille de petite taille

(5.8 a 6.8 mm), largeur légerement infé-

rieure á la moitié de la hauteur, dernier

tour occupant environ 70 % de la

hauteur totale, suture incisée bien

marquée, tours légerement convexes et

régulierement étagés; la protoconque á

environ 1.5 tour, bulbeuse, lisse, blanche

au sommet violet; la sculpture est consti-

tuée de faibles stries d'accroissement

visibles sous fort grossissement et de

cordons spiraux régulierement espacés

sur l'ensemble de la téléoconque, partiel-

lement ou totalement effacés au milieu

du dernier tour et vers la base des tours

de spire; ouverture ovale, canal anal bien

marqué, séparé en deux par une dent

assez forte, canal siphonal court et

profond, labre épaissi extérieurement

portant 5 petites dents, cal columellaire

114

transparent ne dépassant pas l'ouverture

et portant 8 plis de tres petite taille;

décoration de lignes axiales sinusoides

marron sur fond blanchátre, se rejoi-

gnant par groupes de 4 a 7 sous la suture

ainsi que de part et d'autre d'une lacune

spirale vers le milieu du dernier tour,

pour former un systéme de franges;

coquille subtranslucide, fausse suture

modérément distincte par transparence.

Animal blanchátre décoré de petites

taches brun-rouge clair sur la téte et le

pied. Aréte dorsale, liste arriere du pied

et siphon tachés de brun foncé plus mat,

extrémité du siphon portant parfois un

anneau brun foncé, tentacules blanchátres

portant une táche brun-rouge clair dans

sa partie médiane, liseré marron clair sur

le bord du pied, dessus de la sole blanc

légerement moucheté de blanc cru vers

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

20 um

Figures 77, 78. Nassarina procera sp. nov. 77: radula dent latérale et centrale; 78: opercule, lon-

gueur réelle 0,8 mm. Figures 79-82. Nassarina rolani sp. nov. 79: radula dent latérale et centrale;

80: opercule, longueur réelle 0,6 mm; 81, 82: radula.

Figures 77, 78. Nassarina procera sp. nov. 77: lateral and central radular teeth; 78: opercule, real

lenght 0,8 mm. Figures 79-82. Nassarina rolani sp. nov. 79: latearal and central radular teeth; 80:

opercule, real lenght 0,6 mm; 81, 82: radula.

l'extrémité postérieure, dessous blanc.

Opercule subtriangulaire, jaunátre, parfois

finement moucheté de noir, zone d'atta-

chement noire, étirée sur l'axe médian

dans le sens longitudinal et superposée a

la liste foncée du metapodium.

Distribution: Connue uniquement de

la Péninsule du Cap Vert et au large de

la Petite Cóte, Centre Sénégal.

Remarques: M. inflata sp. nov. pré-

sente le méme modele de distribution

que M. fimbriata, avec laquelle elle

partage des systemes de microsculpture

et de décoration de la coquille tres sem-

blables. M. inflata se distingue de M. fim-

briata par une taille supérieure et une

silhouette plus renflée, une protoconque

plus élancée au sommet violet, une

téléoconque aux tours plus convexes et

une décoration constituée de lignes

axiales plus nombreuses et proportion-

nellement plus étroites. Ces 2 especes

peuvent étre considérées comme étroite-

ment apparentées et comme constituant

un groupe d'especes isolé dans le

contexte atlantique.

Genre Nassarina Dall, 1889

Espece type par désignation originale; Nassarina bushii Dall, 1889

Nassarina procera sp. nov. (Figs. 66-69, 77, 78)

Matériel type: Holotype (adulte: 4.3 x 1.4 mm) MNHN (Figs. 66, 67), 3 paratypes juvéniles (3.2 mm,

3.2 mm et 2.9 mm) de la localité type (CFB), 1 paratype adulte (4,1.mm) (Figs. 68, 69) des environs

de Dakar (CJP).

51S

Iberus, 23 (2), 2005

Localité type: Illes des Madeleines, 7-20 m, a l'ouest de Dakar, Centre Sénégal.

Etymologie: Du latin procerus (féminin: -a), allongé, élancé, svelte, par référence a la silhouette

étroite et effilée de la coquille.

Description: Coquille de petite taille

(4.1 a 4.3 mm), allongée, cylindrique,

largeur égale á environ *'/3 de la

hauteur, dernier tour occupant environ

60 % de la hauteur totale, suture incisée

bien marquée, tours convexes; la proto-

conque a environ 1.75 tour, pointue,

lisse, marron tres clair avec trois bandes

de couleur plus foncée; sculpture de la

téléoconque constituée d'environ 17

fortes cótes axiales et de forts cordons

spiraux d'épaisseur comparable, dont 5

'au-dessus de l'ouverture et 8 au-dessous

sur le dernier tour, donnant a la coquille

un aspect réticulé avec des nodules aux

intersections; ouverture rectangulaire,

canal anal bien marqué, canal siphonal

court et profond, labre externe épaissi

extérieurement et portant 5 petites

dents, cal columellaire transparent ne

dépassant pas l'ouverture, sans dent

visible; couleur uniforme marron avec

quelques variations d'intensité sur la

columelle et les cótes axiales.

Animal non-observé. Opercule corné

avec noyau subapical (Fig. 78), radula

composée sur chaque rang d'une dent

centrale rectangulaire et de 2 dents laté-

rales portant 2 cuspides effilées dans

leur partie distale et 1 cuspide massive

en forme de talon vers la base (Fig. 77).

Distribution: Connue uniquement

par le matériel type de la Péninsule du

Cap Vert, récolté sur fonds du:rs.

Remarques: Nassarina procera sp. nov.

apparaít, aux cótés de M. melvilli,

comme l'un des deux Columbellidae les

plus rares de ceux récoltés dans l'infra-

littoral de la Péninsule du Cap Vert. Il

paraít vraisemblable que N. procera se

trouve lá a la limite de sa distribution

géographique ou que sa distribution

bathymétrique principale se situe plutót

dans le domaine circalittoral.

Le placement de cette espece dans le

genre Nassarina est effectué sur la base

des diagnoses proposées par RADWIN

(1978a) et COSTA ET ABSALÁO (1998).

Nassarina rolani sp. nov. (Figs. 8, 70-75, 79-82)

Matériel type: Holotype (4.8 x 2.0 mm) (Figs. 70, 71) et 2 paratypes MNHN, 2 paratypes Z.M.B.,

2 paratypes MNCN, 4 paratypes CER, 9 paratypes CEB, 10 paratypes CJP, tous de la localité type.

Autre matériel examiné: CJP: Sta. 6: 5 spm, 1 cog; Sta. 9: 90 spm, 16 cog; Sta. 14: 2 spm, 8 coqg; Sta.

16: 16 spm, Sta. 20: 2spm, 3 coqg; Sta. 21: 39 spm, 25 coq; Sta. 22: 6 spm, 2 coq; Sta. 25: 1 spm, 1 coq.

CFB: Sta. 6: 10 spm, 10 coq; Sta. 9: 18 coq; Sta. 13: 2 spm;, Sta. 14: 2 spm, 2 cog; Sta. 21: 1 cog; Sta.

22: 2 spm; Sta. 26: 4 spm; Sta. 27: 1 spm.

Localité type: Grand Thiouriba, sud-ouest de Dakar, 40 m, Péninsule du Cap Vert, Centre Sénégal.

Etymologie: L'espece est dédiée au Dr. Emilio Rolán pour l'aide apportée tout au long de cette étude.

Description: Coquille de petite taille

(4.2 a 5.3 mm), allongée, largeur égale a

environ 40 % de la hauteur, dernier tour

occupant environ 63 % de la hauteur

totale, suture incisée bien marquée, tours

bien convexes et régulierement étagés; la

protoconque de 1.75 a 2 tours, pointue,

lisse, blanche; la sculpture de la téléo-

conque est constituée d'environ 11 fortes

cótes axiales qui s'interrompent sur le

dernier tour a la hauteur de l'ouverture,

et de forts cordons spiraux, 6 au-dessus

de l'ouverture et 10 á sa hauteur, les cótes

116

sont plus épaisses que les cordons, qui

tendent a chevaucher celles-ci; ouverture

rectangulaire, canal anal faiblement

marqué, canal siphonal court et profond,

labre épaissi extérieurement et portant 5

dents, cal columellaire ne dépassant pas

l'ouverture et portant 3 plis tres faibles;

couleur de fond variable depuis le blanc

pur jusqu'au marron-violet foncé, avec

bandes spirales plus foncées d'épaisseur

variable (Figs. 72-75).

Décoration de l'animal tres variable,

le pied et la téte jusqu'a la base des ten-

PELORCE ET BOYER: Columbellidae dans l'infralittoral de la Péninsule du Cap Vert

tacules étant généralement marbrés de

gris ou de noirátre sur fond blanc a

creme. Un anneau ou des marques longi-

tudinales gris peuvent étre présents sur

la partie intermédiaire des tentacules.

Chez les animaux sombres, le dessous de

la sole blanche est couramment mou-

cheté de noir, l'avant et l'arriere de la

sole supérieure étant le plus souvent

mouchetés de blanc cru. Parmi les autres

modeles de décoration rencontrés figu-

rent le fond uniforme blanc creme uni-

quement moucheté de blanc cru, le fond

blanc moucheté de petits points blanc

cru et de points et petites lignes ou cou-

lures noirs, accompagné d'un siphon gri-

sátre moucheté de noir, ainsi le fond

blanc orné d'une selle noire encerclant

animal a la hauteur de la nuque.

Opercule jaune moyen, corné, avec

noyau subapical (Fig. 80), radula compo-

sée sur chaque rang d'une dent centrale

carrée et de 2 dents latérales portant 3

cuspides massives régulierement distri-

buées et progressivement plus effilées

vers la partie distale, ainsi qu' une faible

saillie vers la base (Figs. 79, 81, 82).

Distribution: Connue uniquement

des fonds durs de la Péninsule du Cap

Vert, Centre Sénégal.

Remarques: L'extréme variabilité de

Nassarina rolaní sp. nov., rencontrée en

REMERCIEMENTS

Les auteurs remercient Haidar El-Ali

(Dakar), Patrice Petit de Voize

(Quimper) et Patrick Boyer (Saint

Martin de Brome) pour leur aide dans la

récolte de spécimens a Dakar, Emilio

Rolán (Vigo) pour ses dessins d'

animaux vivants ainsi que pour la com-

munication de ses notes et croquis de

terrain et pour l'extraction des radulas

de Nassarina, Andrew Wakefield (Buck-

BIBLIOGRAPHIE

ADANSON, M., 1757. Histoire naturelle du Séné-

gal. Coquillages. Avec la relation abrégée d'un

voyage fait en ce pays, pendant les années 1749-

53. Partie 2, coquillages: 1-275, 19 pls., 1 carte.

Bauche, Paris.

populations localement nombreuses

entre 10 et 40 m sur fonds durs, a donné

lieu a des contróles approfondis qui per-

mettent de confirmer qu'aucun “groupe

de forme” homogene ne peut étre dis-

tingué a l'intérieur de la série, ni en ce

qui concerne les proportions, la sculp-

ture et la décoration de la coquille ni en

ce qui concerne le chromatisme de l'ani-

mal ou la corrélation entre ces variables.

Clathurella polignaci Lamy, 1923 des

iles Bissagos (Fig. 8: figure originale et

Fig. 76: holotype MNHN) montre d'évi-

dentes affinités avec N. rolani, dont elle

se distingue principalement par une

taille plus petite (4 mm), une proto-

conque plus courte, une silhouette plus

pupoide avec des tours de spire moins

convexes et moins anguleux, des cótes

et des cordons plus nombreux et moins

épais.

Le replacement de C. polignaci dans le

genre Nassarina est proposé ici. Ce genre,

représenté par plusieurs espéeces dans

l'Atlantique occidental (RADWIN, 1978a;

COSTA ET ABSALAO, 1998), n'était re-

connu jusqu'a présent dans l'Est Atlan-

tique que par l'holotype de N. polignaci

de Guinée-Bissau (jusqu'alors attribué a

un autre genre) et par les 3 types de N.

rietae Segers et Swinnen, 2004 de La

Palma, Archipel des Canaries.

hurst Hill) pour son aide bibliogra-

phique, Ole Tendal du ZMK pour le prét

des types de Knudsen, Matthias Glau-

brecht du ZMB pour le prét des types de

von Maltzan, Yves Finet du MHNG

pour les photographies des syntypes

probables de B. aurantium, Philippe

Bouchet et Virginie Héros pour leur

accueil et la mise a disposition des types

du MNHN.

COSTA P. M. S. ET ABSALAO, R. S., 1998. Nassa-

rina thetys sp. nov. (Neogastropoda: Colum-

bellidae), a new species from the Brazilian

coast. Basteria, 62: 277-285.

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Iberus, 23 (2), 2005

DAUTZENBERG, P., 1891. Voyage de la goelette

Melita aux Canaries et au Sénégal, 1889-1890.

- Mollusques testacés. Mémoires de la Société

Zoologique de France, 4: 16-65, pl. 3.

DucLos, P. L., 1840. Coquilles univalves marines

a l'état vivant et fossile, publiées par monogra-

phies. Genre Colombella. Didot, Paris. Planches

1-13.

DucLos, P. L., 1846-1850. Genre Colombella. In

Chenu, J. C. (Ed): Illustrations Conchyliologí-

ques. Fortin, Masson, Paris. Planches 1-18:

année 1846; planches 19-27: année 1850.

FIsCHER-PIETTE, E., 1942. Notes critiques et des-

criptives sur les Columbellidae. 1. Sous-genre

Mitrella. Bulletin du Muséum, 2€ s., 14 (3): 223-

226.

FISCHER-PIETTE, E. Er NICKLES, M., 1946. Mo-

llusques nouveaux ou peu comnus des cótes

de l'Afrique occidentale. Journal de Conchy-

liologie, 87: 43-81, pl. 1.

GASKOIN, J. S., 1851. Descriptions of twenty

species of Columbellae and one species of Cy-

praea. Proceedings of the Zoological Society of

London, 19: 2-14.

GMELIN, J. F., 1791. Carolí Linnaei systema natu-

rae per regna tria naturae... Editio decima ter-

tia, aucta, reformata, Vermes Testacea. Leip-

zig, 1 (6): 3021-3910.

HERNÁNDEZ, J. ET BOYER, F., 2005. Notes about

the columbellid fauna from the infralittoral

and circalittoral levels of the Canary Islands.

Iberus, 23 (2): 69-93.

KIENER, L. C., 1834-1841. Iconographie des Co-

quilles Vivantes. Famille des Purpuriferes.

Deuxieme partie. Paris. Pp. 1-112, pls. 1-31.

KNUDSEN, J., 1956. Marine Prosobranchs of

Tropical West Africa (Stenoglossa). Atlantide

Report, 4: 7-110, pls. 1-4.

LAMARCXK, J. B. P. A. de M. de, 1822. Histoire Na-

turelle des Animaux sans Vertebres. Tome Sep-

tieme. Paris. Pp. 1-711.

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Lamy, E., 1923. Campagne du Sylvana, Mis-

sion du Comte Jean de Polignac et de Mr

Louis Gain. Mollusques Testacés. Extrait des

Comptes rendus du Congres des Sociétés savan-

tes en 1922. Sciences: 1-16.

LINNE, C. VON, 1758. Systema naturae per regna

tria naturae... Editio decima, reformata. 1,

Regnum animale. Stockholm. Pp. 1-824.

MC MILLAN, N. F., 1985. Frederick Price Marrat,

“Conchologist, etc* with a list of his type and fig-

ured specimens in Merseyside County Museums

and bibliography of his publications. Mersey-

side County Museums, Liverpool. Pp. 1-33.

MALTZAN, H. F. VON, 1884. Diagnosen neuer Se-

negambischer Gastropoden. Nachrichtsblatt

der deutschen Malakozoologischen Gesellschaft,

16 (5): 65-73.

MARRATr, F. P., 1877. A list of West African

shells, including three new Pleurotomae and

one new Columbella. Quarterly Journal of Con-

chology, 1: 237-244.

MOOLENBEEK, R. ET HOENSELAAR, H. J., 1991. On

the identity of “Columbella rustica” from West

Africa and the Macaronesian Islands. Bulle-

tin Zoologisch Museum, 13 (6): 65-70.

NICKLES, M., 1950. Mollusques Testacés Marins de

la Cóte Occidentale d'Afrique. P. Lechevalier,

Paris. Pp. 1-269.

PACE, S., 1902. Contributions to the study of the

Columbellidae, n*1. Proceedings of the Mala-

cological Society of London, 5 (1-2): 1-154.

RADWIN, G. E., 1978a. The Family Columbelli-

dae in the Western Atlantic. Part lla. The Py-

reninae. The Veliger, 20 (2): 119-133.

RADWIN, G. E., 1978b. The Family Columbelli-

dae in the Western Atlantic. Part Ilb. The Py-

reninae (Continued). The Veliger, 20 (4): 328-

344.

O Sociedad Española de Malacología

Iberus, 23 (2): 119-156, 2005

Columbellidae (Gastropoda, Neogastropoda) of the gulf of

Guinea with the description of eight new species

Columbellidae (Gastropoda, Neogastropoda) del Golfo de Guinea

con la descripción de ocho especies nuevas

Emilio ROLÁN*

Recibido el 1-XI-2204. Aceptado el 11-X-2005

ABSTRACT

The species of the family Columbellidae found in West Africa are studied. Five genera are

represented: Columbella (1 species), Anachis (1), Mitrella (11), Cotonopsis (1) and Strom-

bina (1). Of the 15 species mentioned, 8 are described as new. The genus Strombina is

employed by first time for a West African species.

RESUMEN

Se estudian las especies de la familia Columbellidae encontradas en África occidental.

Cinco géneros están representados en el área de estudio: Columbella (1 especie),

Anachis (1), Mitrella (11), Cotonopsis (1) y Strombina (1). De las 15 especies men-

cionadas 8 se describen como nuevas para la ciencia. El género Strombina es empleado

por vez primera para una especie oeste africana.

KEY WORDS: Columbellidae, Columbella, Anachis, Mitrella, Cotonopsis, Strombina, West Africa, Guinean

Gulf, new species.

PALABRAS CLAVE: Columbellidae, Columbella, Anachis, Mitrella, Cotonopsis, Strombina, África occidental,

golfo de Guinea, especies nuevas.

INTRODUCTION

The Columbellidae of the West Africa

have been seldom studied in recent

years. For most of the West African

species, we can only find some descrip-

tions and records in species lists, in-

cluded in older works such those of Duc-

LOS (1835-40), MENKE (1853), MALTZAN

(1884), MARTENS (1904), DAUTZENBERG

AND FISCHER (1906), DAUTZENBERG (1910,

1927), FISCHER-PIETTE (1942a, b), NICKLES

(1950) and KNUDSEN (1956).

More recently, isolated records have

been mentioned for Sáo Tomé, in FER-

NANDES AND ROLÁN (1993), and Angola,

in GOFAS, PINTO AFONSO AND BRANDAÁO

(1985), with some new records in ROLÁN

AND RYALL (1999a, b) and ROLAN AND

TRIGO (2000); also for Gabon, in

BERNARD (1984).

The Mediterranean species of the

family have been revised in some recent

papers: SCHIRO (1979), SABELLIAND SPADA

(1981), LUQUE (1984), VAN AARTSEN,

MENKHORST AND GITTENBERGER (1984),

MIFSUD (2000), CHIARELLI, MICALI AND

- QUADRI (2003), GIANNUZZI-SAVELLI,

PUSATERI, PALMERI AND EBREO (2003). The

Canary Islands species are treated in

NORDSIECK AND GARCÍA-TALAVERA (1979)

and in HERNÁNDEZ AND BOYER (2005) and

* Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela, Spain.

Yv9

Iberus, 23 (2), 2005

those from the Cape Verde Islands in the

publications of BURNAY AND MONTEIRO

(1977), COSEL (1982a, b, c), ROLÁN AND

LUQUE (2002) and ROLÁN (2002, 2004). The

Senegal species are revised by PELORCE

AND BOYER (2005).

Columbellid classification is still

problematic because of insufficient

information for most species, lack of dis-

crete characters useful for distinguish-

ing species groups, conchological and

radular variation within the family and

the geographically restricted basis of the

most frequently used columbellid classi-

fications (PACE, 1902 and DEMAIN-

TENON, 1999). This makes the generic

placement of species difficult, and thus

we tentatively use here Anachis (in the

sense Of RADWIN, 1977a, b, supported by

the results of DEMAINTENON, 1999) and

Mitrella (a polyphyletic genus according

to DEMAINTENON, 1999), due to the

resemblance of each of the new species

with the type species of both genera.

In this paper, which complements

the study of Dakar species by Pelorce

and Boyer (this volume), we consider the

Gulf of Guinea in a broad sense from

south of Dakar to the south of Angola.

MATERIAL AND METHODS

Most the material studied is in the

author's collection, constituted during

many expeditions to West Africa and by

diving, dredging, and sorting of sedi-

ment. Besides, material was loaned by

the MNHN, some types by ZMUC, and

other material was also studied from the

RESULTS

private collections mentioned in

Acknowledgements.

In order to standardize the criteria

for character descriptions we follow

DEMAINTENON (1999) for descriptions of

operculum and radula.

The radular terminology is based on

BANDEL (1984).

Abbreviations

AMNH American Museum of Natural

History, New York.

BMNH The Natural History Museum,

London.

MNCN Museo Nacional de Ciencias

Naturales, Madrid.

MNHN Muséum National d'Histoire

Naturelle, Paris.

SMF Senckenberg Museum, Frankfurt.

ZMUC Zoologisk Museum, Kobenhavn.

ZSM Zoologische Staatsammlung

Muenchen, Munich.

CCS collection of C. Schroenherr,

Luanda.

CER collection of E. Rolán, Vigo.

CFB collection of Franck Boyer, Sevran.

CJH collection of José María Hernández,

Gran Canaria.

CJP collection of Jacques Pelorce, Le

Grau du Roi.

CJT collection of Juan Trigo, A Coruña.

CHD collection of Juan Horro-Ana

Delgado, Vigo.

CPH collection of Jean Paul Hatten-

berger, France.

sp: live collected specimen.

s: empty shell.

]: juvenile shell.

f: fragment of shell.

Genus Columbella Lamarck, 1799

Type species: Voluta mercatoria Linné, 1758. Recent, Caribbean. Designated by Lamarck, Mém.

Soc. Hist. Nat. Paris: 70.

Columbella adansoni Menke, 1853 (Figs. 1-14, 149, 160)

Columbella adansoni Menke, 1853. Zeitschr. Malakozool., 10 (5-6): 74-75. [Type locality: Sáo Vicente

Island, Cape Verde archipelago].

Columbella rufa Menke, 1853. Zeitschr. Malakozool., 10 (5-6): 75.

120

ROLÁN: Columbellidae of the gulf of Guinea

Columbella rustica auct. non (Linnaeus, 1758).

Columbella spongiarium Duclos sensu ROCHEBRUNE (1881a).

Columbella rustica striata (Duclos) sensu PÉREZ SÁNCHEZ AND MORENO BATET (1991).

Type material: Lectotype, designated by MOOLENBEEK AND HOENSELAAR (1991), in SMF.

Other material examined: Many shells and specimens from Madeira; Canaries; Azores; Cape Verde

Islands; Ghana; Sáo Tomé; Angola.

Description: See NORDSIECK AND

GARCÍA-TALAVERA (1979, as Columbella

rustica striata). Shell (Figs. 1-11) solid,

with a large and wide last whorl, and a

conical pointed spire with 4-6 whorls.

Protoconch (Figs. 12-14) of several

whorls. Colour of teleoconch very variable,

pink, orange, brown, usually with blotches

of several colours frequently with small

oval light spots disposed spirally. Aperture

elongate, straightened at the middle by

an enlargement of the external lip.

Dimensions: Up to 21 mm, many

populations composed by smaller speci-

mens of about 8-12 mm only.

Soft parts pigmented but variable

according to shell colour.

Radula (Fig. 149) with a central tooth

four times as wide as long and with

very acute posterior corners. Lateral

teeth about three times as long as wide,

with twisted base; cutting edge of later-

als with the basal cusp low and long, the

central cusp wide and not sharpened,

and the apical cusp acute and wide.

Operculum (Fig. 160) light brown,

ovoid-elongate, with an ovoid mark of

insertion with a small prominence in the

middle.

Distribution: The species is known

from the Macaronesian archipelagos

and the mainland West African coast

from the Gulf of Guinea to Angola

(ROLÁN AND RYALL, 1999a).

Remarks: In the Gulf of Guinea, the

genus Columbella is represented by a

single species, whose shells, of medium

size, and abundant in shallow water

areas. This species was confused until

recently with C. rustica (Linné, 1758), so

references to C. rustica for the area

(NICKLES, 1950 or BERNARD, 1984,

among others) must be considered to

belong to C. adansoni Menke, 1853.

MOOLENBEEK AND HOENSELAAR

(1992) established the differences be-

tween both species, based on protoconch

and radula. These authors also explained

the reasons for their distribution, on the

basis of oceanic currents and interspe-

cific competition. They concluded that

the first, Columbella adansoni, is present in

the Macaronesian archipelagos and the

second, Columbella rustica, in the

Mediterranean Sea and along the north-

ern coast of West Africa. OLIVERIO (1995)

confirmed these differences on allozyme

based studies. ROLÁN AND RYALL (1999a)

established the geographic range of dis-

tribution of both species along the

African coast. In the present work the

radula (Fig. 148) and operculum (Fig.

159) of Columbella rustica are illustrated

to point out the differences.

The reasons for the confusion

between both species are due to the great

intraspecific variability which is more

important than the differential characters

between the two species. Therefore, with

the simple observation of the shells

without protoconch is impossible to

know to which taxon correspond and

from where they come from. The proto-

conch of C. rustica is paucispiral (Figs.

15-19). The planktotrophic multispiral

protoconch of C. adansoni supposes a

pelagic larval development making fea-

sible a distribution along archipelagos

and main coast. Columbella adansoni has a

larger larval dispersal ability ranging

from from the Azores to South Angola, a

geographical range of distribution

greater than that of C. rustica.

C. adanson1 is very variable in differ-

ent habitats. One form has shells with a

very prominent aperture, while another

has very small one; the colour can be

uniform or spotted. Sometimes the

shells are minute, in other places double

the size. In the Cape Verde archipelago,

specimens of different size often live

together.

121

Iberus, 23 (2), 2005

Figures 1-11. Columbella adansoni. 1-3: shells, 20.7, 19.1, 19.1 mm, Lanzarote, Canary ls.; 4-6:

shells, 14.6, 14.5, 13.4, mm, Mordeira, Sal, Cape Verde Islands; 7, 8: shells, 11.7, 10.0 mm, Sal

Rei, Boavista, Cape Verde Islands; 9: shell, 10 mm, Praia Amelia, Angola; 10, 11: shells, 14.5, 14.4

mm, Luanda, Angola (all from CER).

Figuras 1-11. Columbella adansoni. 1-3: conchas, 20,7, 19,1, 19,1 mm, Lanzarote, Islas Canarias;

4-6: conchas, 14,6, 14,5, 13,4, mm, Mordeira, Sal, Islas de Cabo Verde; 7, 8: conchas, 11,7, 10,0

mm, Sal Rei, Boavista, Islas de Cabo Verde; 9: concha, 10 mm, Praia Amelia, Angola; 10, 11: conchas,

14,5, 144 mm, Luanda, Angola (todas de la CER).

122

ROLÁN: Columbellidae of the gulf of Guinea

500 um

500 um 500 um

Figures 12-14. Protoconchs of C. adansoni, Cape Verde Islands. Figures 15-19. Protoconchs of C.

rustica. 15, 16: UEtoile, Nouadhibou, Mauritania; 17-19: Cullera, Valencia, Spain (all from CER).

Figuras 12-14. Protoconchas de C. adansoni, /slas de Cabo Verde. Figuras 15-19. Protoconchas de C.

rustica. 15, 16: L'Etoile, Nouadhibou, Mauritania; 17-19: Cullera, Valencia, España (todas de la

CER).

123

Iberus, 23 (2), 2005

Genus Anachis H. Adams and A. Adams, 1853

Type species: Columbella scalarina G. B. Sowerby II, 1832, from Panama (Chiriquí), by subsequent

designation (TATE, 1869).

Diagnosis: RADWIN (1977a, p. 120).

Anachis ryalli spec. nov. (Figs. 20-27)

Type material: Holotype (Fig. 20) in MNHN. Paratypes: AMNH (1, Fig. 21), MNCN (1, 15.05/46628,

Fig. 22), BMNH (1, Fig. 23), ZSM (1, Fig. 24), CJP (3), CJH (3), CFB (6), CER (5), CPR (137), all from

the type locality.

Type locality: Off Sekondi, Ghana, trawled around 40 m.

Etymology: The species is named after Peter Ryall, malacologist who lived for many years in Ghana

and collected the type material.

Description: Shell solid (Figs. 20-24),

up to 9.3, mm in. length, broadly

fusiform with a moderately high spire.

Protoconch (Figs. 26, 27) of 1 */4

whorls, sharply pointed, and about 500

um in maximum diameter, smooth and

usually whitish in colour but cream on

the upper part.

Teleoconch of 4-4 */2 hardly convex

spiral whorls, with distinct suture and

small subsutural shelf. Sculpture

formed by axial ribs, well defined sub-

suturally, between 18 and 22 on the last

whorl. Many spiral threads visible only

in the interspaces of the ribs, but more

evident near the base, where the axial

ribs disappear.

Aperture narrow and axially elon-

gate, usually white inside. Columella

curved, S-shaped, with 5-6 small denti-

cles in the lower part, the uppermost

being the most prominent. Inner part of

external lip with about 7-8 denticles, the

second one larger than the others.

Shell colour uniformly cream or

whitish. Periostracum thin, smooth and

transparent.

Soft parts, operculum and radula

unknown.

Distribution: Only known from the

type material, from Ghana.

Remarks: A. ryalli most resembles the

Senegalese species A. aurantia (Lamarck,

1822) (= A. cancellata Gaskoin, 1851) but

the latter is larger, with a wider and darker

shell. Also it may be differentiated from

A. freytagi Maltzan, 1884 [= Columbella

(Anachis) bubakensis Lamy, 1923], from

Senegal and Bissagos Islands, because its

shell is smaller and narrower, uniformly

subhyaline, light tan colour or brownish,

with a narrow aperture and a contracted

base. Even samples of A. freytagí with

lighter colour and almost erased colour

pattern, retain the typical colour in the first

whorls and have a more rectilinear profile.

Anachis cuspidata Marrat, 1877 (= A.

emergens Fischer and Nickles, 1946),

from Senegal, is wider and has a more

colourful shell.

Anachis turbita (Duclos, 1840) (= rac

Dautzenberg, 1891), from Senegal and

Canary Islands, is more colourful, has a

wider shell and its protoconch is not so

pointed.

Anachis ryalli may be differentiated

from A. valledori Rolán and Luque, 1999,

from the Cape Verde Islands, because

the latter species has a smaller and nar-

rower shell, is uniformly light in colour

without brown axial lines, has a narrow

aperture, and the protoconch is evi-

dently more elevated and whitish.

Anachis avaroides Nordsieck, 1975,

from the Canary and Selvagens Islands,

is wider, has constant colouration of

axial bands and ocelli, and lacks spiral

sculpture, except at the base.

Genus Mitrella Risso, 1826

Type species: Murex scriptus Linnaeus, 1758 (= Mitrella flaminea Risso, 1826), by subsequent des-

ignation of MÓRCH (1859).

124

ROLÁN: Columbellidae of the gulf of Guinea

Figures 20-27. Anachis ryalli spec. nov. 20: holotype, 6.2 mm; 21-24: paratypes, 6.5 (AMNH), 6.0

(MNCN), 6.4 (BMNH) and 6.7 mm (ZSM), Sekondi, Ghana, 40 m. 25: detail of the aperture;

26, 27: detail of the protoconch of paratypes of Figures 21 and 24.

Figuras 20-27. Anachis ryalli spec. nov. 20: holotipo, 6,2 mm; 21-24: paratipos, 6,5 (AMNH), 6,0

(MNCN), 6,4 (BMNA) y 6,7 mm (ZSM), Sekondi, Ghana, 40 m. 25: detalle de la abertura; 26, 27:

detalle de la protoconcha de los paratipos de las Figuras 21 y 24.

125

Iberus, 23 (2), 2005

Diagnosis: RADWIN (1977b, p. 337), but see remarks of the same author and DEMAINTENON (1999,

p. 267)

Mitrella pallaryi (Dautzenberg, 1927) (Figs. 28-34, 152, 163)

Columbella (Mitrella) vulpecula Pallary. 1900 ex Monterosato ms. Cog. Mar. litt. Dép. d'Oran. ].

Conchyl., 48(3): 279, pl. 6. fig. 8 (also var. minor Pallary and var. albida Pallary). [Type locality:

Oran, Algeria].

Pyrene pallaryi Dautzenberg, 1927. nom. nov. pro Columbella vulpecula Pallary, 1900 non C-. B.

Sowerby, 1844. Res. Camp. Sci. Albert [, 72: 89.

Type material: Not examined.

Other material examined: Atlantic: Spain: Galicia: 75 sp, 10 s, several j, from numerous localities

(CJT, CHD). Portugal: 4 sp, between Salema and Praia da Luz, Algarve, 37” 00' N, 08* 45" W, 70 m

(MNHN); 1 s, N/O Faial, 37” 01.3" N, 09% 05.7W, 135 m (MNHN). Mediterranean: Spain: Mar de

Alborán, Almería: 30 sp and s, 100-130 m (CER); 16 sp, Almería Bay (CER); Málaga: 4 sp, Marbella,

50 m (CER). Algeria: 5 s, near Orán, diving, 10 m (CER); 5 s, Orán, (MNHN). Malta: 6 s, 40 m (CER).

Morocco: 6 s, Tangiers, expiscis Lepidotrigla (MNHN); 3 sp, dredgings, 60-100 m (CER). Sahara: 25

sp, and s (CER). Azores archipelago: 2 c, 37” 03" N, 25” 09” W, N Santa Maria, 110 m (MNHN); 1 s,

38” 03' 40"N, 30% 55 W, 98 m (MNHN). Bay of Biscay: 2 s, 43? 46.51 N, 02* 00.58" W, 165 m (MNHN).

Madeira: N/O “Jean Charcot ZARCO, St. 21, 33” 00.7” N, 16” 25.5W, 220-290 m (MNHN). Canary

Islands: 2 s, La Palma, nets, 100-150 m (CER); 4 sp, La Palma, nets, 100-200 m (MNHN); 40 sp, 5 s,

Tasaente, La Palma, 150-200 m (CER); 1 s, Tenerife, 100-150 m (CER); 1 s, Gran Canaria (CER). Mau-

ritania: continental plateau, 18” 18” N, 16? 31” W, 134 m (MNHN); 1 s, “N'Diago” Stn. 306, 19* 6' N,

167 40” W, 93 m (MNAHN). Senegal: 7 s, Gorée (CES); 4 s, Cap Manuel, 50 m (MNAHN); 3 s, Cap de

Naza, 50 m (MNHN). Congo: 2 sp, Pointe Noire, Plage Mendame (CPH). Angola: 1 sp, 2 s, Ilha de

Luanda, 75-90 m (MNHN); 2 sp, Luanda, 20 m (CCS); 4 s, Luanda, dragados, 60 m (CER); 1 sp, 1

s, Mussulo, Luanda, 90-100 m (MNHN >); 10 s, Palmeirinhas (CER); 5 s, Saco Mar (CER).

Description: For shell description and

colour pattern of the head-foot, proto-

conch and operculum, see LUQUE (1986:

236, pl. 2, figs. 9, 15). Shell (Figs. 28-33)

elongate, solid, sharply pointed, smooth,

shiny and with a curved profile of the

whorls.

Protoconch (Fig. 34) of about three

whorls, also illustrated with SEM in

ROLÁN AND TRIGO (2000, figs. 6, 7).

Radula (Fig. 152, see also LUQUE, 1986,

pl. 3, fig. g, and ROLÁN AND TRIGO, 2000,

fig. 8), with a central tooth four times as

wide as long and with acute posterior

corners. Lateral teeth about four times as

long as wide; with twisted base; cutting

edge of laterals with the basal cusp short

and prominent, and both central and

apical cusps narrow and acute.

Operculum (Fig. 163) light brown,

ovoid, almost circular with an ovoid

mark of insertion, which is almost

divided in two by an elongate promi-

nence at the middle part.

Distribution: Described from the

Mediterranean (Oran) by PALLARY

126

(1900, as Columbella vulpecula), also cited

by SABELLI, GIANNUZZI-SAVELLI AND

BEDULLI (1990) and CHIARELLI, 2002; in

Europa by BOUCHET, LE RENARD AND

GOFAS (2001), DAUTZENBERG (1927)

recorded it from Azores, PASTEUR-

HUMBERT (1962) from Casablanca,

Morocco, NORDSIECK (1968) from Orán,

and SCHIRÓ (1979) from Alboran Sea.

MACEDO, MACEDO AND BORGES

(1999) also recorded it from Portugal. In

Spain it has been found by SIERRA, GAR-

CÍA AND LLORíÍS (1978) and SABELLI AND

SPADA (1981) from the Canaries, Gibral-

tar and other places in South Spain. CE-

CALUPO AND GIUSTI (1989) mention it for

Capraia, at 400-440 m. POPPE AND GOTO

(1991: 152, pl. 31, fig. 6) mention those

records already known.

ROLÁN AND RYALL (1999b) and

ROLÁN AND TRIGO (2000) recorded it for

Angola.

Mitrella pallaryi has a multispiral

protoconch indicating planktotrophy. Its

distribution area is large and includes

the whole Mediterranean, the Atlantic

ROLÁN: Columbellidae of the gulf of Guinea

Figures 28-34. Mitrella pallaryí. 28, 29: 17.0, 14.5 mm, Camariñas, Galicia, Spain (CER); 30:

14.7 mm, Oran, Algeria (CER); 31: 16.1 mm, La Palma, Canary 1. (CER); 32: 17.3 mm, Luanda,

Angola (CER); 33: 15.5 mm, A Guarda, Galicia, Spain; 34: protoconch, Luanda, Angola.

Figuras 28-34. Mitrella pallaryi. 28, 29: 17,0, 14,5 mm, Camariñas, Galicia, España (CER); 30:

14,7 mm, Orán, Algeria (CER); 31: 16,1 mm, La Palma, Islas Canarias (CER); 32: 17,3 mm,

Luanda, Angola (CER); 33: 15,5 mm, A Guarda, Galicia, España; 34: protoconcha, Luanda, Angola.

127

Iberus, 23 (2), 2005

0) 1 dl

a 1

Pei

Figures 35-45. Mitrella psilla. 35-44: shells, between 4.6 and 5.0 mm (CER). Baia de Etoile,

Mauritania, intertidal; 45: protoconch.

Figuras 35-45. Mitrella psilla. 35-44: conchas, entre 4,6 y 5,0 mm (CER). Bahía de l'Etoile, Maurita-

nia, intermareal; 45: protoconcha.

128

ROLÁN: Columbellidae of the gulf of Guinea

Figures 46-54. Mitrella psilla. 46-52: shells, 4.7, 5.3, 4.8, 4.8, 4.9, 4.1, 4.3 mm, Sacomar, Angola,

4-11 m (CER); 53: 4.6 mm, Baia das Pipas, Angola; 54: protoconch, Sacomar.

Figuras 46-54. Mitrella psilla. 46-52: conchas, 4,7, 5,3, 48, 48, 49, 4,1, 4,3 mm, Sacomar,

Angola, 4-11 m (CER); 53: 4.6 mm, Baía das Pipas, Angola; 54: protoconcha, Sacomar.

Iberus, 23 (2), 2005

up to north Spain, Archipelagos of

Azores and Canaries, and Angola. Curi-

ously there are no records for the west

African coast of the Gulf of Guinea.

Remarks: In the present work we

include photographs of specimens from

Galicia, north Spain, Camariñas, (Figs.

28, 29) and A Guarda, (Fig. 33), Oran,

Algeria (Fig. 30), Canary Islands (Figs.

31), Angola (Fig. 32), and the protoconch

of another shell from the Sahara (Fig.

34), some of these previously figured in

ROLÁN AND TRIGO (2000). All of them

show a similar form and even colour

pattern within some population vari-

ability. The wide distribution of M. pal-

laryi is explained if we consider the mul-

tispiral protoconch. The species is vari-

able in colour and pattern, but the main

characters are similar in shells from

distant localities.

The lack of collecting records in

some areas may be due to its peculiar

habitat, in deep and rocky bottoms, and

probably this species will be collected in

other localities from West Africa which

are still scarcely sampled. Anyway, up

to now it was not collected between

Senegal and Angola and may have a

bipolar distribution.

Mitrella psilla (Duclos, 1846) (Figs. 35-54, 132-135, 143, 156, 165, 169)

Colombella psilla Duclos, 1846. Hist. natur...Genre Colombelle, pl. 15, fig. 5-6. [no type locality].

Colombella japix Duclos, 1850. Hist. natur...Genre Colombelle, pl. 22, fig. 13-14. [no type locality].

Type material: Syntypes of C. psilla and C. japix in MNHN.

Other material examined: Mauritania: 30 s, 50 j, Baie de L'Etoile, intertidal (MNHN); 217 sp, 14 s,

38 j, Baie de l'Etoile, intertidal, in algae (CER); 16 s, Banc d” Arguin, in beach sediment (CER).

Senegal: 200 sp, 8 c, Region de Dakar (MNHN). Congo: 1 s, Pointe Indienne, 1 m (CPH). Angola: 3

sp, Lobito, 4 m (CER); 63 sp, 4 s, 8 j, Sacomar, 4-11 m (CER); 4 sp, Limagens, 5 m (CER); 1 s, Santi-

ago, 5 m (CER); 53 sp, 10 s, 10j, Praia Amelia, 5-11 m (CER); 1 sp, 15 c, 6j, Praia Amelia, Namibe

(MNHN); 3 j, Ponta de Noronha (MNHN); 5 j, Baia de Lucira (Bissonga), Namibe (MNHN); 2 sp,

1 e, 3j, Baia do Cesar, intertidal (CER); 8 s, 2j, Chapeu Armado, 5 m (CER); 10 s, 4 j, Baia das Pipas,

2-10 m (CER); 5 j, Sáo Nicolau, Namibe (MNHN); 1 s, Bentiava [formerly Sáo Nicolau], Namibe

(CER);3 sp, 2 c, Praia das Conchas, Namibe (MNHN).

Description: As there are different

populations, this description is general

for all of them. The differences are

pointed out below.

Shell (Figs. 35-44, 46-53) ovoid elon-

gate, solid, with a last whorl larger than

half the height.

Protoconch (Figs. 45, 54, 132, 133)

apparently smooth, with 1 */4 spiral

whorls, clearly demarcated from the

beginning of the teleoconch, diameter

about 600 um with nucleus 260 um.

Colour light with a dark brown spiral

band, except in the Mauritanian popula-

tion which is totally dark brown.

Microsculpture of protoconch (Figs. 134,

135) only visible with high magnifica-

tion and forming a rough surface.

Teleoconch with 4 whorls, shiny, su-

ture not impressed, with whorl profile

scarcely convex, without any sculpture.

130

Aperture relatively elongate with small

teeth on the columella and about 4-5 den-

ticles on the inner part of the outer lip.

Colour light brown to yellowish on

background, lighter near the suture and

with a band in the middle of the last

whorl; numerous, whitish or cream ocel-

late patterns superimposed on this back-

ground all over the surface. Aperture

elongate and narrow. External lip wide,

thicker externally.

Dimensions: Usually between 4.5 and

5.3 mm.

Soft parts in Mauritanian population

(Fig. 169) creamy-white, with numerous

circular grey ocelli and dark rings in the

middle of the tentacles and the extrem-

ity of the siphon.

Radula (Fig. 156) studied in speci-

mens from l'Etoile, Mauritania and

Sacomar, Angola, with central tooth

ROLAÁN: Columbellidae of the gulf of Guinea

Figures 55-58. Mitrella dartevelleí. 55-57: holotype, 8.9 mm, Gabon, (ZMC) ; 58: protoconch.

Figures 59-65. Mitrella melvilli. 59-61: shells, 8.5, 7.2, 7.4 mm, Palmeirinhas (CER); 62-65:

shells, 10.7, 10.3, 9.8, 10.7 mm, off Luanda, 40-60 m (CER).

Figuras 55-58. Mitrella dartevellei. 55-57: Holotipo, 8,9 mm, Gabón, (ZMC) ; 58: Protoconcha.

Figuras 59-65. Mitrella melvilli. 59-61: conchas, 8,5, 7,2, 7,4 mm, Palmeirinhas (CER); 62-65:

conchas, 10,7, 10,3, 9,8, 10,7 mm, Luanda, 40-60 m (CER).

131

Iberus, 23 (2), 2005

three times as wide as long with hardly

acute posterior corners. Lateral teeth

about three times as long as wide, with

strongly twisted base; cutting edge of

laterals with basal cusp relatively wide

at the base and blunt distally, central

cusp narrow and sharpened, apical cusp

longer and acute.

Operculum (Figs. 143, 165) corneous,

ovoid, light brown, with the insertion

partially divided by a prominence.

Distribution: M. psilla is known from

Mauritania, Senegal and Angola.

It is curious that we have not found

shells of the present species in the area

located between Senegal and Angola,

mainly in Ghana, well sampled by Peter

Ryall (pers. comm.) and by the author,

neither is it referred to by BERNARD

(1984) in his book on Shells of Gabon.

The presence of this species in some

. parts of this poorly known area is very

probable, but also possibly the species

has a bipolar distribution.

Remarks: The shells from Mauritania

(Figs. 35-44) are frequently darker in

colour, more translucent, and with an

evanescent colour pattern, in which the

circles are not well delimited; sometimes

these shells are almost without any

pattern. The colour of the protoconch

and the first whorls is dark brown.

The shells from Dakar are similar

but the colour pattern of circles is

usually well marked and evident.

The shells from Angola (Sacomar

and Praia Amelia) (Figs. 46-52) are more

defined in colour and in ocellate

pattern; the colour of the protoconch is

light yellow with a darker sutural band;

the shells from Baia das Pipas are

similar but more translucent (Figs. 53).

In spite of the differences found

between populations from Mauritania,

Senegal and Angola, the comparison of

the characters of the shell, size, proto-

conch, microsculpture of the protoconch

and radula did not show differences. We

therefore consider all these populations

conspecific.

This species has been synonymized

with doubts by WAGNER AND ABBOTT

(1978) with Mitrella baccata (Gaskoin,

1852), from the Caribbean, but this

species is different, with only some sim-

ilarities in the shell.

Mitrella broderipi (G.B. Sowerby,

1844), described from Alborán Sea but

present in north Africa, is larger and

wider, more solid, has a wider proto-

conch (see Figs. 131 and 132), and differ-

ent radula (Figs. 155 and 156) and oper-

culum (Figs. 142 and 143).

M. alvarezí Rolán and Luque, 2001,

from the Cape Verde Islands, is another

small species and sometimes has ocelli,

but is smaller and the radulae (Figs. 151

and 156) and operculum (Figs. 162 and

165) are also different.

Mitrella denticulata (Duclos in Chenu,

1840), living in Senegal, is similar but

larger, darker, with ocelli only in few

bands, larger light blotches below the

suture and a light apex.

Mitrella dartevellei (Knudsen, 1956) (Figs. 55-58)

Pyrene dartevelli Knudsen, 1956. Atlantide Report, 4: 31, plate 2, figs. 8, 9. [Type locality: Stn. 123,

2” 03'S, 9” 05'E, 50 m off Gabon].

Type material: Holotype in ZMUC (Figs. 55-58).

Other material examined: Gabon: 2 s, “N' kondo” oilfield, 2? 34.1" S, 9 00' E, 120 m (MNAHN).

Description: See KNUDSEN (1956).

Shells have been illustrated in

KNUDSEN (1956) and in BERNARD

(1984). The protoconch is paucispiral

(Fig. 58).

Dimensions: Type material 8.9 to 9.3

mm in length.

132

Distribution: Only known from

Gabon, where it is probably endemic.

Remarks: The narrow base, the deep

suture and the peripheral groove,

always very evident, are diagnostic

characters that clearly differentiate it

from other species of the area of study.

ROLÁN: Columbellidae of the gulf of Guinea

Figures 66-77. Mitrella melvilli. 66: holotype, 8.9 mm (ZMUC); 67-75: shells, 9.5, 11.0, 11.0,

8.6, 9.4, 9.5, 10.0, 9.6, 8.9 mm, Corimba, Luanda, 20 m (CER); 76: detail of the aperture; 77:

protoconch.

Figuras 66-77. Mitrella melvillz. 66: holotipo, 8,9 mm (ZMUC); 67-75: conchas, 9,5, 11,0, 11,0,

8,6, 94, 9,5, 10,0, 9,6, 8,9 mm, Corimba, Luanda, 20 m (CER); 76: detalle de la abertura; 77: pro-

toconcha.

133

Iberus, 23 (2), 2005

Mitrella melvilli Knudsen, 1956 (Figs. 59-77, 140, 158, 167)

Pyrene melvilli Knudsen, 1956. Atlantide Report, 4: 33, pl. 2, fig. 11. [Type locality: Stn. 145, 9” 20'N,

14? 15'W, 32 m, Guinea Conakry].

Type material: Holotype (Fig. 66) of Mitrella melvilli in ZMUC.

Other material examined: Senegal: 1 s, Dakar, 14” 23.5" N, 17? 24.5" W, 65-70 m (MNHN). Ivory

coast: 7 s, Abidjan region (MNHN). Nigeria: 2 s, 04? 03' N, 06” 12” E (MUNHN). Congo: 1 s, Pointe

Noire, Plage Mardamie (CPH). Angola: 56 sp, 10 s, Corimba, Luanda, 10-20 m (MNHN); 106 sp, 12

s, 23 j, Corimba, Luanda, 20 m (CER); 12 s, 26 j, Mussulo (MNHN); 24 sp, 4 s, 2 j, Ilha de Luanda,

40-60 m (MNAHN); 10 sp, 5 s, 6j, off Mussulo, Luanda, 90-100 m (MNHN); 2 s, Palmeirinhas, Luanda

(MNHN); 20 sp, 19 s, 10j, Palmeirinhas, Luanda, 4-8 m (CER); 4 s, 6j, Piambo, 3 m (CER); 1 s, Punta

das Lagostas, 5-20 m (MNHN); 10 sp, 3 s, Luanda, 10 m (CER); 3 sp, 10 s, 5j, off Luanda, 40-60 m

(CER); 8 sp, 6 s, 8 s, Praia Amelia, 5 m (CER).

Description: See also KNUDSEN (1956).

Shell (Figs. 59-75) rather solid, elongate

and sharply pointed.

Protoconch (Figs. 77, 140) with 3 */4

smooth spiral whorls, scarcely convex

with a nucleus of about 130 jm and a

diameter of about 650 um, difficult to

measure because the protoconch and the

beginning of the teleoconch are not

clearly demarcated. Colour of proto-

conch light brown, sometimes with a

more evident spiral band.

Teleoconch with about 6-7 spiral

whorls which are smooth, shiny, almost

flat in profile, with a superficial suture,

and a narrow shelf.

Aperture (Fig. 76) narrow, columella

curved with very small tubercles, about

6 in number. External lip with about 6

teeth, the second and the third from the

upper part larger than the rest.

Colour pattern consisting of a cream

background and yellowish or light

brown irregular ovoid figures, forming

a net, larger below the suture. In some

parts this pattern changes to one of

brown blotches.

Dimensions: holotype 8.9 mm, largest

shells studied 11.5 mm.

Soft parts unknown.

Radula (Fig. 158), studied in

Angolan specimens, with a central tooth

two and a half times as wide as long

and almost rectangular with hardly

acute posterior corners. Lateral teeth

about three times as long as wide, with

strongly twisted base; cutting edge of

laterals with basal cusp wide in the base

and rounded distally, central cusp short

134

and sharpened, and apical cusp much

longer and acute.

Operculum (Fig. 167) ovoid and

somewhat elongate, light brown, almost

transparent, with an ovoid mark of

insertion which has an angular promi-

nence in the middle.

Distribution: Described from Guinea

Conakry, this species is also known

from Senegal, Ivory Coast, Nigeria and

Angola. Probably not present in islands

of the Gulf of Guinea (Sáo Tomé,

Principe, Annobon) never found in spite

of many samplings. |

Remarks: The holotype of M. melvilli

is a slightly faded shell but it still retains

the typical pattern of the species in the

first whorls.

HEDLEY (1899) described a Colum-

bella melvilli from Micronesia (Funa-

futi). Hedley's species is not a true

Columbella and if it is indeed a Mitrella

this could mean that the name for the

African species is pre-occupied. The

characters of the Hedley species prob-

ably fit better within Pyrenola Iredale,

1918, which was considered a

subgenus of Mitrella, but may be a true

valid genus. For this reason, and

pending of further study, we keep the

name M. melvilli for the African

species.

The closest species are M. africana

spec. nov. (see below) and M. aemulata

spec. nov. (see below for differences). M.

africana has a multispiral protoconch,

but is different in colouration, and lives

sympatrically in some areas; M. aemulata

has a paucispiral protoconch.

ROLÁN: Columbellidae of the gulf of Guinea

Figures 78-84. Mitrella africana spec. nov. 78: holotype, 8.8 mm, Palmeirinhas, Luanda, Angola

(MNCN); 79: shell, 7.5 mm, Miamia, Ghana (CER); 80, 81: shells 9.1, 7.9 mm, Mbini, Equator-

ial Guinea (CER); 82: shell, 7.9 mm, Palmeirinhas, Angola (CER); 83: protoconch of the shell of

the Figure 82; 84: detail of the aperture, shell of the Figure 82.

Figuras 78-84. Mitrella africana spec. nov. 78: holotipo, 8,8 mm, Palmeirinhas, Luanda, Angola

(MNCN); 79: concha, 7,5 mm, Miamia, Ghana (CER); 80, 81: conchas 9,1, 7,9 mm, Mbini, Equa-

torial Guinea (CER); 82: concha, 7,9 mm, Palmeirinhas, Angola (CER); 83: protoconcha de la concha

de la Figura 82; 84: detalle de la abertura, concha de la Figura 82.

183

Iberus, 23 (2), 2005

Mitrella africana spec. nov. (Figs. 78-84, 141, 157, 164, 168)

Type material: Holotype (Fig. 78) in MNCN (15.05/46629). Paratypes in the following: AMNH (1),

BMNH (1), MNHN (1), ZSM (1), CJH (1), CFB (1), CER (8), CPR (1), all from the type locality.

Other material examined: Senegal: 5 s, Region de Dakar (MNHN); 1 s, Fleuve Casamance, Zin-

guinchor, 3-4 m (MNHN). Guinea Conakry: 3 s, Mission Gruvel (MNHN); 3 s, Iles Bissagos, Mission

Gain (MNAHN); 15 sp, 47 s, 10 j, Ile de Los, 8-18 m (MNHN). Ivory Coast: 3 s, drag. continental shelf

(MNHN). Ghana: 8 s, Miamia, 8-12 m (CER); 5 sp, 3 s, Takoradi, 4-8 m (CER); 1 sp, Busua, between

the beach and Abokwa Islet, 5 m (CER). Equatorial Guinea: 3 s, Mbini, intertidal (CER). Gabon: 4

sp, Cap Esterias, 0-3 m (MNHN). Sáo Tomé e Principe: 4 s, Baia de Santo Antonio, Principe, 8 m

(CER). Congo: 1 sp, 12 s, ORSTOM beach, Pointe Noire, 5-7 m (MNHN). Angola: 3 s, region Ambrizete

07? 17.49 N, 12* 53.05” E (MNHN); 40 sp, 68 s, 27 j, Barra do Dande, Bengo, infralitoral rocks

(MNAHN); + 300 sp, + 100 s, Cacuaco, Bengo, infralittoral rocks (MNHN), 1 s, off Luanda, 40-60 m

(CER); 1 sp, Cacuaco, Luanda, 2-6 m (CER); 2 s, Cabo Ledo, Luanda, 40 m (MNHN); 1 sp, Morro

dos Veados, intertidal (CER); 1 s, 1 j, Palmeirinhas, 3-6 m (CER).

Type locality: Luanda, Angola.

Etymology: The name reflects the widespread distribution of this species along the West

African coast.

Description: Shell (Figs. 78-82) solid,

lanceolate, with a pointed spire.

Protoconch (Figs. 83, 141) smooth, of

about 2 */2 slightly convex spiral

whorls, with a diameter of about 520

um and a nucleus of 120 um, difficult to

measure because the protoconch and the

beginning of the teleoconch are not

clearly demarcated. Colour of proto-

conch light brown or cream, sometimes

with a more evident dark, spiral band.

Teleoconch with about 6 spiral

whorls which are smooth, shiny, almost

flat in profile, with a superficial suture,

almost smooth.

Aperture (Fig. 84) narrow, columella

curved with very small, inconspicuous

tubercles. External lip with about 6 teeth

of similar size in the upper part, and

smaller ones in the lower part.

Colour pattern formed by a whitish-

cream background and a reddish reticu-

lation forming ovoid figures from which

the subsutural are larger.

Dimensions: Holotype

largest shell studied 10 mm.

Animal (Fig. 168) drawn from mater-

ial collected in Ghana: background

colour cream with small yellowish spots;

irregular blotches of violet-reddish

(similar to that of the shell) along the

lateral sides of the foot, on the tentacles,

behind the eyes and on the siphon, with

a ring of this colour near the tip.

Radula (Fig. 157) with a central

tooth three times as wide as long and

8.8 mm,

136

with hardly acute posterior corners.

Lateral teeth about three times as long

as wide, with strongly twisted base;

cutting edge of laterals with basal cusp

relatively wide basally and blunt dis-

tally, placed close to the central one,

which is close to the apical, both being

short and acute.

Operculum (Fig. 164) light brown,

ovoid, rather transparent, with a mark

of insertion of similar form, which is

almost divided in two by an elongate

prominence along the middle part of the

longer axis.

Distribution: From Senegal to

Angola, including the islands of the

Guinean Gulf.

Remarks: This species has been illus-'

trated from Gabon in KNUDSEN (1956)

and BERNARD (1984) as Pyrene parvula

(Dunker, 1847). The taxon M. parvula

Dunker, 1847 is (after Rios, 1985) a

synonym of M. argus (Orbigny, 1842)

and this species is considered by DE

JONG AND COOMANS (1977) synonym of

M. dichroa Sowerby, 1844. The type local-

ity mentioned in the original description

is “Ind. occid.?” and this is commented

in PACE (1902) and in VAN AARTSEN ET

AL. (1984). The type material is not

known. The description of this taxon is

not corresponding clearly with any West

African species and so we agree with

van AARTSEN ET AL. (1984) considering it

nomen dubium. Furthermore, this name

probably is pre-occuped by Fusus bucci-

ROLÁN: Columbellidae of the gulf of Guinea

Figures 85-88. Mitrella aemulata spec. nov. 85-87: holotype, 7.7 mm, Annobon (MNHN); 88:

protoconch.

Figuras 85-88. Mitrella aemulata spec. nov. 85-87: holotipo, 7,7 mm, Annobón (MNHN); 88: proto-

concha.

noides var. parvula Grateloup, 1833 and

by Buccinum columbelloides var. parvula,

Grateloup, 1847. The American shell of

M. dichroa is similar but a little smaller

and the brownish spiral part among the

ocelli is on the lower part of the whorls

instead to be subsutural.

The West African closest species to

M. africana is the sympatric M. melvilli

Knudsen, 1956 which is similar in size

and pattern, both having a multispiral

protoconch. M. africana has only about 2

1/2 whorls of protoconch, while M.

melvilli has 3 */4 (see Figs. 140, 141). The

colour is yellowish in M. melvilli but

reddish in M. africana. Finally, the colour

pattern is similar, but better differenti-

ated in M. africana where the subsutural

oval figures are always very well

marked, and more constant, while in M.

melvilli it may even disappear, being

more variable, with parts bearing only

light brown blotches. The apertural

tubercles on the external lip are differ-

ent, the second-third are larger in M.

africana. In Angola, both species live

sympatrically in several places.

M. ocellata Gmelin, 1791, from the

north of the West African coast, has a

pattern with ocelli but its size is larger,

the ocelli more uniform, and the opercu-

lum (see Figs. 144 and 164) and radulae

are different (see Figs. 153, 154 and 157).

Some colour forms of Anachis valle-

doríi Rolán and Luque, from the Cape

Verde Islands, may be confused with M.

africana, but that species always has

axial ribs, a paucispiral protoconch and

differences in radula (see Figs. 150 and

157) and operculum (Figs. 161 and 164).

Mitrella aemulata spec. nov. (Figs. 85-88)

Type material: Holotype (Figs. 85-87) in MNHN (from A. Crosnier coll.).

Other material examined: Equatorial Guinea: 3 f, 1j, San Antonio de Palé, Annobon (CER), 2-3 m.

Type locality: Equatorial Guinea, Annobon, 1” 26' S, 5” 37' 30”E, 20-40 m.

Etymology: The specific name is derived from the latin word “aemulatus” which means “imitate”,

and makes allusion to the similarity of this species with M. melvilli.

137

Iberus, 23 (2), 2005

Figures 89-97. Mitrella inesitae spec. nov., Esprahinha, Sáo Tomé. 89: holotype, 10.0 mm (MNCN);

90-95: paratypes: 10.8 (MNHN), 9.6 (BMNH), 10.5 (AMNH), 10.1 (ZSM), 10.5 (CER) and 9.4

mm (CER); 96: detail of the aperture of the holotype; 97: protoconch of the holotype.

Figuras 89-97. Mitrella inesitae spec. nov., Esprahinha, Santo Tomé. 89: holotipo, 10,0 mm

(MNCN); 90-95: paratipos: 10,8 (MNAN), 9,6 (BMNH), 10,5 (AMNH), 10,1 (25M), 10,5

(CER) y 9,4 mm (CER); 96: detalle de la abertura del holotipo; 97: protoconcha del holotipo.

138

ROLÁN: Columbellidae of the gulf of Guinea

Figures 98-102. Mitrella saotomensis spec. nov. 98-100: holotype, 4.4 mm, Praia Mutamba, Sáo

Tomé (MNCN); 101: detail of the aperture of a paratype (CER); 102: protoconch of the holotype.

Figuras 98-102. Mitrella saotomensis spec. nov. 98-100: holotipo, 4,4 mm, Praia Mutamba, Santo

Tomé (MNCN); 101: detalle de la abertura de un paratipo (CER); 102: protoconcha del holotipo.

Description: Shell (Figs. 85-87) ovoid

elongate, solid, with a last whorl more

than half the height.

Protoconch (Fig. 88)) with only one

smooth spiral whorl, with a diameter of

about 750 pm. Colour of protoconch

light brown with two dark bands above

and below the suture.

Teleoconch with about 5 spiral

whorls which are smooth, shiny, almost

flat in profile, with a superficial suture.

Aperture narrow, columella S-

shaped with very small tubercles,

about 5 in number. External lip, with

about 6 teeth, the second and third

from the top being a little larger than

the rest.

Colour pattern formed by a cream

background and a yellowish or light

brown irregular reticulation forming

ovoid figures, which are larger below

the suture, with a suprasutural spire

band formed by small, white, axial

blotches.

Dimensions: Holotype 7.7 mm in

length.

Distribution: Only known from the

holotype and some fragments from

139

Iberus, 23 (2), 2005

Figures 103-108. Mitrella tenebrosa. 103: holotype, 8.4 mm, Esprainha, Sáo Tomé (MNCN); 104,

105: paratype, 8.0 mm, (CPR); 106: paratype, 7.6 mm, (MNHN); 107: protoconch; 108: detail

of the aperture.

Figuras 103-108. Mitrella tenebrosa. 103: holotipo, 8,4 mm, Esprainha, Santo Tomé (MNCN); 104,

105: paratipo, 8,0 mm, (CPR); 106: paratipo, 7,6 mm, (MNAN); 107: protoconcha; 108: detalle de

la abertura.

Amnobon island, from where it is proba-

bly endemic.

Remarks: The most similar species to

M. aemulata is M. melvilli, from conti-

nental Gulf of Guinea, but the proto-

conch of this latter species is multi-

spiral, while that of M. aemulata is

paucispiral with only one spiral whorl.

It is probable that this species origi-

nated from a common ancestor with M.

140

melvilli and, living in isolated condi-

tions in an insular area, evolved from a

planktotrophic to a lecitotrophic larval

development.

Mitrella bruggeni van Aartsen,

Menkhorst and Gittenberger, 1984, from

the Mediterranean and Canary Islands,

has a similar aspect but is larger and

wider, with a more ovoid aperture and

more denticles on the outer lip.

ROLÁN: Columbellidae of the gulf of Guinea

Figures 109-117. Mitrella annobonensis. 109-111: holotype, 4.8 mm, San Antonio de Palé,

Annobon, Guinea Equatorial (MNCN); 112-114: paratypes: 4.7 mm (MNHN), 4.5 mm

(AMNH), 4.0 mm (CER), all from type locality; 115, 116: protoconch; 117: detail of the aperture.

Figuras 109-117. Mitrella annobonensis. 109-111: holotipo, 4,8 mm, San Antonio de Palé,

Annobón, Guinea Equatorial (MNCN); 112-114: paratipos: 4,7 mm (MNAN), 4,5 mm (AMNA,),

4,0 mm (CER), todas de la localidad tipo; 115, 116: protoconcha; 117: detalle de la abertura.

141

Iberus, 23 (2), 2005

Mitrella inesitae spec. nov. (Figs. 89-97, 136, 145-147, 166)

Type material: Holotype (Fig. 89) deposited in the MNCN (15.05/46630). Paratypes in the follow-

ing: AMNH (1, Fig. 92), BMNH (1, Fig. 91), MNHN (1, Fig. 90), ZSM (1, Fig. 93), CJH (4), CEB (10),

CER (75, Figs. 94, 95), CPR (1), all from the type locality.

Other material examined: Equatorial Guinea: 12 f, 7 j, San Antonio de Palé, Annobon, 10-15 m

(CER). Sáo Tomé: 1 s, Morro Peixe (MNHN); 8 j, Praia Mutamba, infralittoral rocks (MNHN); 3 s,

Calypso, st. 14 40? 34' N, 08” 32" W (MNHN); 20 sp, 21 s, 193, Esprainha (Neves), infralitoral (MNHN);

136 sp, 14 s, 41 j, Lagoa Azul, 4-8 m (CER); 37 sp, 6 s, 37 j, Esprainha, 6-10 m (CER), 8 s, Praia

Mutamba, 4-8 m (CER).

Type locality: Esprainha, east coast of Sáo Tomé, Republic of Sio Tomé and Principe.

Etymology: The specific name is after the author” s niece Inés Alvarez Torres, of Trubia, Oviedo,

Spain, companion on many collecting trips.

Description: Shell (Figs. 89-95)

fusiform, solid, with a last whorl larger

than half the height.

Protoconch (Figs. 97, 136) short,

smooth, difficult to distinguish from the

beginning of the teleoconch, with about

one and a half whorls, yellowish in

colour, sometimes darker, scarcely

lighter in the suture, about 620 um in

diameter with nucleus about 230 um.

Teleoconch with 6-7 whorls, smooth,

shiny, suture not impressed, profile

almost flat.

Aperture (Fig. 96) elongate and

narrow. External lip thickened exter-

nally, internally with some teeth

present, the first one standing out alone

and separated from the second. Second

tooth largest, sometimes seeming to be

formed by the fusion of two, third tooth

a little smaller and the remaining three

or four down to the base, very small.

Inner lip with small tubercles on its

lower part. Siphonal canal short and

wide.

Background colour brown, with

numerous yellowish or cream ocellate

patterns overall, which are small and a

little elongated spirally. Subsutural area

frequently with larger brown blotches

and between them areas with white or

cream colour. In some shells, another

similar band in the middle of the last

whorl, partially visible above the suture

of spire whorls. Basal area with 12-16

spiral cords.

Dimensions: Most shells between 9.00

and 11.0 mm, holotype 10.0 mm in

height.

Soft parts unknown.

142

Radula (Fig. 147) with a central tooth

two times as wide as long and with

rounded corners. Lateral teeth about

three times as long as wide with

strongly twisted base; cutting edge of

laterals with basal cusp wide basally

and blunt distally, central and apical

cusps narrow and acute.

Operculum (Fig. 145, 146, 166) cor-

neous, ovoid, light brownish, with the

insertion mark also ovoid elongate and

with a very small prominence at the

middle.

Distribution: Only known from Sáo

Tomé from where it is probably

endemic.

Remarks: Mitrella inesitae spec. nov. is

different from most of the west African

species which are smaller, except M.

pallaryi which is wider, has a multispi-

ral protoconch and a different pattern.

The most similar Mediterranean species

is M. scripta (Linné, 1758) which is

larger and with a more variable colour

pattern. They can be differentiated

because the aperture of M. inesitae is

relatively more elongate, and the tuber-

cles on the columella are smaller, while

those on the external lip are larger, the

upper one being more separated from

the second.

Mitrella lanceolata (Locard, 1886),

from the Mediterranean, is more elon-

gate, with a wider and relatively shorter

aperture and a simpler colour pattern.

Some elongate specimens of the

Mediterranean M. gervillii (Payraudeau,

1826) may have a similar aspect, but

they are very large, with a wider aper-

ture and smaller denticulation in it.

ROLÁN: Columbellidae of the gulf of Guinea

Figures 118-121. Mitrella condeí spec. nov. 118: holotype, 16.5 mm, Santa María, Angola

(MNHN); 119-120: paratype, 16.2 mm, Santa Maria, Angola (MNCN); 121: protoconch of the

holotype. Figures 122, 123. Cotonopsis molfinsi, 40 mm, (CJH).

Figuras 118-121. Mitrella condei spec. nov. 118: holotipo, 16,5 mm, Santa María, Angola (MNAN);

119-120: paratipo, 16,2 mm, Santa Maria, Angola (MNCN); 121: protoconcha del holotipo. Figuras

122, 123. Cotonopsis molfinsi, 40 mm, (CJH).

Iberus, 23 (2), 2005

Mitrella saotomensis spec. nov. (Figs. 98-102, 137)

Type material: Holotype (Figs. 98-100) in MNCN (15.05 /46631); paratypes: CER (1 s, without apex,

Fig. 101), CJH (2 s).

Other material examined: Sáo Tomé: 14 f, Praia Mutamba, 3-8 m (CJH); 1 f, Praia Mutamba, 2-8 m

(CER); 15 f, Lagoa Azul (CHD).

Type locality: Praia Mutamba, Sáo Tomé, 5 m, Republic of Sáo Tome and Principe.

Etymology: The specific name is after the island where the species was collected.

Description: Shell (Figs. 98-100)

fusiform, solid, with a last whorl larger

than half the height.

Protoconch (Figs. 102, 137) short,

smooth, with 1 */4 spiral whorls, clearly

differentiated from the beginning of the

teleoconch, because the spiral striation

appears, diameter about 500 qm with

nucleus 250 qm. Colour of protoconch

yellowish with a darker sutural band,

and a dark point in the apex.

Teleoconch with 4 whorls, shiny,

suture not impressed, whorl profile

almost flat. Spiral striae appearing at the

beginning of the teleoconch, respec-

tively, 6 in the first whorl, 10 in the

second, 16 in the following and more

that 60 on the last whorl; very small and

closely spaced, except at the base of the

shell, where the 17 lower ones are wider

and define small spiral cords.

Aperture (Fig. 101) elongate and

narrow. External lip widely thickened

externally, internally with some teeth, the

uppermost standing out alone separated

from the upper extreme and from the

second one. Second tooth largest, the fol-

lowing one a little smaller, the remaining

four teeth, down to the base, very small.

Inner lip with 5 small tubercles on its lower

part. Siphonal canal short and wide.

Colour yellowish brown on back-

ground, darker near the suture with a

band on the last whorl which continues

the suture; this background covered by

numerous yellowish or cream ocelli

overall.

Dimensions: Holotype 4.5 mm in

height.

Distribution: Only known from Sáo

Tomé Island, from where it is probably -

endemic.

Remarks: The main differences with

the most similar species are the follow-

ing:

Mitrella broderipi (Sowerby, 1844) and

M. bruggen1t van Aartsen, Menkhorst and

Gittenberger, 1984, from the Alboran

Sea, are larger, wider and with a wider

aperture with smaller denticulation.

Mitrella alvarezi Rolán and Luque,

2001, from Cape Verde Islands, is larger,

darker, with a more irregular pattern

with parts without ocelli, and a totally

smooth surface.

Mitrella psilla, from West Africa, is

similar in size or slightly larger, but the

shell is wider; its pattern is formed by

larger ocelli, the aperture is wider, the

teeth of the outer lip are regularly

spaced, and it lacks the evident tuber-

cles on the columella.

Mitrella tenebrosa spec. nov. (Figs. 103-108)

Type material: Holotype (Fig. 103) in MNCN (15.05/46632). Paratypes: CER (1), CPR (1, Fig. 104,

105), both from the type locality; MNHN (1, Fig. 106) from Neves, Sáo Tomé.

Type locality: Esprainha, Sáo Tomé, Republic of Sáo Tome and Principe.

Etymology: The specific name alludes to its dark colour.

Description: Shell (Figs. 103-106)

fusiform, solid, with a last whorl larger

than half the height.

Protoconch (Fig. 107) studied in few

shells and not in good condition; appar-

144

ently short, smooth, perhaps with 1 */4

to 1 1/2 spiral whorls, but difficult to

distinguish from the beginning of teleo-

conch; diameter about 600 um with

nucleus about 250 qm. Colour of proto-

ROLÁN: Columbellidae of the gulf of Guinea

17.7 mm, Praia

Figures 124-130. Strombina descendens, 124-127: shells of 20.2, 18.8, 20.2,

Amelia, Namibe, 40-60 m (MNHN); 128: Detail of the aperture; 129, 130: protoconch.

Figuras 124-130. Strombina descendens, 124-127: conchas de 20,2, 18,8, 20,2, 17,7 mm, Praia

Amelia, Namibe, 40-60 m (MNHN); 128: detalle de la abertura; 129, 130: protoconcha.

145

Iberus, 23 (2), 2005

conch dark brown in the best preserved

specimen.

Teleoconch with 4-6 smooth and

hardly convex whorls, shiny, with

suture impressed. Basal area with 8-10

spiral cords.

Aperture (Fig. 108) elongate and

narrow. External lip slightly thickened

externally. Internally, 6 strong teeth with

a regular separation among them. Inner

lip with 6 small tubercles on its lower

part. Columella straight in the middle

and curved at the extremes. Siphonal

canal short and wide.

Background colour dark brown,

darker near the suture and with very

numerous small oval ocelli overall; in

other shells this pattern is not seen

because of erosion.

Dimensions: Holotype 8.4 mm in

height, other shells slightly smaller.

Distribution: Only known from Sáo

Tomé, deeper than 15-20 m. As some

crabbed material was collected in

Esprainha, it is supposed that the

species lives in deeper water.

Remarks: The closest species to M.

tenebrosa and their main differences are

the following:

Mitrella inesitae is sympatric, but its

shell is larger, more elongate, wider, and

with a more colourful pattern. The aper-

ture is narrower and the upper teeth on

the external lip are larger and more sep-

arated.

Mitrella pallaryi, from the European

and West African coast, is a larger

species with a multispiral protoconch.

Mitrella saotomensis is sympatric, and

it has a smaller shell and its pattern is

formed by circular ocelli. The teeth on

the external lip are not regular.

Mitrella africana and M. melvillí, from

the West African continental coast, have

multispiral protoconch, are lighter in

colour and the teeth on the external lip

are different in size, the upper ones

being larger.

Mitrella annobonensis spec. nov. (Figs. 109-117, 138, 139)

Type material: Holotype (Figs. 109-111) in MNCN (15.05/46633); paratypes in MNAHN (1, Fig. 112),

AMNH (1, Fig. 113) and CER (1, Fig. 114).

Type locality: San Antonio de Palé, Annobon, 10-15 m, Equatorial Guinea.

Etymology: The specific name is after the island where the species was found.

Description: Shell (Figs. 109-114)

ovoid-conical, very solid, with a last

whorl which represents ?/3 of the height.

Protoconch (Figs. 116, 138), short,

with 1 */4 whorls, brown in colour,

darker along the suture, diameter of

about 520 um with nucleus 230 um,

apparently smooth, but spiral lines with

irregular nodules can be seen with high

magnification (Fig. 139).

Teleoconch with about 4 whorls, shiny,

suture not impressed, whorl profile almost

flat. Termination of protoconch well

defined because spiral striae appear at the

(Right page) Figures 131-139. Protoconchs and microsculpture. 131: Mitrella broderipi, VEtoile,

Nouadibou, Mauritania; 132: Mitrella psilla, 'Etoile, Nouadibou, Mauritania; 133: Mitrella psilla,

Sacomar, Angola; 134: microsculpture of the protoconch of M. psilla, Sacomar; 135: microsculpture

of the protoconch of M. psilla, DEtoile; 136: Mitrella inesitae sp. nov., Lagoa Azul, Sáo Tomé; 137:

Mitrella saotomensis sp. nov., Praia Mutamba, Sáo Tome; 138: Mitrella annobonensis sp. nov., San

Antonio de Palé, Annobon; 139: microsculpture of the protoconch of M. annobonensis sp. nov.

(Página derecha) Figuras 131-139. Protoconchas y microsculpture. 131: Mitrella broderipi, /'Etoile,

Nouadibou, Mauritania; 132: Mitrella psilla, Etoile, Nouadibou, Mauritania; 133: Mitrella psilla,

Sacomar, Angola; 134: microscultura de la protoconcha de M. psilla, Sacomar; 135: microscultura de

la protoconcha de M. psilla, L'Etoile; 136: Mitrella inesitae sp. nov., Lagoa Azul, Santo Tomé; 137:

Mitrella saotomensis sp. nov., Praia Mutamba, Santo Tome; 138: Mitrella annobonensis sp. noz,

San Antonio de Palé, Annobón; 139: microscultura de la protoconcha de M. annobonensis sp. nov.

146

nea

Columbellidae of the gulf of Gui

ROLÁN

0 um

147

Iberus, 23 (2), 2005

beginning of the teleoconch, 8 on the first

whorl, 17 on the second, 27 on the fol-

lowing and very numerous and difficult

to count on the last whorl. Spiral striae

very small and very attenuated in some

places. Basal area with about 15 well

defined spiral cords. Irregular axial ribs

present on all the whorls, about 15 on the

first whorl and 25 on the following, less

noticeable on the last whorl.

Aperture (Fig. 117) elongate and

narrow. External lip widely thickened

externally, internally with about 7 teeth,

the uppermost being the largest, and the

lower smaller. Inner lip with no tuber-

cles. Siphonal canal short and wide.

- Background colour reddish brown

with numerous yellowish or cream

ocelli overall, sometimes less marked on

the convexity of the last whorl.

Dimensions: Holotype 4.8 mm in

height.

Soft parts, operculum and radula

unknown.

Distribution: Only known from

Annobon, Equatorial Guinea, from

where it is probably endemic.

Remarks: This species may be differ-

entiated from any of the other known

West African species of the genus

because it is a very solid, short and rela-

tively wider shell, with narrow aper-

ture, and spiral striae:

Mitrella alvarezi Rolán and Luque,

2001 is larger, more fragile, darker, with

a more variable pattern partially

without ocelli, and a totally smooth

surface.

Mitrella psilla is of similar size, but

less solid and globose, the aperture is

wider and the colour lighter, yellowish,

with larger ocelli.

Mitrella broderipi (G. B. Sowerby,

1844) is larger, relatively wider, more

fragile, the pattern is formed by larger

ocelli, the aperture is wider, the teeth of

the outer lip are regularly distant, and

lacks tubercles on the columella.

Mitrella condei spec. nov. (Figs. 118-121)

Type material: Holotype (Fig. 118) in MNHN. Paratypes: 1 s, Baia de Lucira (Bissonga), Namibe,

Angola (MNHN); 1 s, Santa Maria, Angola, (MNCN 15.05/46634).

Type locality: Praia Amelia, Namibe, Angola.

Etymology: The species is dedicated to the Spanish malacologist Javier Conde, Associate Editor of

Iberus, for his continuous help in our work.

Description: Shell (Figs. 118-120)

solid, conical elongate, smooth, with a

large last whorl.

Protoconch (Fig. 121) difficult to see

due to the lack of separation from teleo-

conch, probably of about 1 whorl, with

diameter about 700 um.

Teleoconch of about 7 whorls, totally

flat, with suture incised but not deep.

Last whorl large, with the same height

as the spire, and with a peripheral angu-

lation; from this angulation the profile

of the shell is concave, the last whorl

ending in a narrow base presenting 13

narrow spiral ribs.

Aperture rhomboidal, columella with

its central part in the same direction as the

axis of the shell, deviated above and below

(Right page) Figures 140, 141. Protoconchs. 140: Mitrella melvillí, Luanda, Angola; 141: Mitrella

africana, Palmeirinhas, Angola. Figures 142-147. Opercula. 142: Mitrella broderipi, VEtoile, Mauri-

tania; 143: Mitrella psilla, Sacomar, Angola; 144: Mitrella ocellata albine. Dakar, Senegal; 145, 146:

Mitrella inesitae sp., Lagoa Azul, Sio Tomé. Figure 147. Radula of Mitrella inesitae sp. nov. A. Lagoa

Azul, Sáo Tome.

(Página derecha) Figuras 140, 141. Protoconchas. 140: Mitrella melvilli, Luanda, Angola; 141: Mitrella

africana, Palmeirinhas, Angola. Figuras 142-147, Opercula. 142: Mitrella broderipi, Etoile, Mauritania;

145: Mitrella psilla, Sacomar, Angola; 144: Mitrella ocellata albine. Dakar, Senegal: 145, 146: Mitrella inesi-

tae sp., Lagoa Azul, Sáo Tomé. Figure 147. Rádula de Mitrella inesitae sp. nov. A. Lagoa Azul, Sáo Tome.

148

ROLÁN: Columbellidae of the gulf of Guinea

149

Iberus, 23 (2), 2005

forming an open S. On the external lip

there are 8 teeth continued by lirae,

number 2 and 3 being the largest.

- Colour light brown formed by small

oval ocelli oriented spirally.

Dimensions: Holotype 16.5 mm in

height, other known specimens of

similar size.

Soft parts, radula and operculum

unknown.

Distribution: Only known from

Angola.

Remarks: We consider that this

species has similarity with Mitrella minor

Scacchi, 1836, type species of the sub-

genus Columbellopsis, due the character-

istic profile and the narrow base. At

present, there is not complete agreement

about the validity of the genus Columbel-

lopsis, as is commented on by BOYER

AND ROLÁN (2005).

From its form and size it could be

included in the genus Strombina Mórch,

1852, but the species of this latter genus

usually have clear axial sculpture and

spiral striations, no present in M. condei.

The juvenile shells of M. condei may

be similar to those of Strombina descendens

(Martens, 1904) (see below) since this

species has 3-5 smooth whorls and a

similar pattern of small oval ocelli. They

can be differentiated because S. descendens

have a wider protoconch (about 150-200

um more); also the suture is shallow in M.

condei while it is very deep in S. descendens

and with an evident separation.

Genus Cotonopsis Olsson, 1942

Type species: Strombina (Cotonopsis) panacostariceus Olsson, 1942. [Type locality: Pliocene of

Burica Peninsula, Costa Rica, Charco Azul Formation]. By original designation.

Cotonopsis monfilsi Emerson, 1993 (Figs. 122, 123)

Cotonopsis monfilsi Emerson, 1993. The Nautilus, 106 (4): 147. [Type locality: off St. Louis (16” 02'N

16” 30'W), Senegal, 300 m.]

Type material: Not examined.

Other material examined: Senegal: 2 s, “Louis Sauger” at 600-1000 m (CJH). Guinea Conakry: 3 s,

from 40-50 m (CJH); 15 s, Saint Louis, 300-1000 m (MNHN); 3 s, Senegal, 250-300 m (MNHN);

Angola: 2 s, from South Angola fishermen (MNHN).

(Right page) Figures 148-158. Radulae of Columbellidae. 148: Columbella rustica, Antalya, Turky,

shell of 18.6 mm; 149: Columbella adansoni, Lanzarote, Canary islands, shell of 19.5 mm; 150:

Anachis valledori, Sal Rei, Boavista, Cape Verde Ís., shell of 6.8 mm; 151: Mitrella alvarezz, Sal Rei,

Boavista, Cape Verde Islands, shell of 4.0 mm; 152: Mitrella pallaryi, Camarinas, Galicia, shell of

14.5 mm; 153: Mitrella ocellata, N'gor, Dakar, Senegal, shell of 11.0 mm (decolate); 154: Mitrella

ocellata albine form, Dakar, Senegal, shell of 11.8 mm (not decolate); 155: Mitrella broderipz, Baie

de Etoile, Nouadhibou, Mauritania, shell of 7:5 mm; 156: Mitrella psilla, Sacomar, Angola, shell

of 4.0 mm; 157: Mitrella africana, Rio Muni, Equatorial Guinea, shell of 8.6 mm; 158: Mitrella

melvillz, Corimbo, Luanda, shell of 9.5 mm.

(Página derecha) Figuras 148-158. Radulas de Columbellidae. 148: Columbella rustica, Antalya,

Turquía, concha de 18,6 mm; 149: Columbella adansoni, Lanzarote, Islas Canarias, concha de 19,5

mm; 150: Anachis valledori, Sal Rei, Boavista, Islas de Cabo Verde, concha de 6,8 mm; 151: Mitrella

alvarezi, Sal Rei, Boavista, Islas de Cabo Verde, concha de 4,0 mm; 152: Mitrella pallaryi, Camariñas,

Galicia, concha de 14,5 mm; 153: Mitrella ocellata, N'gor, Dakar, Senegal, concha de 11,0 mm

(decapitada); 154: Mitrella ocellata forma albina, Dakar, Senegal, concha de 11,8 mm (no decapi-

tada); 155: Mitrella broderipi, Bahía de Etoile, Nouadhibou, Mauritania, concha de 7,5 mm; 156:

Mitrella psilla, Sacomar, Angola, concha de 4,0 mm; 157: Mitrella africana, Río Muni, Guinea Ecua-

torial, concha de 8,6 mm; 158: Mitrella melvilli, Corimbo, Luanda, concha de 9,5 mm.

150

ROLÁN: Columbellidae of the gulf of Guinea

157

Iberus, 23 (2), 2005

Description: See EMERSON (1993). We Angola, mainly found in deep

illustrate a typical shell (Figs. 122, 123). water.

Distribution: It is a species with a Remarks: No other species is similar

wide distribution from Senegal to to this one.

Genus Strombina Mórch, 1852

Type species: Strombina lanceolata (G. B. Sowerby, 1832), by subsequent designation Bucquoy,

Dautzenberg and Dollfus, 1882, p. 78.

Diagnosis: H. and A. ADAMS (1858: 186), THIELE (1935: 457) and JunG (1989).

Strombina descendens (Martens, 1904) (Plate 19) (Figs. 124-130)

Mangelia descendens von Martens, 1904. Ergebn. dtsch. Tiefsee-Exp., 7: 7, pl. 3, fig. 20.

Type material: Not examined.

Other material examined: Equatorial Guinea: 4 s, 2 j, St. 45 0? 25' N, 9% 00' E (col. Marche-

Marchad, MNHN). Gabon: 6 s, Calypso, stn. 451, 0? 25' N, 09% 00 E, 73 m (MNHN); 3 s, Port-

Gentil, 0% 47.4' S, 8” 43.6' E, 25 m (MNHN>); 1 s, St. 123, 2* 03.5” S, 9” 05' E, (MNHN exZMUC).

Congo: 1 s, 3j, Pointe Noire, Plage Mendame (CPH). Angola: 12 s, Ilha de Luanda, 40-60 m

(MNHN); 5 s, Palmeirinhas, Luanda (MNHN); 5 s, Praia Amelia, Namibe, Angola (CER); 8 s, 4 j,

Praia Amelia, Namibe, 40-60 m (MN HN).

Description: This species has been

described and illustrated in KNUDSEN,

(1956: 36, pl 3, fi8- 21), as “Pyrene descen-

dens. Some specimens are here illus-

trated (Figs. 124-127) as well as the aper-

ture (Fig. 128).

The protoconch (Figs. 129, 130) is

short, but it is difficult to see the whorls

since there is no clear separation with

the teleoconch; its diameter varies

between 750 and 900 um.

Dimensions: Between 14 and 20 mm.

Soft parts, Ooperculum and radula

unknown.

Distribution: Known from Equatorial

Guinea to Angola.

Remarks: The species is tentatively

included here in the genus Strombina

Moóorch, 1852 because of its similarity with

CONCLUSIONS

The family Columbellidae is repre-

sented by a high number of species in

the Gulf of Guinea, at a similar level to

the Mediterranean, where 13 species in

4 genera are known (GIANNUZZI-SAVELLI

ET AL., 2003). In the present work we

considered 15 species in 5 genera. Of

152

some West Atlantic and Pacific species of

this genus. If further studies prove this

relationship, it should be the only known

species of this genus in West African.

The specimens from the north of the

distribution area (Gabon, Equatorial

Guinea) are smaller (usually 10-12 mm)

and the dimensions given by KNUDSEN

(1956) are about 14 mm. By contrast, the

Angolan shells are larger and may reach

more than 20 mm.

The comparison of shells from

Gabon and Angola, very different in

size, did not show other significant dif-

ferences: the protoconch is slightly

smaller, and the smooth whorls may be

1-2 lesser than the Angolan specimens.

Anyway, the main characters are equal

and we consider them conspecific.

course, if we consider the complete West

African area including the species living

in the Canary Islands and in Senegal,

the total number of species would be

notably increased (see SEGERS AND

SWINNEN, 2003, HERNANDEZ AND BOYER,

2005, and PELORCE AND BOYER, 2005).

ROLÁN: Columbellidae of the gulf of Guinea

Figures 159-167. Opercula (from the same specimens from which the radula was studied). 159: C.

rustica; 160: C. adansont;, 161: A. valledori, 162: M. alvarezi; 163: M. pallaryi; 164: M. africana;

165: M. psilla; 166: M. inesitae; 167: M. melvillí. Figures 168, 169. Soft parts of Mitrella species.

168: M. africana; 169: M. psilla.

Figuras 159-167. Opérculos (de los mismos ejemplares de los que fue estudiada la radula). 159: C.

rustica; 160: C. adansoni; 161: A. valledori; 162: M. alvarezi; 163: M. pallaryi; 164: M. africana;

165: M. psilla; 166: M. inesitae; 167: M. melvilli. Figuras 168, 169. Partes blandas de especies de

Mitrella. 168: M. africana; 169: M. psilla.

153

Iberus, 23 (2), 2005

Some of the species studied have a

large distribution area (Columbella adan-

soni, Mitrella pallaryi, M. psilla, M.

melvilli, M. africana and C. monfilsi). Of

these species, only one (M. pallary) has a

very extended area (all the Mediter-

ranean, most of the Macaronesian archi-

pelagos and all West Africa); one other

species (C. adansoni) extends to the Mac-

aronesian islands besides the West

african coast, but not to the Mediter-

ranean. Four more species (M. psilla, M.

melvilli, M. africana and C. monfilsi) have

a range approximately equivalent to the

extension of the studied area (from

Senegal to south Angola).

The single species with a large range

which was undescribed is M. africana;

this species was known but it was con-

sidered as part of an American taxon

(M. parvula).

Some other species have a smaller

range (Strombina descendens, from Gabon

to Angola). The rest of the species has a

short range, being probably endemic to

small areas of coast (as Anachis ryalli,

only found in Ghana, and Mitrella

condei, in Angola); or they are endemic

to islands (M. inesitae, M. saotomensis

and M. tenebrosa, endemic in Sáo Tomé,

and M. annobonensis and M. aemulata, in

Amnobon).

Of these 15 species, 8 are described

as new. These species were not known

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The study of the protoconchs showed

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SABELLI, B. AND SPADA, G., 1981. Guida illus-

trata all'identificazione delle conchiglie del

Mediterraneo. Suppl. Bollettino Malacologico,

17 (11-12). No pagination.

SABELLI, B, GIAANNUZZI-SAVELLI, R. AND BEDULLI,

D., 1990. Catalogo annotato del molluschi marini

del Mediterranean. Libreria Naturalistica

Bolognese, Bologna. 348 pp.

SCHIRO, G., 1978. Il genere Mitrella Risso, 1826,

nel Mediterranean (Prosobranchia-Bucci-

noidea). I. La Conchiglia, 10 (114-115): 8-10.

SCHIRO, G., 1979. The genus Mitrella Risso, 1826

in the Mediterranean. La Conchiglia, 11 (120-

121): 7-8.

SEGERS, W. AND SWINNEN, F., 2003. On the ocur-

rence of Zafra exilis (Philippi, 1849) on the Ca-

nary Islands. Gloria Maris, 42 (4-5): 101-103.

SIERRA, A., GARCÍA, L. AND LLoORíS, D., 1978.

Trofismo y competencia alimentaria en as-

teroideos de la bahía de Almería. Investiga-

ciones Pesqueras, 42 (2): 485-499.

TATE, R., 1869. Appendix [to] S. P. Woodward,

A manual of the Mollusca, 2nd ed. separately

paged 1-86; 27 text figs. London.

VAN AARTSEN, J. J., MENKHORST, H. P. M. G. AND

GITTENBERGER, E., 1984. The marine Mol-

lusca of the Bay of Algeciras, Spain, with

general notes on Mitrella, Marginellidae and

Turridae. Basteria. Suppl. 2: 1-135.

WAGNER, R.J. L. AND ABBOTT, R. T., 1978. Stan-

dard Catalog of Shells. American Malacologist,

Greenville, pagination by sections.

O Sociedad Española de Malacología Iberus, 23 (2): 157-163, 2005

A new Fusinus (Gastropoda: Fasciolariidae) from Japan

Un nuevo Fusinus (Gastropoda: Fasciolariidae) de Japón

Roland HADORN* and Mitsuo CHINO**

Recibido el 21-1-2004. Aceptado el 12-X-2005

ABSTRACT

Examination of the holotype of Fusinus hyphalus M. Smith, 1940 reveals that FE. hyphalus

is a subjective junior synonym of F. graciliformis (Sowerby, 1880). The species often called

F. hyphalus and illustrated by authors is without a name. Fusinus satsumaensis spec. nov.

[type locality: off Akune, Kagoshima Prefecture, Japan] is proposed as the name for this

misidentified species in the subgenus Chryseofusus Hadorn and Fraussen, 2003 and com-

pared with F. (Chryseofusus) graciliformis (Sowerby, 1880), F. (C.) chrysodomoides

(Schepman, 1911), F. (C.) jurgeni Hadorn and Fraussen, 2002 and F. (C.) westralis

Hadorn and Fraussen, 2003.

RESUMEN

El examen del holotipo de Fusinus hyphalus M. Smith, 1940 revela que F. hyphalus es un

sinónimo juvenil de F. graciliformis (Sowerby, 1880). La especia a menudo denominada F.

hyphalus e ilustrada por varios autores carece de nombre. Se propone el nombre de Fusi-

nus satsumaensis spec. nov. para esta especie inidentificada, dentro del subgénero Chry-

seofusus Hadorn y Fraussen, 2003. Se compara con F. (Chryseofusus) graciliformis

(Sowerby, 1880), F. (C.) chrysodomoides (Schepman, 1911), FE. (C.) jurgeni Hadorn y

Fraussen, 2002 y F. (C.) westralis Hadorn y Fraussen, 2003.

KEY WORDS: Mollusca, Gastropoda, Fasciolariidae, Fusinus, Chryseofusus, Japan, new species.

PALABRAS CLAVE: Mollusca, Gastropoda, Fasciolariidae, Fusinus, Chryseofusus, Japón, nueva especie.

INTRODUCTION

HADORN AND FRAUSSEN (2003) We have since found the holotype in

recently described the subgenus the Florida Museum of Natural History

Chryseofusus in Fusinus to accommodate

a number of species sharing conchologi-

cal characteristics different from typical

Fusinus and described new species in

this subgenus. One of the treated species

was F. hyphalus M. Smith, 1940. At that

time of that paper the two authors were

unable to locate the holotype of F.

hyphalus.

* Dreihubelweg 23, CH-3250 Lyss, Switzerland.

in Gainesville, USA. Examination of the

holotype reveals that E. hyphalus is a

juvenile specimen of E. graciliformis.

Based on the original description and

the poor figure of the holotype,

HADORN AND FRAUSSEN (2003) failed to

recognize this synonymy and confused

it with the misidentified species often

called FE. hyphalus by authors and

** 6-23-18-202, Arima, Miyamae-ku, Kawasaki, 216-0003, Japan.

IS7

Iberus, 23 (2), 2005

recorded from Japan, the East China Sea

and the Philippine Islands.

Abbreviations used:

ANSP Academy of Natural Sciences,

Philadelphia, USA.

BMNH The Natural History Museum,

London, United Kingdom.

FLMNH/UF Florida Museum of

Natural History, Gainesville, Florida,

USA.

MC Collection of Mitsuo Chino,

Kawasaki, Japan.

NSMT National

Tokyo, Japan.

RH Collection of Roland Hadorn, Lyss,

Switzerland.

RMNH National Museum of Natural

History — Naturalis, Leiden, the

Netherlands.

WAM Western Australian Museum,

Perth, Australia. .

ZMA Zoologisch Museum, University

of Amsterdam, Amsterdam, the

Netherlands.

dd dead collected specimen.

juv juvenile specimen.

Science Museum,

MNHN Muséum national d'Histoire

naturelle, Paris, France.

lv live collected specimen.

subad subadult specimen.

SYSTEMATICS

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.

Type species: Murex colus Linnaeus, 1758, by typification of replaced name.

Subgenus Chryseofusus Hadorn and Fraussen, 2003

Chryseofusus Hadorn and Fraussen, 2003. The deep-water Indo-Pacific radiation of Fusinus.

Iberus, 21 (1): 207-240.

Type species: Fusus chrysodomoides Schepman, 1911.

Fusinus (Chryseofusus) satsumaensis spec. nov. (Figs. 1-8)

Fusinus (Simplicifusus) hyphalus M. Smith, 1940. — Kira (1962: 85); Springsteen and Leobrera (1986:

177+179, pl. 48, fig. 2). non M. Smith, 1940.

Simplicifusus hyphalus (M. Smith, 1940). — Higo, Callomon and Goto (1999: 263). non M. Smith,

1940.

Fusinus (Chryseofusus) hyphalus M. Smith, 1940. — Hadorn and Fraussen (2003: 218-219, figs. 22,

23). non M. Smith, 1940.

Type material: Holotype (78.1 x 24.7 mm, lv), NSMT-Mo 73704: 350-400 m, off Akune, Kagoshima

Prefecture, Japan.

Paratype 1 (68.4 x 22.2 mm, lv), MC; paratype 2 (65.9 x 21.3 mm, lv), RH: 350-400 m, off Akune,

Kagoshima Prefecture, Japan. Paratype 3 (72.9 x 24.4 mm, lv), MC; paratype 4 (75.9 x 25.1 mm, lv),

MNHN, paratype 5 (73.5 x25.1 mm, lv), ANSP 412950: 250 m, southwest of Cape Noma, Kagoshima

Prefecture, Japan.

Other material examined: Japan, from type locality, 1 dd, MC. - Japan, off Cape Noma, Kagoshima

Prefecture, 1 dd subad, MC. - Taiwan, deep water, 1 lvjuv/1 lv, RH. - Taiwan, Keelung, 1 lv/1 dd

subad, RH. - Unknown locality, 1 dd, RH.

Type locality: Off Akune, Kagoshima Prefecture, Japan.

Etymology: F. (C.) satsumaensis spec. nov. is named after the type locality. Satsuma is the ancient

name of Western Kagoshima which is famous in the Japanese history with regard to the Meiji

Restoration 1868.

158

HADORN AND CHINO: A new Fusinus (Gastropoda: Fasciolariidae) from Japan

Figures 1-8. Fusinus (Chryseofusus) satsumaensis spec. nov. 1, 2: holotype NSMTI=Mo 73704, Japan,

off Akune, Kagoshima Prefecture, 78.1 mm; 3,4: paratype MC, Japan, off Akune, Kagoshima Pre-

fecture, 68.4 mm; 5, 6: paratype RH, Japan, off Akune, Kagoshima Prefecture, 65.9 mm; 7: holo-

type NSMT-Mo 73704, spire tip; 8: Operculum.

Figuras 1-8. Fusinus (Chryseofusus) satsumaensis spec. nov. 1, 2: holotipo NSMI-Mo 73704, Japón,

frente a Akune, Kagoshima Prefecture, 78,1 mm; 3,4: paratipo MC, Japón, frente a Akune, Kagoshima

Prefecture, 68,4 mm; 5, 6: paratipo RH, Japón, frente a Akune, Kagoshima Prefecture, 65,9 mm; 7:

holotipo NSMI-MO 73704, ápice de la espira; 8: opérculo.

159

Iberus, 23 (2), 2005

Description: Shell rather large for

subgenus (up to 80 mm), thin, light-

weight but solid, fusiform with elongate

spire, uniformly white, pale or light

yellow, consisting of about 9 slightly

convex whorls with weak subsutural

concavity. Spire long and pointed, body

whorl inflated and ventricose in adult

specimens, siphonal canal narrow, occa-

sionally curved. Aperture including

intact siphonal canal slightly longer

than spire. Suture fine but distinct, only

weakly incised.

Protoconch relatively large, white,

glossy, bulbous, consisting of 1 */4 - 1

1/2 whorls, final */4 whorl with 3 or 4

narrow axial riblets, reaching from

suture to suture. Transition to teleo-

conch marked by a varix. Diameter 1.1-

1.2 mm.

Axial sculpture inconspicuous, only

visible on upper teleoconch whorls.

Axial ribs weak, narrow, extending

from suture to suture, interspaces

narrow. 10 or 11 ribs on 3 uppermost

teleoconch whorls, up to 15 on fourth

whorl, becoming weaker, irregular and

disappearing on fourth or fifth whorl.

Axial growth lines fine but distinct on

all whorls, crossing spiral sculpture

and giving the surface the texture of

linen.

Spiral sculpture weak. Teleoconch

beginning with 5 close-set primary

spiral cords. Starting with third whorl,

one finer secondary spiral cord appears

between each pair of primary cords.

From fourth whorl on, fine tertiary

spiral threads appear at both sides of the

secondary cords. Their number increas-

ing to up to 5 on body whorl, while sec-

ondary cords become as strong as the

primary ones.

Aperture large, ovate, pointed at

both ends, white, smooth inside. Outer

lip convex, thin, simple. Inner lip

smooth, parietal callus thin, glossy,

appressed to parietal wall, not detached,

columellar folds absent.

Siphonal canal long, relatively

narrow, usually curved, about as long as

aperture. Outer side sculptured with

fine, close-set spiral cords and interca-

lated fine threads of different strength.

Periostracum thin, well-adherent,

straw-brown.

Operculum (Fig. 8) typical of genus,

corneous, red-brown, ovate, rounded

above and pointed below, shape and

size corresponding to aperture, outer

side ornamented with concentric

growth lines, with terminal nucleus.

Range and habitat: Japan, Enshu-nada

and westwards, East China Sea at 100-

300 m on sandy bottom (HIGO ET AL.,

1999: 263); Taiwan (collection RH),

Philippines, Cebu and Bohol (SPRING-

STEEN AND LEOBRERA, 1986: 177).

(Right page) Figures 9-11. Fusinus (Chryseofusus) hyphalus M. Smith, 1940, holotype F.MNH/UE

174301, Japan, Shikoku Island, Kochi Prefecture, off Tosa, 36.5 mm. 9, 10: shell; 11: spire tip.

Figures 12, 13. Fusus graciliformis Sowerby, 1880, holotype BMNH 1880.10.15.2, Japan, 52.5

mm. Figures 14, 15. Fusus sieboldi Schepman, 1891, holotype RMNH 86858, Japan, 40.0 mm

(=junior synonym of E graciliformis). Figure 16. Fusinus (Chryseofusus) westralis Hadorn and

Fraussen, 2003, holotype WAM S10876, northwest Australia, Rottnest Island, 114.4 mm. Figure

17. Fusinus (Chryseofusus) jurgeni Hadorn and Fraussen, 2002, holotype MNHN, southwest

Madagascar, 94.2 mm. Figure 18. Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911), lecto-

type ZMA, Indonesia, Molucca-Passage, 70.7 mm.

(Página derecha) Figuras 9-11. Fusinus (Chryseofusus) hyphalus M. Smith, 1940, holotipo

FLMNH/UF 174301, Japón, isla Shikoku, Kochi Prefecture, frente a Tosa, 36,5 mm. 9, 10: concha;

11: ápice de la espira. Figuras 12, 13. Fusus graciliformis Sowerby 1880, holotipo BMNH

1880.10.15.2, Japón, 52,5 mm. Figuras 14, 15. Fusus sieboldi Schepman, 1891, holotipo RMNH

86858, Japón, 40,0 mm (=sinónimo juvenil de E. graciliformis). Figura 16. Fusinus (Chryseofusus)

westralis Hadorn y Fraussen, 2003, holotipo WAM S10876, NO de Australia, isla Rottnest, 114.4

mm. Figura 17. Fusinus (Chryseofusus) jurgeni Hadorn y Fraussen, 2002, holotiyo MNHN, SO de

Madagascar, 94,2 mm. Figura 18. Fusinus (Chryseofusus) chrysodomoides (Schepman, 1911), lec-

totipo ZMA, Indonesia, estrecho de las Molucas, 70,7 mm.

160

HADORN AND CHINO: A new Fusinus (Gastropoda: Fasciolariidae) from Japan

161

Iberus, 23 (2), 2005

Comparison: F. (C.) satsumaensis spec.

nov. was often misidentified as F.

hyphalus M. Smith, 1940 by authors,

including the paper of Chryseofusus by

HADORN AND FRAUSSEN (2003). Shortly

after publication of that paper the first

author was able to locate the holotype of

E. hyphalus [FLMNH/UF 174301: type

locality: Japan, Shikoku Island, Kochi

Prefecture, off Tosa, 100 fms, M. Smith's

collection] (Figs. 9-11). FE. hyphalus is a

subjective junior synonym of F. gracili-

formis (Sowerby, 1880). This specimen is

a dead collected juvenile specimen of

36.5 x 12.5 mm and is identical in shape,

sculpture and protoconch structure but

somewhat more slender than the holo-

type of E graciliformis [BMNH

1880.10.15.2, 52.5 x 18.0 mm, dd, type

locality: Japan] (Figs. 12, 13) and F.

sieboldi (Schepman, 1891) [RMNH 86858,

40.0 x 15.2 mm, dd, type locality: Japan

(= junior synonym of F. graciliformis)]

(Figs. 14, 15). Most specimens referred

to E. hyphalus by authors belong, as far

as we are able to determine, to this new

species.

The holotype of F. hyphalus (Figs. 9-

11) differs from the new species by the

smaller size, the more convex whorls,

the more constricted suture, the reddish-

brown coloured shell, and the smaller

protoconch (diameter 0.8 mm instead of

1.1-1.2 mm). In general, F. (C.) gracili-

formis can be separated by the some-

what larger adult size, the deeper and

more pronounced subsutural concavity,

the less inflated and shorter body whorl,

the clearly smaller and differently sculp-

tured protoconch, and by often having

rather strong, broad axial ribs on the

upper whorls.

FE. (C.) westralis Hadorn and

Fraussen, 2003 (Fig. 16) differs by its

larger shell (up to 140 mm), the longer

spire, the larger number of whorls (11-

13), the more pronounced subsutural

concavity, the finer spiral sculpture con-

sisting of a larger number of fine spiral

cords and intercalated threads, and the

shorter and broader siphonal canal.

F. (C.) jurgeni Hadorn and Fraussen,

2002 (Fig. 17) can be distinguished by

162

its larger shell (up to 100 mm), the

longer spire, the more numerous (11 or

12) and more convex whorls, the more

constricted suture, the wider spire

angle, the stronger spiral sculpture, the

often reddish-brown tinged spiral

cords, the stronger and broader axial

ribs on upper teleoconch whorls, the

more rounded aperture and the broader

siphonal canal.

F. (C.) chrysodomoides (Schepman,

1911) is similar in size, but differs by

having a longer spire, a heavier shell, a

larger number of whorls, a wider spire

angle, a more pronounced axial sculp-

ture on the upper whorls, a smaller and

more rounded aperture, and a shorter

and broader siphonal canal. The lecto-

type designated by HADORN AND

FRAUSSEN (2003: 211) is figured here

(Fig. 18).

F. satsumaensis is placed in the sub-

genus Chryseofusus based on the smooth

adapical whorls, the weak, close-set,

regular spiral sculpture crossed by dis-

tinct growth lines, giving the surface the

texture of linen, the relatively short spire

and siphonal canal, the less convex

whorls with subsutural concavity, and

the simple, thin, adherent parietal

callus.

FE satsumaensis was, as misidentified

as E hyphalus, often placed in the genus

Simplicifusus Kuroda and Habe in

Kuroda, Habe and Oyama, 1971 [type

species: Simplicifusus noguchii Habe and

Masuda, 1990]. Simplicifusus was con-

cluded to be a subjective junior

synonym of Granulifusus by SNYDER

(2003: 87-88). Granulifusus has an oper-

culum which is completely different

from all other species belonging to the

subgenus Chryseofusus and all other sub-

genera in Fusinus. Species of the genus

Fusinus have an operculum with termi-

nal nucleus, corresponding to the aper-

ture in size and shape. Granulifusus has

a small, round-ovate, thin operculum,

not filling the aperture, with nucleus sit-

uated at lower outer side. For a detailed

discussion we refer to HADORN AND

FRAUSSEN (2003: 211) and to SNYDER

(2003: 88).

HADORN AND CHINO: A new Fusinus (Gastropoda: Fasciolariidae) from Japan

ACKNOWLEDGMENTS

We thank K. Way (BMNH), J. Pick-

ering (BMNH), J. Goud (RMNH), E.

Gittenberger (RMNH), and F. G.

Thompson (FLMNH/UF) for the loan

ot type material, K. Fraussen

BIBLIOGRAPHY

BIELER, R. AND PETIT, R.E., 1990. On the various

editions of Tetsuaki Kira's “Coloured Illus-

trations of the Shells of Japan” and “Shells of

the Western Pacific in Color, Vol. 1”, with an

annotated list of new names introduced.

Malacología, 32: 131-145.

HADORN, R. AND FRAUSSEN, K., 2003. The deep-

water Indo-Pacific radiation of Fusinus (Chry-

seofusus subgen. nov.) (Gastropoda: Fascio-

lariidae). Iberus, 21 (1): 207-240.

HIGO, S., CALLOMON, P., AND GOTO, Y., 1999.

Catalogue and bibliography of the marine shell-

bearing Mollusca of Japan. Elle Scientific Pub-

lications, Japan, 749 pp.

(Belgium) and M. A. Snyder (USA) for

comments, corrections and important

help to the manuscript, D. Monsecour

(Belgium) for correcting the English

text.

KIRa, T., 1962. Shells of the Western Pacific in

Color (1). Hoikusha Publishing Co., Ltd., Os-

aka, Japan, 224 pp. [for additional printings,

see BIELER AND PETIT, 1990].

SNYDER, M. A., 2003. The genera Simplicifusus

and Granulifusus (Gastropoda: Fasciolari-

idae) with the description of two new species

in Granulifusus. Journal of Conchology, 38 (1):

87-93.

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

Shells of the Philippines. Carfel Seashell Mu-

seum, Manila, 377 pp.

163

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O Sociedad Española de Malacología —__—_——— Iberus, 23 (2): 165-181, 2005

Checklist of the opisthobranchs (Mollusca: Gastropoda)

from the Chilean coast deposited in the “Colección de Flora

y Fauna Profesor Patricio Sánchez Reyes” from the

“Pontificia Universidad Católica de Chile”

Catálogo de los opistobranquios (Mollusca: Gastropoda) de la costa

Chilena depositados en la “Colección de Flora y Fauna Profesor Patricio

Sánchez Reyes” de la Pontificia Universidad Católica de Chile

María Angélica FISCHER*? and Juan Lucas CERVERA**

Recibido el 20-IV-2004. Aceptado el 27-X-2005

ABSTRACT

The specimens of opisthobranch molluscs included in the “Colección de Flora y Fauna Pro-

fesor Patricio Sánchez Reyes”, housed in the Pontificia Universidad Catolica de Chile have

been examined. As result of this review, a checklist of 18 opisthobranch species (2

sacoglossans, 1 pleurobranchid and 15 nudibranchs] is given.

To date, a total of 75 opisthobranch species, arranged in 51 genera and 34 families,

have been recorded in Chile. In this paper, the geographical distribution of the studied

species is commented on and extended in some cases.

RESUMEN

Se han revisado los ejemplares de moluscos opistobranquios presentes en la “Colección

de Flora y Fauna Profesor Patricio Sánchez Reyes”, depositada en la Pontificia Universi-

dad Católica de Chile. Como resultado de dicha revisión, se presenta una lista de 18

especies de opistobranquios [2 sacoglosos, 1 pleurobránquido y 15 nudibranquios).

Hasta el momento, en Chile se han citado un total de 76 especies de opistobranquios, dis-

tribuídas en 51 géneros y 34 familias. En el presente trabajo se comenta y se amplía en

algunos casos la distribución geográfica de las especies abordadas.

KEY WORDS: Opisthobranchia, Sacoglossa, Nudipleura, Pleurobranchoidea, Nudibranchia, Chilean coast.

PALABRAS CLAVE: Opisthobranchia, Sacoglossa, Nudipleura, Pleurobranchoidea, Nudibranchia, costa Chilena.

INTRODUCTION

Information on the opisthobranch century and originates mainly from sci-

molluscs from the Chilean coast goes entific expeditions of European origin

back to records in the nineteenth and one of North-American origin,

* Department of Cellular Animal Physiology, Nijmegen Institute for Neuroscience, Radboud University

Nijmegen, Toernooiveld 1, 6525 ED Nijmegen, The Netherlands.

** Departamento de Biología, Facultad de Ciencias del Mar y Ambientales, Universidad de Cadiz, Polígono del

Rio San Pedro, s/n. Apdo. 40, 11510 Puerto Real (Cádiz) Spain.

' Correspondence author. E-mail: m.fischerOscience.ru.nl

165

Iberus, 23 (2), 2005

Table 1. Main scientific expeditions in which opisthobranch species from the Chilean coast were

collected.

Tabla 1. Principales expediciones en las que se recolectaron opistobranquios en las costas chilenas.

Expedition

Voyage autour du monde sur la corvette La Coquille

Voyage dans l'Amérique Méridionale

Voyage de /'Astrolabe

The Vogage of HMS Challenger

Mission Scientifique du Cap Horn

Albatross

Expedition of 2 years to the west coast of South Ámerica

Swedish Antarctic Expedition

Expedition Natural History

Expedition to Juan Fernández and Eastern Island

The Lund University Chile Expedition

which also yielded general collections of

various taxonomic groups.

The results of these expeditions have

been published as checklists and

descriptions of molluscs in various jour-

nals and books, including descriptions

of about 60 species, several of which

were described for the first time (see

Table I). Other publications not specifi-

cally devoted to opisthobranch molluscs

but dealing with some Chilean species

are PÓPPIG (1829), GOULD (1852), CUN-

NINGHAM (1871), ABRAHAM (1877),

PFEFFER (1886), DALL (1889; 1909), PLATE

(1894), PiLSBRY (1895), STREBEL (1905),

POowELL (1951) and REHDER (1980).

The opisthobranchs of South-

America need a detailed revision,

because until now most species have

been described on the basis of preserved

material, sometimes with a single refer-

ence specimen, and in many cases only

as to their external morphology. More-

over, many of these descriptions are

fragmentary or ambiguous.

In recent years, interest in the opisto-

branch molluscs of South-America has

revived, and several faunistic and taxo-

nomic studies have been published,

mostly based on the study of living

material and mainly concerning

opisthobranch species of the Chilean

coast (MILLEN, SCHRÓDL, VARGAS AND

INDACOCHEA, 1994; SCHRÓDL, 19%6a; b,

166

Denmark

Sweden

Great Britain

Sweden

Country Years Author

France 1822-1825 lesson (1831)

France 1826-1833 — D'OrBicNY (1835-1846)

France 1826-1829 Quov AND GAMARD (1832)

1873-1876 BercH (1884)

France 1882-1883 RocueBrune AND MaBiLLE (1891)

U.S.A. 1885-1888 — Dan (1890)

1893-1895 BercH (18980)

1901-1903 — ObHNer (1926)

1901-1904 — Enor (1907)

Sweden 1921 ObHner (1921)

1948-1949 Marcus (1959)

1997a; b; c, 1999a; b; c, 2000a; b; e, 2001,

2003; MUÑOZ, VALDÉS AND ORTEA, 1996;

FISCHER AND ORTEA, 1997; FISCHER,

CERVERA AND ORTEA, 1997; VALDÉS AND

GOSLINER, 2001; VALDÉS, 2002; VALDÉS

AND MUNIAÍN, 2002; FISCHER AND

CERVERA, 2005; FISCHER, VAN DER VELDE

AND ROUBOS, 2005; SCHRÓDL, ALARCÓN,

BEDRIÑANA, BRAVO, BUSTAMANTE, CAR-

VALHO, FÓRSTERRA, GALLARDO, HAUS-

SERMANN AND SALMEN, 2005).

Some of the species reported for the

Chilean coast have also been recorded

from the Antarctic coast (WAGELE, 1990;

1995; CATTANEO-VIETI, 1991; GARCÍA,

TRONCOSO, GARCÍA-GÓMEZ AND

CERVERA, 1993) and from Argentina

(MUNIAÍN, ORTEA AND RODRÍGUEZ, 1991;

MUNIAÍN, VALDÉS AND ORTEA, 1996;

SCHRÓDL, 1996b) and Peru (D'ORBIGNY,

1837; MILLEN ET AL., 1994; SCHRÓDL,

1996b).

In 1960 Professor Patricio Sánchez

Reyes, from the Pontificia Universidad

Católica de Chile, in Santiago de Chile,

founded the Room for Systematics (“la

Sala de Sistemática”) where biological

material collected from several field

expeditions was deposited. After his

decease in 1999, and in recognition of

his exhaustive work, this systematic

room was named “Colección de Flora y

Fauna Profesor Patricio Sánchez Re-

yes”.

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

In this paper we study the opistho-

branch material gathered from the

Chilean coast between 1960 and 1971,

present in this institution. It has not

been studied until now, due to a lack of

Chilean specialists.

MATERIAL AND METHODS

Specimens were identified by study-

ing their external morphology as well as

their anatomy. Results were compared

with descriptions in the literature. Infor-

mation about the collection localities of

the material along the Chilean coast and

RESULTS

Opisthobranchs of the Chilean

coast are distributed over five higher

taxa according to the new classifica-

tions: Cephalaspidea, Aplysiomorpha,

Sacoglossa, Tylodinoidea and Nudi-

the geographic distribution of the

species has been included.

The classification follows recent and

comprehensive studies of the

phylogeny of various opisthobranch

groups (JENSEN, 1996, 1997; WAGELE

AND WILLAN, 2000; SCHRÓDL, WAÁGELE,

AND WILLAN, 2001; “VALDÉS AND

GOSLINER, 2001; VALDÉS, 2002).

According to modern insights in phylo-

genetic classification, taxa higher than

“family” have been not assigned to a

taxonomic category (DE QUEIROZ AND

GAUTHIER, 1994) but the hierarchical

structure of the classification has been

maintained.

pleura (see Table II). Among our mate-

rial, we only identified species

belonging to Sacoglossa and Nudi-

pleura, as will be described in the

following list.

OPISTHOBRANCHIA

SACOGLOSSA Von lIhering, 1876

Family PLAKOBRANCHIIDAE Gray, 1840

Genus Elysia Risso, 1818

Elysia hedgpethi Marcus, 1961

Material: Los Molles (32* 15' S; 71” 30” W), four specimens (N* SSUC-729). Duao, X región (34* 55"

S; 71” 33 W), one specimen (N* SSUC-612).

Previous records from Chile: SCHRÓDL

(1996a): Seno Otway (53* 07" S; 71? 22 W),

Fuerte Bulnes (53* 39 S; 70? 56” W), Bay

of Mansa (537 32' S; 70? 55 W) in the South

of Chile. The present material is the first

record from the central coast of Chile.

General distribution: Also known

from Tomales Bay (California) (MARCUS,

1961) and from Vancouver Island

(MILLEN, 1980) to Bahía de los Angeles

and Bahía de San Quintín (Baja Califor-

nia, Mexico) (BEHRENS, 1991).

Family HERMAEIDAE H. Adams and A. Adams, 1854

Genus Aplysiopsis Deshayes, 1839/1853 non Bergh, 1872

Aplysiopsis brattstrómi Marcus, 1959

Material: La Portada, Bay of Antofagasta (2>” 38' S; 70? 31” W), one specimen (N* SSUC-2608).

Previous records from Chile: Marcus

(1959): Antofagasta (type locality); SCHRÓDL

(1996a) recorded this species from the cen-

tral and southern Chilean coast: Bay of

167

Iberus, 23 (2), 2005

Coliumo (36* 32* S; 72% 57 W), Fuerte

Bulnes, extending its geographic distribu-

tion significantly towards the south.

General distribution: Also known

from Comodoro Rivadavia, Patagonia

Argentina (MUNIAÍN, 1997).

NUDIPLEURA Waágele and Willan, 2000

PLEUROBRANCHOIDEA Férussac, 1822

Family PLEUROBRANCHIDAE Férussac, 1822

Genus Berthella Blainville, 1824

Berthella platei (Bergh, 1898)

Material: Los Molles, two specimens (N* SSUC-183). Poza Arrecifes, Los Molles, one specimen, (N*

SSUC-470). El Tabo (33* 27” S; 71” 38” W), one specimen (N* SSUC-32).

Previous records from Chile: BERGH

(1898a): Quirina; ODHNER (1926): Gulf of

Reloncaví (41? 44” S; 72% 55 W), Punta

Pelluco (41? 30' S; 72% 53 W), North

from Bay of Quellín (41? 51 S; 72% 55'

W>, South from Gulf of Ancud (42* 26' S;

72% 59 W); BurdwoodBank (53* 45” S;

61” 10” W); MARCUS (1959): Bay of Cal-

buco (41* 56' S; 73% 08” W), North from

Gulf of Ancud, between Tres Cruces and

Punta Piedras (41? 50” S; 73% 28” W);

SCHRÓDL (1999): Island Picton, Beagle

Channel, Lenca, Bay of Reloncaví;

SCHRÓDL ET AL. (2005): Comau Fjord

(429 05" /429 305; 729 37"/727 21 W:

General distribution: This species has

not been reported outside the Chilean

coast.

NUDIBRANCHIA Blainville, 1814

Family POLYCERIDAE Alder and Hancock, 1845

Genus Thecacera Fleming, 1828

Thecacera darwini Pruvot-Fol, 1950

Material: Coliumo, Los Morros (36* 29 S; 72* 58” W), three specimens (N” SSUC-1102), three specimens

(N* SSUC-1109). Caleta Leandro, Tumbes (36* 37' S; 73 07” W), two specimens (N* SSUC-1538).

Previous records from Chile: PRUVOT-

FoL (1950): Bay of Naranja, Islote Hoste

(55* 10” S; 69% 20” W); Marcus (1959):

Chiloé-North, Punta Ahui (41* 49” S; 73

51" W), Islote de Chonos (45 00' S; 74

00” W); SCHRÓDL (199%6a; 2003): Juan

López (23” 30' S; 70? 32” W), Bay Inglesa

(27 07 570% 53 WI) Los blornos:

Coquimbo (29* 38” S; 71? 29 W), Pichi-

dangui (32* 08” S; 71* 33” W), Bay of

Coliumo; SCHRÓDL (2003): Seno Ventis-

quero (44? 30" S; 72% 35" W); SCHRÓDL ET

AL. (2005): Comau FEjord (42* 05'/42* 30'

S; 72% 37'/72* 21* W); FiscHER' ETLAR:

(2005): Bay of Antofagasta (23” 29” S; 70*

25” W), Bay of La Herradura (29* 58” S;

71 22” W), Bay Tongoy, Coquimbo (309

15" S; 719 30” W),-Bay Horcón? Walls

paraíso (33* S; 71” W); Las Cruces, Val-

paraíso (33%:29":S;1 7121381 WbBa

Hueihue, Ancud, Chiloé (41* 54” S; 731

31” W); Bay Putemún, Castro, Chiloé

(429 25' S; 73% 45' W).

General distribution: This species has

not been recorded outside the Chilean

coast.

Family Chromodorididae Bergh, 1891

Genus Cadlina Bergh, 1878

Cadlina sparsa (Odhner, 1922)

168

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

Material: Coliumo, Los Morros, one specimen, 29 mm in length (N* SSUC-1102).

Previous records from Chile: ODHNER

(1922): Archipelago of Juan Fernández

(537 37 5,78 99 - W)), MARCUS(1959):

Bay of Quetalmahue (41” 50” 40” S; 73

57" 10” W); SCHRÓDL (1996a; 2003): Bay

of Coliumo; SCHRÓDL ET AL. (2005):

Comau Fjord (42? 05'/42* 30” S; 72*

DT pza zac WN)»

General distribution: This species is

also known from California (JAECKLE,

1984; BEHRENS, 1991) and Argentina

(SCHRÓDL, 2000b, 2003).

Genus Tyrinna Bergh, 1898

Tyrinna nobilis Bergh, 1898

Material: Iquique (20? 12” S; 70? 10” W), two specimens (N* SSUC-3334).

Previous records from Chile: BERGH

(1898a), ODHNER (1921): Juan Fernández

Island; Pajargo Island (probably Pájaros

Island); MARCUS (1959): Strait of Chacao

(41? 46" S; 73 45” W), Punta de Tenaún

(429 20' 50” S; 73 22” 00” W) and Bay of

Calbuco; SCHRÓDL (1996a): Bay of

Coliumo, Faro Corona (41* 50' S; 73* 50'

W), Seno Ventisquero (44? 30" 5; 72? 35”

W”; SCHRÓDL AND MILLEN (2001): Beach

Posiriqueros (267 125,705.59 W), Los

Hornos, Pichidangui (32” 08” S; 71? 33'

W), Bay of Coliumo, Lenca (41* 40' S;

72% 40” W); SCHRÓDL (2003): Strait of

Magellan, Chilean Patagonia to Los

Hornos (northern Chile) and Juan Fer-

nández Islands; SCHRÓDLE ET AL. (2005):

Comau Fjord (42 05'/42* 30" S; 729

SI UZAZAINN):

General distribution: This species is

also known from several localities in

Argentinian Patagonia (MUNIAÍN ET AL.,

1996; SCHRÓDL, 1996a, 2003; SCHRÓDL

AND MILLEN, 2001).

Family DORIDIDAE Rafinesque, 1815

Genus Doris Linnaeus, 1758

Doris fontainei d'Orbigny, 1837

Material: Mehuín (39* 26' S; 73” 12” W), one specimen (N* SSUC-2129). Poza Verde, Coliumo, one

specimen (N* SSUC-1141). Punta Moquehua, Caleta Mansa, five specimens (No. SSUC-5313). Chiloé

(43? 00” S; 74? 00' W), one specimen (N* SSUC-5411).

Previous records from Chile: D'OR-

BIGNY (1835-1846): Southern Montemar,

northern of Valparaíso (32” 57' 24” S; 71*

33" 25” W); ODHNER (1926): Coquimbo,

Puerto Montt (41? 28” S; 72* 57” W) and

Dichato (36" 33" S; 72% 56” W); MARCUS

(1959): Los Molles; SCHRÓDL (1996a):

Bay of Coliumo, Seno Ventisquero.

SCHRÓDL (2003): from Chilean Patagonia

to Arica (northernmost of Chile) (18? 25”

S; 70% 16" W); SCHRÓDL ET AL. (2005):

Comau Fjord (42 05”/42* 30” S; 72"

IIESIZ ZEN):

General distribution: Also known

from Argentina: Northern Argentina

(ODHNER, 1926); and Patagonian shores

(SCHRÓDL, 1996a; VALDÉS AND MUNIAÍN,

2002). Moreover, SCHRÓDL (2003) sup-

ports its presence in the Peruvian coast.

Remarks: Doris fontainei has been

recently reported from the coasts of

Argentina and Chile under different

names (MUNIAÍN ET AL., 1991; SCHRÓDL,

19%6a; 1997b; 2000c). But, according to

VALDÉS AND MUNIAÍN (2002), the correct

specific name for this species should be

D. fontainei. Recently. SCHRÓDL (2003)

transferred this species to the genus

Archidoris based on the absence of an

acrembolic penis as described for the

169

Iberus, 23 (2), 2005

Table H. Opisthobranch species recorded from the Chilean coast. Taxa higher to Family level are

not assigned to any category, as explained in the text. Species included in our material are marked

with an asterisk.

Higher Taxa Families Genera Species

CEPHALASPIDEA S./' Acteonidae Actaeon A. delicatus Dall, 1889

A. curtulus Dall, 1889

A.vagabundus Rochebrune and Mabille, 1891

Diaphanidae Diaphana D. paessleri Strebel, 1905

Toledonia [. limnaeaeformis (Smith, 1879)

[. perplexa Dall, 1902

Scaphandridae Scaphander S. interruptus Dall, 1890

Aglajidae Aglaja A. maculata (d'Orbigny, 1837)

APLYSIOMORPHA Aplysiidae Aplysia A. parvula Bergh, 1898

Dolabella D. auricularia (Lightfoot, 1786)

| Dolabriferidae Dolabrifera D. dolabrifera (Rang, 1828)

SACOGLOSSA

Oxynoacea Juliidae Julia J. exquisita Gould, 1862

Berthelinia B. pseudochloris Kay, 1964

Plakobranchacea Plakobranchidae Elysia *E. hedgpethi Marcus, 1961

Hermaeidae Aplysiopsis *A. bratistrómi Marcus, 1959

Limapontidae Limapontia Limapontia sp.

Ercolania E. evelinae Marcus, 1959

TYLODINOIDEA Umbraculidae Umbraculum U. umbraculum (Lightfoot, 1786)

NUDIPLEURA

Pleurobranchoidea ? Pleurobranchidae Berthella *B. platei (Bergh, 1898)

Berthellina B. citrina (Rippel and Leuckart, 1828)

Pleurobranchaea P maculata (Quoy and Gaimard, 1832)

Nudibranchia

Anthobranchia

Doridoidea

“Phanerobranchia” * Onchidorididae Acanthodoris A. folklandica Eliot, 1907

Goniodorididae Okenia 0. angelensis Lance, 1966

0. luna Millen, Schródl, Vargas and Indacochea, 1994

Áncula A. fuegiensis Odhner, 1926

Corambidae * Corambe C. lucea Marcus, 1959

Polyceridae Kaloplocamus K. maculatus (Bergh, 1898)

Holoplocamus H. papposus Odhner, 1926

Polycera P priva Marcus, 1959

P alabe Collier and Farmer, 1964

Thecacera *T. darwini Pruvot-Fol, 1950

Cryptobranchia Chromodorididae > Cadlina *(. sparsa (Odhner, 1922)

Iyrinna *T. nobilis Bergh, 1898

Dorididae Doris / Neodoris *D. fontaineid'Orbigny, 1837

*D. kerguelenensis (Bergh, 1884)

Doris sp. 1

Ñ. claurina Marcus, 1959 incertae sedis

Discodorididae * Geitodoris 6. patagonica Odhner, 1926

Baptodoris *B. peruviana (d'Orbigny, 1837)

Gargamella *6. inmaculata Bergh 1894

Diaulula D. hispida (d'Orbigny, 1837)

*D. punctuolata (d'Orbigny, 1837)

*D. variolata (d'Orbigny, 1837)

Rostanga *R. pulchra MacFarland, 1905

170

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

Tabla II. Opistobranquios citados en la costa chilena. No se asignan taxones a categorias superiores a

familia, tal y como se explica en el texto. Las especies incluidas en nuestro material se indican con un

Asterisco.

Higher Taxa Families Genera Species

Dexiarchia ”

Cladobranchia

Dedronotoidea Tritoniidae Tritonia T. challengeriana Bergh, 1884

T. vorax (Odhner, 1926)

*T. odhneri Marcus, 1959

Iritonia sp. |

Dendronotoidae Dentronotus Dendronotus sp.

Dotoidae Doto *D. uva Marcus, 1955

Hancockidae Hankockia H. schoeferti Schródl, 1999

Phylliroidae Phylliroe P bucephala Péron and Lesueur, 1810

Arminoidea Arminidae Armina A. cuvieri (d'Orbigny, 1837)

Proctonotidae Janolus $ J. rebeccae Schródl, 1996

: J. chilensis Fischer, Cervera and Ortea, 1997

Aeolidoidea Flabellinidae Flabellina E folklandica Eliot, 1907

Flabellina sp. 1

Flabellina sp. 2

Tergipedidae Cuthona €. georgiana (Ptefter in Martens and Pfeffer, 1886)

C. pusilla (Bergh, 1898)

€. odhneri Marcus, 1959

€. valentini (Eliot, 1907)

Cuthona sp. 1

Cuthona sp. 2

Eubranchidae Eubranchus E. agrius Marcus, 1959

Eubranchus sp. |

Eubranchus sp. 2

Tergipedidae lergipes [. valentini (Eliot, 1907)

Facelinidae Facelina E cyanella (Couthovy in Gould, 1852) insertae sedis

Phidiana *P lottini (Lesson, 1831)

P patagonica (d'Orbigny, 1837)

Glaucidae Glaucus 6. atlanticus Forster, 1777

Aeolidiidae Aeolidia *A. papillosa (Linnaeus, 1761)

A. collaris Odhner, 1922

Fionidae Fiona *E pinnata (Eschscholtz, 1831)

1 In view of the analyses of MikKELSEN (1996; 2002) and the molecular analyses of GRANDE, TEMPLADO, CERVERA AND ZARDOYA (2004;

b) it seems clear that this taxon ¡s paraphyletic, although we used it in the present study because the phylogenetic relations

between the different groups are still not definitively established.

2 There are three species of Pleurobranchoidea reported from Chile, but only Berthella plateiis reported in this study, and is the only

one occurring in both the Magellan and Peruvian provinces. Two other pleurobranchid species, Berthellina citrina (Rijppell and

Leuckart, 1828) and Pleurobranchaea maculata (Quoy and Gaimard, 1823) have been recorded from the Chilean coast, but

they are restricted to Eoster Island and the Juan Fernández Islands, respectively. However, both records should be considered

very doubtful according to MARCUS AND GOSLINER (1984), ScurónL (19960) and Cervera, GOSLINER AND GARCÍA-GÓMEZ (1999).

2 Recent studies support the non-monophyly of Phanerobranchia (THoLLesson, 1999; WoLischelo AnD WaGELE, 1999; WoLtschElo-

Lenceuin6, Boore, Brown AND WaGeLE, 2001; VaLnés, 2002; WiceLe, VONNEMANN AND WGELE, 2003; GRANDE, TEMPLADO, CERVERA

AND ZARDOYA, 20040, b; FAHEY AND GOSLINER, 2004) although a deep phylogenetic analysis of this taxon is still lacking.

4 ScuróoL Ano WáceLe (2001) hove recently clorified the systematic position of the Family Corambidae and redescribe the Chilean

species Corambe lucea Marcus, 1959.

171

Iberus, 23 (2), 2005

Table Il. Continuation.

Tabla II. Continuación.

5 In Chile the Chromodorididae are represented by two genera, each one with one species: Cadlina sparsa and Tyrinna nobilis.

Another nominal chromodoridid species, Chromodoris juvenca Bergh, 1898 was attributed to the genus Cadlina by Marcus

(1959) and SchróoL (19960). However, this species is currently considered as a junior synonym of 7. nobilis (SCHRÓDL,

2000b, 2003; SchróoL Ano MiLLen, 2001). Thus, the only valid species of Cadlina in Chile appears to be C. sparsa.

6 After the recent revision and phylogenetic study of the dorid criptobranchs VaLoés (2002) considered the Families Platydorididae,

Baptodorididae and Diaululidae as synonyms of the Family Discodorididae. Consequently, the genera Baptodoris, Gargamella

and Digulula must be included within the family Discodorididae.

7 Dexiarchia Schródl, Wigele and Willan, 2001 is a Nudibranchia clade recently erected, which includes the Doridoxoidea, Dendrono-

toidea, Aeolidoidea and Arminoidea.

8 There are two valid species at the Chilean coast, Janolus rebeccae Schródl, 1996 and J. chilensis Fischer, Cervera and Ortea, 1997.

A posterior comparative study of both species let us see that they are different species (Fischer, Schródl and Cervera, unpub-

lished data).

genus Doris. In the genus Archidoris the

prostate gland is reduced or absent,

whereas in Doris fontainei it is well

developed, which agrees with the genus

Doris in general. Nevertheless, in this

study, we consider that based on most

of the morphological characters it

belongs to the genus Doris.

Doris kerguelenensis (Bergh, 1884)

Material: Island Nueva, Magallanes (55? 15' S; 66” 32* W), one specimen (N* SSUC-5155).

Previous records from Chile: BERGH

(1884): Puerto Otway (Chilean Patagonia);

BERGH (1898a): Bay Tumbes and Punta

Arenas; ODHNER (1926): Punta Arenas,

Puerto Sofía, River Condor, Tierra del

Fuego, Puerto Harris; MARCUS, 1985: 61?

15 S; 55 05 W; SCHRÓDL (1996a): Bay Pos-

esión (52” 13' S; 69* 17" W).

Distribution: Also known from Royal

Sound and Morbihan Bay, Kerguelen

Islands (BERGH, 1884; VICENTE, 1974);

Almirante Buck, Antartic Territory

(VAYSSIERE, 1917); South Georgias, South

from Falklands Islands, Ushuaia

(Argentina) (ODHNER, 1926); MCMURDO

Sound, Antarctic Territory (ODHNER,

1934); Davis Sea, Antarctic Territory

(MINICHEV, 1972); Macquarie and Heard

Islands (Burn, 1973); Scotia Sea (GARCÍA

ET AL., 1993); Wedell Sea, Antarctic

Peninsula and South Georgias (WAGELE,

1990); New Caledonia (VALDÉS, 2001).

Remarks: VALDÉS (2002) has argued that

Austrodoris is synonymous with Doris.

Accordingly, the species Austrodoris ker-

guelenensis should be denominated Doris

kerguelenensis and be included within the

Dorididae, as was also suggested by

SCHRÓDL (1996a). Recently, however,

SCHRÓDL (2003) referred to this species as

Austrodoris kerguelenensis overlooking the

statement of VALDÉS (2002).

Family DISCODORIDIDAE Bergh, 1891

Genus Baptodoris Bergh, 1384

Baptodoris peruviana (d'Orbigny, 1837)

Material: Iquique (20? 12' S; 70? 10' W), three specimens (N” SSUC-3332). La Portada, Antofagasta,

three specimens (N* SSUC-2607). South-East of the Mejillones Peninsula (23 20' S; 70% 34 W), twelve

specimens, (N” SSUC-3032). Los Molles, three specimens (N* SSUC-183).

EX2

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

Previous records from Chile: Isla

Pájaros (BERGH, 1898); Valparaíso,

(DaLL, 1909).

General distribution: This species is

also known from Peru: San Lorenzo

(D'ORBIGNY, 1837) to Pucasana

(SCHRÓDL, 1996a). Its record from Galá-

pagos Islands (PILSBRY AND VANATTA,

1902) has been considered dubious (Fis-

CHER AND CERVERA, 2005) and not con-

sidered here.

Remarks: The nominal species Doris

peruviana d'Orbigny, 1837 was trans-

ferred to the genus Platydoris by

Schródl (2003), considering P. punc-

tatella Bergh, 1898 as a junior synonym.

DORGAN, VALDÉS AND GOSLINER (2002)

were not able to assign a generic name

to P. punctatella on the basis of the orig-

inal description by BERGH (1898), but

using the photograph of a living

animal in SCHRÓDL (1996a), they stated

that it is not a Platydoris species.

FISCHER AND CERVERA (2005) agree

with this view. Thus, since the type

material of D. peruviana cannot be

located, these authors compare speci-

mens of a doridoidean species from

Iquique to Los Molles (Chilean coast)

with the holotype of P. punctatella, and

conclude that both are conspecific and,

according to the radular teeth features,

belong to the genus Baptodoris rather

than Platydoris.

Genus Gargamella Bergh, 1894

Gargamella inmaculata Bergh, 1894

Material: Cabo Metalqui, Chiloé (41? 50” 30” 5; 73” 28” 30” W), two specimens (N” 55UC-3899).

Previous records from Chile: ODHNER

(1926): Ultima Esperanza, Tierra del

Fuego (53% 00” S; 69% 20” W); MARCUS

(1959): Cabo de San Antonio (53% 55' S;

70% 52” W), Cabo Delgado (50* 06” S; 74?

55 W), Gulf of Ancud, between Quenu

and Islote de Calbuco (41* 48” 50” S, 732

09” 40” W); SCHRÓDL (1996a): Seno

Ubway Queule (89235773 13 W),

Bay of Coliumo. This is a common

species from the southern Chilean coast.

General distribution: Also recorded

from Northern Argentina (BERGH, 1894;

ODHNER, 1926), as well as the Argen-

tinian Patagonia and the Burdwood

Bank (ODHNER, 1926; SCHRÓDL, 2003).

The records from Peru, Southern Africa

and New Zealand by ZAGAL AND

HERMOSILLA (2001) are considered

erroneous (SCHRÓDL, 2003).

Remarks: SCHRÓDL (1996a)

included this species erroneously in

the family Kentrodorididae, but

recently VALDÉS (2002) transferred

it again to the family Discodoridi-

dae.

Genus Diaulula Bergh, 1878

Diaulula punctuolata (d'Orbigny, 1837)

Material: Los Molles, two specimens (No. SSUC-183). Mehuín, one specimen (N* SSUC-2129). Punta

Moquehua, Caleta Mansa, eleven specimens (N* SSUC-5278).

Previous records from Chile: D'OR-

BIGNY (1835-1846): Valparaíso (33* 02” S;

71 38” W); BERGH (1898a): Bay Talc-

ahuano (36 40” S; 73 03 W), South Bay

Tumbes; ODHNER (1926): Melinka,

Guaitecas Islands; SCHRÓDL (1996a): Bay

Coliumo, Faro Corona, Seno Ventis-

quero; VALDÉS AND GOSLINER (2001):

Península Lacuay, Chiloé Island; VALDÉS

AND MUNIAÍN (2002): Lota, Peninsula

Lacuay, Chiloé Island and ABRAHAM

(1877): Strait of Magallanes.

General distribution: Apart from

Chile, this species is also known from

173

Iberus, 23 (2), 2005

Callao, Peru (DaLL, 1909); Argentinian

Patagonia (SCHRÓDL, 1996a, 2003) and

Falklands Islands (ELIOT, 1907a).

Remarks: The generic status of the

nominal species Doris punctuolata d'Or-

bigny, 1837 and Anisodoris marmorata

Bergh, 1898 (non Archidoris marmorata

Bergh, 1891) has received much atten-

tion (VALDÉS AND GOSLINER, 2001;

VALDÉS AND MUNIAÍN, 2002; SCHRÓDL,

2003). Thus the former of this species

was transferred to Diaulula recently by

the above authors. On the other hand,

the second species is transferred to

Peltodoris Bergh, 1880 by VALDÉS AND

MUNIAIN (2002).

Diaulula variolata (A'Orbigny, 1837)

Material: El Tabo, V región (33” 27' S; 71” 38" W), four specimens (N* SSUC-32). El Tabo, Valparaíso,

one specimen (N” SSUC-53). Iquique, one specimen (N* SSUC-3333).

Previous records from Chile: BERGH,

(1898a): Bay of Coquimbo (29* 57” S; 71*

22" W); MARCUS (1959): Bay of San Vicente

(36? 44' S; 73? 11” W), South-Eastern Punta

Gualpén, northern Chile (36? 44” 54” 5; 732

11" 02” W); SCHRÓDL (1996a): Bay Inglesa,

Los Hornos, Guanaqueros (30* 10' S; 71

26" W), Bay of Coliumo; SCHRÓDL (1997):

Bay Inglesa (27? 07' S: 70 53" W); SCHRÓDL

(2003): Arica (18? 25' S; 70” 16' W); VALDÉS

AND MUNIAÍN (2002): Coquimbo and Lota.

General distribution: This species has

not been recorded outside Chile.

Remarks: VALDÉS AND MUNIAIN

(2002) consider the nominal species

Doris variolata d'Orbigny, 1837 within

Peltodoris, but distinct from the nominal

species Anisodoris marmorata Bergh,

1898. SCHRÓDL (2003) demonstrated that

both species belong to the genus Diaul-

ula, based on the presence of caryophyl-

lidia on the notum, as well as that both

are conspecific. The present study

extends the known distribution of D.

variolata much more northwards, up to

Iquique.

Genus Rostanga Bergh, 1879

Rostanga pulchra MacFarland, 1905

Material: Totoralillo, two specimens (N” SSUC-938).

Previous records from Chile: MARCUS

(1959): Playa Brava, Chiloé (41? 51* 35”

S; 73 49 20” W), Punta El Morro,

Ancud (41? 52 42” S; 73” 50' 46” W);

SCHRÓDL (1996a): Coliumo, Queule,

Bay of Ancud, Faro Corona. The

present study extends the geographical

range towards the northern Chilean

coast.

General distribution: Also recorded from

Alaska (LEE AND FOSTER, 1985) to Mexico

(LANCE, 1966; MARCUS AND MARCUS,

1970). It has been also collected from

Argentina (MARCUS AND MARCUS, 1969).

Family "TRITONIDAE Lamarck, 1809

Genus Tritonia Cuvier, 1797

Tritonia odhneri Marcus, 1959

Material: Mehuín (39* 26' S; 73” 12” W), one specimen (N”. SSUC-2129). Duao, one specimen (N?

SSUC-577). Ensenada de Tumbes (36? 37' S; 73% 07” W), one specimen (N* SSUC-1381). Corral (32*

52" S; 737 37" W), two specimens (N* SSUC-3807).

174

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

Previous records from Chile: Punta Tenaún,

Gulf of Ancud (42* 20” S; 73% 22” W):

MARCUS (1959); Bay of Coliumo, Queule

(36? 32* S; 72? 57" W): SCHRÓDL, 199%6a.

General distribution: This species has

also been recorded from northern

Argentina to southern Brazil (SCHRÓDL,

2003).

Family DOTOIDAE Gray, 1853

Genus Doto Oken, 1815

Doto uva Marcus, 1955

Material: Punta Blanca, Tocopilla (22? 04” S; 70? 12” W), one specimen (N* SSUC-3038).

Previous records from Chile: MARCUS

(1959): Channel of Calbuco, Gulf of Ancud

in Chile; SCHRÓDL (1997a): Bay of Coliumo.

SCHRÓDL ET AL. (2005): Comau Fjord (42?

WDRERR305 722 37/12) 21 W). l'he pre=

sent study extends the geographical range

towards the northern Chilean coast.

General distribution: This species is

also known from Sao Paulo, Brazil

(MARCUS, 1955, 1957, 1959).

Family FACELINIDAE Bergh, 1889

Genus Phidiana Gray, 1850

Phidiana lottini (Lesson, 1831)

Material: Duao, six specimens (N* SSUC-610). Caleta Leandro, Tumbes, one specimen (N* SSUC-

1534). Yerbas Buenas, Tumbes, four specimens (N” SSUC-1665). El Morro, Tomé (36* 37" S; 72*

57" W), four specimens (N* SSUC-1936). Mehuín, five specimens (N* SSUC-2129). Punta Moque-

hua, Caleta Mansa, eight specimens (N* SSUC-5279).

Previous records from Chile: Lesson

(1831): San Vicente; PLATE (1894):

Coquimbo; BERGH (1898): Coquimbo,

Cavancha, Iquique; MARCUS (1959):

Punta Liles in Gulf of San Vicente (36*

43936" 9, 73-08" 10” W), South-East

Prat Aa (AI SO 10 S; 737 51 20%

W”); SCHRÓDL (199%6a, 2003): Los Hornos

(29” 38" S; 71” 29 W), Queule, Lenca (41*

40" S; 72% 40” W), Bay of Ancud (41* 52*

S; 739 55” W), Bay of Coliumo; SCHRÓDL

ET AL. (2005): Comau Fjord (42? 05'/422

30 S; 72? 37'/72* 21” W). Records from

the Strait of Magellan are considered

erroneous (SCHRÓDL, 2003).

General distribution: Apart from

Chile, this species is also known from

Callao, Peru (D'ORBIGNY, 1835-1846).

Family AEOLIDIIDAE d'Orbigny, 1834

Genus Aeolidia Cuvier, 1798

Aeolidía papillosa (Linnaeus, 1761)

Material: Los Molles, two specimens (N” SSUC-183). Los Morros, Coliumo, two specimens (N*

SSUC-1113). Puerto Inglés, Tumbes (36* 37" S; 73% 07" W), six specimens (N* S5SUC-1619). Traiguén,

Tumbes, four specimens (N* SSUC-1743). El Morro, Tomé, one specimen (N* SSUC-1936). Punta

Moquehua, Caleta Mansa, six specimens (N* SSUC-5279).

Previous records from Chile: BERGH

(1898a): In the Magellanic province, Seno

Almirantazgo; MARCUS (1959): North-West

from Islote Quenu (41? 49 15” S, 73* 10'

15” W), East from Punta Corona (41? 47"

12” S, 73% 52* 23” W), North from Islote

Cochinos (41? 49” 25” S, 73” 48' 58” W),

North from Gulf of Ancud between Tres

143

Iberus, 23 (2), 2005

Cruces and Punta Piedras, Bay Tumbes

and Punta María in Agua Fresca, Strait of

Magellan (53* 22 S, 70% 57" W); SCHRÓDL

(1996a): Faro Corona; SCHRODL (2003): Bay

of Coliumo. With the present study the

distribution has been markedly extended

northwards, up to Los Molles.

General distribution: Aeolidia papillosa

is a worldwide species in temperate

waters (SCHRÓDL, 2003; THOMPSON AND

BROWN, 1984). In southern America it

has been also recorded from Falkland

Islands (BERGH, 1898) and Argentinian

Patagonia (SCHRÓDL, 1996a).

Family FIONIDAE Alder and Hancock, 1851

Genus Fiona Alder and Hancock, 1851

Fiona pinnata (Eschscholtz, 1831)

Material: Mehuín, Valdivia, two specimens (N* SSUC-2129).

Previous records from Chile: BERGH

(1898a): Juan Fernández Islands and

Tumbes, Talcahuano (Chile Central).

General distribution: Fiona pinnata is

a cosmopolitan species (SCHMEKEL

DISCUSSION

To date the number of known

opisthobranchs in Chile amounts to 75

species distributed in 51 genera and 34

families, all of them recorded in Table II.

The present study provides records of 18

opisthobranch species: 2 sacoglossan, 1

pleurobranchid and 16 nudibranchs, all

previously known from Chile.

The Nudibranchia form the most

diverse clade of opisthobranchs in

Chile, consisting of 57 species. Among

them Thecacera darwini is a well-known

and very common species along the

entire Chilean coast, just as Phidiana

lottini and Diaulula variolata.

Chilean opisthobranch species

descriptions too poor to warrant reiden-

tification, or currently considered as

nomen dubium or nomen nudum, are not

included in Table II. These are: Doris del-

icata Abraham, 1877, Doris tomentosa

Cuvier, 1804 (sensu Abraham, 1877),

Doris amarilla Póppig, 1829, Doris magel-

lanica Cunningham, 1871, Doris chilensis

Abraham, 1877 and Acanthodoris vathelei

Rochebrune and Mabille, 1891. The

holotype of the latter species has proba-

bly been mislaid (V. Heros, personal

communication).

176

AND PORTMANN, 1982; GOSLINER,

1987); recorded off northern Peru

(D'ORBIGNY, 1835-46) and from Lobos

Islands, northern Peru (SCHRÓDL,

199%6a; 2003).

Eleven species reported in the litera-

ture are still unidentified: one sacoglos-

san (Limapontia sp., SCHRÓDL, 1999a) and

ten nudibranchs (Doris sp., SCHRÓDL,

199%6a; Dendronotus sp., SCHRÓDL, 1996a;

Tritonia sp1., SCHRÓDL, 2003; Flabellina

sp. 1, SCHRÓDL, 19%6a, Flabellina sp. 2,

SCHRÓDL, 19%6a; Eubranchus sp. 1,

SCHRÓDL, 19%6a; Eubranchus sp. 2,

SCHRÓDL, 1996a, Cuthona sp. 1, SCHRÓDL,

1996a; Cuthona sp. 2, SCHRÓDL, 1996a;

Tergipedidae sp. 1, SCHRÓDL, 2003).

On the other hand, the opistho-

branch collection of Couthouy reported

on by GOULD (1852) has been lost. Due

to the poor description of this material,

both species Doris luteola Couthouy in

Gould, 1852 nomen dubium, and Doris

plumulata Couthouy in Gould, 1852

nomen dubium could not be included in

Table II.

ACKNOWLEDGEMENTS

This publication is a tribute to Pro-

fessor Patricio Sanchez in memoriam,

who founded the Systematic Room of

the “Pontificia Universidad Católica de

FISCHER AND CERVERA: An opistobranch checklist from the Chilean coast

Chile” that made available the material

for this study. We thank the Curator of

the same institution Mr. Patricio Zabala,

Professors Jesús Ortea (University of

Oviedo) and Cecilia Osorio (University

of Chile). Also to Dr. Virgine Heros

(MNHN, París) for kindly providing

information about some type material.

Finally, we thank Professors Heike

Waágele and Dietrich Hofmann (Univer-

sity of Bochum), for the supplied facili-

ties in their laboratories, and Professor

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genus Cadlina (Gastropoda: Opisthobranchia)

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SCHRÓDL, M., 2000c. Taxonomic revision of the

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SCHRÓDL, M., ALARCÓN, M. A., BEDRIÑANA, L.

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W., 2003. Towards a phylogeny of the Opistho- W. AND WAGELE, H., 2001. The phylogeny of

branchia. In Lydeard, C. and Lindberg, D. Nudibranchia (Opisthobranchia, Gastropoda,

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181

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ÁviLA, S. P, SANTOS, A. C., PENTEADO, A. M., RODRIGUES, A. M., QUINTINO,

[. AND MACHADO, M. I. The molluscs of the intertidal algal turf in the Azores. /be-

rus, 23 (1): 67-76.

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Figure 2. Transect performed at Poga do Pano (Lajes do Pico, Pico island) and vertical distribution

of rocky shore organisms. HWST: mean high water level at spring tides; LWST: mean low water

level at spring tides.

Figura 2. Transecto realizado en Poga do Pano (Lajes do Pico, isla de Pico) y distribución vertical de

organismos de costa rocosa. HWST: nivel medio superior del agua en mareas de primavera: LWST: nivel

medio inferior del agua en mareas de primavera.

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vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se den las sinonimias

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Dendrodoris limbata (Cuvier, 1804)

Sinonimias

Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Localidad tipo: Marsella].

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275.

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Fretter, V. y Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.

Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W. EF.

(Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166.

Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas.

Miscelánea Zoolgica, 3 (5): 21-51.

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INSTRUCTIONS TO AUTHORS

e Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. Papers are

manuscripts 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: Serge Gofas, Editor de Publicaciones,

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+ 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

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* 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. This must include a concise but informative title, with mention of family of higher taxon when appropri-

atte, 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. Yhese should content the rest of the paper, divided into sections under short headings. Whenever

possible 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.

e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical

Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its authori-

ty including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN FULL, includ-

ing the periodical, 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. If 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

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» 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 CAP-

ITALS. The references need not be cited when the author and date are given only as authority for a taxonomic name.

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other unpublished 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.

Miscelánea Zoológica, 3 (5): 21-51.

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Figure 1. Neodoris carvi. A: animal crawling; B: rinophore; C: gills.

If abbreviations are to be used in illustrations, group them alphabetically after the Legends for Figures section.

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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 20093:

Socio numerario (en España): 40 euros

(en Europa) 40 euros

(fuera de Europa): 48 euros

Socio estudiante (en España): 23 euros

(en el extranjero): 29 euros

Socio Familiar: (sin recepcion de revista) 4 euros

Socio Protector: (mínimo) 48 euros

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(fuera de Europa): 54 euros

INSCRIPCIÓN: 6 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-

llos 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.

A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio-

nal postal money orders in euros sent to the Treasurer). Members 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.

SMITHSONIAN INSTIT

Ll 00071

ÍNDICE

Iberus 23 (2) 2005

REDFERN, C. AND ROLÁN, E. A new species of Lodderena (Gastropoda: Skeneidae) from the

Bahamas

Una nueva especie de Lodderena (Gastropoda: Skeneidae) de las Bahamas ...... A 1-6

MARTÍNEZ-ORTÍ, A. Y ROBLES, E Los caenogasterópodos terrestres (Mollusca, Orthogastropoda)

de la Comunidad Valenciana (España)

The land caenogastropods (Mollusca, Orthogastropoda) of the “Comunidad Valenciana”

o O O 7-24

TRONCOSO, J. S., MOREIRA, J. AND URGORRI, V. Soft-bottom mollusc assemblages in the Ría de

Ares-Betanzos (Galicia, NW Spain)

Asociaciones malacológicas de substratos blandos de la Ría de Ares-Betanzos ( Gia NO

VE A a A E O II 25-38

ESPINOSA, F., Fa, D. A. Y OCAÑAa, T. M. J. Estado de la especie amenazada Patella ferruginea

Gmelin, 1791 (Gastropoda: Patellidae) en la bahía de Algeciras y Gibraltar

Status of the endangered limpet Patella ferruginea Gmelin, 1791 (Gastropoda: Patellidae) in

the Algeciras bay and Gibraltar ........ O O nono 39-46

BOYER, E The discovery of a radula in a Dentimargo species and its taxonomic implications

Descubrimiento de la rádula en una especie de Dentimargo y sus implicaciones taxonómicas

47-52

BOYER, E. AND ROLÁN, E. About a sibling species of Mitrella minor (Scacchi, 1836)

Sobre una especie gemela de Mitrella minor (Scacchi, 1836) ......ccncnncooconnnnnenrannnnnnos 53-67

HERNÁNDEZ, J. M. AND BOYER, E Notes on the columbellid fauna from the infralittoral and cir-

calittoral levels of the Canary Islands

Notas sobre los columbelidos del infralitoral y circalitoral de Canarias .....umommmmmmm. 69-93

PELORCE, J. Er BOYER, E La famille Columbellidae (Gastropoda: Muricoidea) dans Pinfralittoral

de la Péninsule du Cap Vert (Sénégal)

The family Columbellidae (Gastropoda: Muricoidea) in the infralittoral of the Peninsula of

RN A A A A A caco 95-118

ROLÁN, E. Columbellidae (Gastropoda, Neogastropoda) of the gulf of Guinea with the descrip-

tion of eight new species

Columbellidae (Gastropoda, Neogastropoda) del Golfo de Guinea con la descripción de ocho

A A A A A A io 119-156

HADORN, R. AND CHINO, M. A new Fusinus (Gastropoda: Fasciolariidae) from Japan

Un nuevo Eusinus (Gastropoda: Fasciolariidae) de Japón .......mcmionimnmmommmmomm*m*m*”.*.”«.s 157-163

FISCHER, M2 A. AND CERVERA, J. L. Checklist of the opisthobranchs (Mollusca: Gastropoda)

from the Chilean coast deposited in the “Colección de Flora y Fauna Profesor Patricio

Sánchez Reyes” from the “Pontificia Universidad Católica de Chile”

Catálogo de los opistobranquios (Mollusca: Gastropoda) de la costa Chilena depositados en la

“Colección de Flora y Fauna Profesor Patricio Sánchez Reyes” de la Pontificia Universidad

E A A A 165-181

ISSN 0212-3010