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 (ASSOCIATE EDITORS)

Francisco Javier Conde de Saro

Benjamín Gómez Moliner

Angel Antonio Luque del Villar

Emilio Rolán Mosquera

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

Rúdiger Bieler

Sigurd v. Boletzky

Jose Castillejo Murillo

Karl Edlinger

Antonio M. de Frias Martins

José Carlos García Gómez

Gonzalo Giribet de Sebastián

Edmund Gittenberger

Angel Guerra Sierra

Gerhard Haszprunar

Yuri 1. Kantor

María Yolanda Manga González

Jordi Martinell Callico

Ron K. 0'Dor

Takashi Okutani

Marco Oliverio

Pablo E. Penchaszadeh

Winston F. Ponder

Carlos Enrique Prieto Sierra

M* de los Ángeles Ramos Sánchez

Francisco Javier Rocha Valdés

Paul G. Rodhouse

Joandoménec Ros ¡ Aragones

María Carmen Salas Casanovas

Gerhard Steiner

Victoriano Urgorri Carrasco

Anders Warén

PORTADA DE lberus

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, Banyuls-sur-Mer, 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 Universitát Wien, Viena, Austria

Universidad de Santiago de Compostela, Santiago de Compostela, España

Swedish Museum of Natural History, Estocolmo, Suecia

lberus 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

Vol. 22 (2) Oviedo, diciembre 2004

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, P.O. 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, 22 (2): 1-8, 2004

Observations on the reproductive period of the freshwater

mussel Potomida littoralís (Unionidae)

Observaciones sobre el periodo reproductor del mejillón de agua

dulce Potomida littoralis (Unionidae)

Karl-Otto NAGEL*

Recibido el 17-X11-2003. Aceptado el 5-11-2004

ABSTRACT

Data on the occurrence of gravid Potomida littoralis are presented from museum collec-

tions, field samplings and published literature. Gravid animals and released glochidia are

recorded to occur between March and October. The smallest gravid animals were four or

five years old. The length of the brooding period apparently follows a latitudinal cline.

Potomida littoralis is most likely a tachytictic consecutive brooder with unsynchronised

broods.

RESUMEN

Se aportan datos sobre la presencia de ejemplares grávidos de Potomida littorales, a par-

tir de colecciones de museos, muestreos de campo y bibliografía. Los reproductores y la

liberación de gloquidios se produce entre marzo y octubre. Los reproductores menores

eran animales de cuatro o cinco años de edad. La duración del periodo de cría sigue, en

apariencia, una tendencia latitudinal. Potomida littoralis se reproduce en primavera y en

verano, con puestas no sincronizadas.

KEY WORDS: Reproduction, Mollusca, Potomida littoralis, Unionidae

PALABRAS CLAVE: Reproducción, moluscos, Potomida littoralis, Unionidae.

INTRODUCTION

Freshwater mussels (Unionoida) are

faced with a serious decline in both their

numbers and distribution in many parts

of the world (BOGAN, 1993). Reproduc-

tion biology and ecology of juvenile

mussels are key factors that determine

the survival abilities of freshwater

mussel populations (BAUER, 2001).

Several species occurring in Europe that

are more seriously threatened have

received considerable attention in the

past decades (Margaritifera margaritifera:

BAUER, 1988; YOUNG, COSGROVE AND

HAsTIE, 2001; Unio crassus: HOCHWALD,

2001; Pseudunio auricularius (syn: Mar-

garitifera auricularia): ARAUJO AND

RAMOS, 2001). The biology and ecology

of most other European species remain

insufficiently known.

The occurrence of Potomida littoralis

(Cuvier, 1798) (authorship of this taxon

according to FALKNER, BANK AND VON

PROSCHWITZ, 2001 and FALKNER, RIPKEN

AND FALKNER, 2002) continues to be

* pod hrádzou 3011, SK - 900 01 Modra (Harmónia), Slovenska Republika - Slovakia. konagelOgmx.de

Iberus, 22 (2), 2004

Figure 1. Orientation map for localities from Table 1. F: France; E: Spain; P: Portugal; TR: Turkey;

SYR: Syria; IL: Israel.

Figura 1. Mapa de las localidades de la Tabla 1. E: Francia; E: España; P: Portugal; TR: Turquía; SYR:

Siria; IL: Israel.

widespread in Western Europe (France,

Spain, Portugal), however for other

parts of the species” distribution range

(Greece, Turkey, Near and Middle East,

Northern Africa) no information is cur-

rently available. Data on the general bi-

ology of this species date back to the be-

ginning of the last century. Haas (1917)

discovered that the species has 4 marsu-

pial gills (= tetrageny). He examined the

anatomy of a single male specimen from

France and 11 males and 3 sterile (=

non-gravid) females of Rhombunio rothi

Bourguignat, 1863 from Jaffa /Tel Aviv,

Israel (Haas, 1920). Later, he considered

this species to be a synonym of Potomida

littoralis (HAas, 1969). It might be noted

that the material of R. rothi studied by

Haas should still be present in the

Senckenberg Museum, Frankfurt/M.,

Germany. Lot n. SNF 000811 contains 25

specimens preserved in alcohol, none of

which are gravid. The label reads:

“Rhombunio rothi (Bgt) - Jaffa — J. Aha-

roni S. 1913”. Haas (1924) later exam-

ined the condition of the marsupial gills

of animals collected in Spain. He proba-

bly determined the sex of these speci-

mens by histological studies but this is

not stated explicitly in his paper. His

findings however suggest that Potomida

littoralis is dioeceous.

This paper is a report on the occur-

rence of gravid specimens of P. littoralis

in the river Ognon (eastern France) and

on the presence of gravid mussels and

glochidia of this species based on pub-

lished data and museum material. The

data is culled from three distinct areas

within the species” distribution range.

MATERIAL AND METHODS

Sampling localities and collection

dates are given in Table I. All localities

are shown on an orientation map (Fig.

1). Data on gravid specimens were

obtained from examination of the fol-

lowing collections:

1. Collection Kinzelbach, University

of Rostock, Germany (Prof. Dr. R.

Kinzelbach). Reference numbers in

Table II: VO-78/

NAGEL: Reproductive period of the freshwater mussel Potomida littoralis (Unionidae)

Table I. List of localities.

Tabla I. Lista de localidades.

No. Sampling locality

France

1 Cireyles-Bellevaux, 19 km NNE of Besancon

2 Marnay, ca 20 km WNW of Besancon

3 canal near Montfrin, 20 km WSW of Avignon

4 Clerey, 13 km SE of Troyes

5 le Guétin, about 1,5 km upstream of confluence with Loire,

9 km SW of Nevers

6 Brehemont, 27 km WNW of Tours

7 Candes-St. Martin, 50 km WNW of Tours

g Chauvigny, 9 km E of Poitiers

Spain and Portugal

Lake of Bañolas/Banyoles, ca 16 km NNW of Geron/Girona

10 Acequia (ditch) de San Adrián, ca 44 km ESE of Logroño

11 Zaragoza

12 Sástago, app. 56 km SE of Zaragoza

13 near Ruidera, ca 26 km SSE of Tomelloso

14 Goyán/Goián, 12 km upstream of the river mouth at

La Guardia/A Guarda (at 3 different but very close sites)

15 near Vila Nova de Cerveira (opposite of Goyán),

ca 12 km upstream of the river mouth

16 Tuy/Tui, 27 km upstream of the river mouth

17 Salvaterra do Miño, 39 km upstream of the river mouth

lurkey, Syria and Israel |

18 near Samanda_i, ca 25 km SW of Hatay (Antakya)

19 (1) confluence of the two outlets of the Lake of Homs

(Buhairat al-Qattina or B. al-Hims) and (2) backwater

20 atbridge of road Homs-lartus

21 stagnant backwater in the al-Ghab ENE of Ain al-Kurum

22 canal at crossing of the road Ain al-Kurum to

Sagalibiya (Sqelbié)

23 Lake Tiberias (Kinneret)

2. Collection of recent freshwater

invertebrates at the Museo Nacional de

Ciencias Naturales, Madrid, Spain. Ref-

erence numbers in Table 1l: MNCN FW-

3. Senckenberg-Museum, Frankfurt

am Main, Germany. Reference numbers

in Table II: SNF-

Gills of living or preserved speci-

mens were inspected to see whether they

were filled. The developmental status of

the content of the gills was not always

determined and is not reported here.

River Or River Departmement,

stream basin Province, country

Ognon Rhóne Haute-Saóne

Gardon Rhóne Gard

Seine Seine Aube

Allier Loire Nievre

Indre Loire Indre-et Loire

Vienne Loire Indre-et Loire

Vienne Loire Vienne

endorheic Girona

Ebro Navarra

Canal Imperial de Aragón Ebro laragoza

Ebro Ebro laragoza

Lagunas de Ruidera Guadiana Albacete and Ciudad Real

Mino Mino Pontevedra

Minho Minho Viana do Castelo, Portugal

Mino Miño Pontevedra

Miño Miño Pontevedra

Orontes Orontes Hatay, Turkey

Orontes Orontes Syria

Jordan Israel

Live adult mussels were gently opened

by hand to approximately 1 cm. The soft

parts of most museum specimens were

already removed from the shell and sep-

arately stored. It is important to remem-

ber that in Potomida littoralis the inner

and outer gills become filled with eggs.

In this case they are swollen and non-

transparent. Filled marsupia are cream

white to light brown in live animals,

while unfilled gills are semi-transparent,

thin and greyish colour.

Iberus, 22 (2), 2004

Mussels from the river Ognon at

Cirey-les-Bellevaux were repeatedly

inspected by hand while wading on the

riverbanks. The maximum depth

sampled was 1.5 m. Sampling dates are

given in Table III. The mussels were

painted with numbers for individual

recognition and measured to the nearest

1/10 mm with a veneer calliper.

RESULTS

Table II reports the findings from

collections and field observations. All

gravid specimens were encountered

from April to October.

The smallest gravid specimen was

39.3 mm in length (June 6, 1990, river

Miño at Goyán). The smallest gravid

mussel from the Ognon measured 41.0

mm (Table II). In both cases, three,

perhaps four, darkly coloured growth

lines were evident on the outside of the

shell. This means that the animals were

in their fourth (or fifth) spring /summer

period when they reproduced.

Table III reports the findings of re-

peated observations of marked individ-

ual mussels in one locality in eastern

France. Mussel with length 55.3 mm

was not found gravid in the year 2000

and became gravid in 2001. Since the an-

imal was among the largest in that pop-

ulation it was most likely also sexually

mature in the first year of observation.

This individual may have paused from

reproduction in 2000 or may have repro-

duced exceptionally late in that year.

Excluding animals that are less than

39.3 mm, which was the size of the

smallest gravid individual, the number

of mussels found gravid in the river

Ognon was 33%. Similar values were

found for the pooled samples from the

river Orontes (33%) and the Ebro at

Sástago on July 18, 1916 (30%).

DISCUSSION

Little is known about the brood

behaviour of unionoidean bivalves. In

many cases inferences from circumstan-

tial observations and the lack of distinc-

tion between individual and population

behaviour have prevented us from

detecting the actual patterns until

recently (HEARD, 1998). Specific infor-

mation on the reproductive period of

Potomida littoralis is scattered. The only

published records of gravid animals are

from Haas (1917, 1920) and PRASHAD

(1919).

Haas (1917) conducted regular sam-

plings between July 18 and September 5,

1916 in the river Ebro at Sástago. He

found individuals carrying eggs and

larvae at the beginning of this period

and he recognized freshly metamor-

phosed mussels on the last date.

However, Haas obviously failed to

account for the presence of gravid

animals in a sample he had taken the

year before (September 7-8, 1915; Haas

1916). A small proportion of those

mussels, still preserved in the Sencken-

berg Museum, are gravid (Table II). For

the other samples carried out in 1916

(Sástago: August 15, 22, 29, September 5;

Gallur, ca 42 km NW of Zaragoza: Sep-

tember 3) Haas did not give the number

of P. littoralis collected. It remains

unclear whether he inspected them for

gravidity at all.

PRASHAD (1919) made observations

on 3 specimens of Unio (Rhombunio)

semirugatus collected in October, 1912, in

Lake Tiberias (Sea of Galilee). The

species name is considered by HAAS

(1969) to be a synonym of Potomida lit-

toralis. One mussel was gravid in the

sample of Prashad. Apparently, it con-

tained glochidia only in the outer gills.

In fact, Prashad regarded the outer gills

as the only structure having marsupial

function (= ectobranchy). Apart from

the presence of larvae, Prashad based

his identification of marsupial gills on

the spacing of the gill septa. HAAS

(1924) expressed doubts about the valid-

ity of this interpretation since he identi-

fied four marsupial gills in Rhombunio

rothi by histological examinations

(Haas, 1920, 1924). Considering all

observations reported in Table II, it

might be concluded that the specimen

of Prashad was collected at the end of

NAGEL: Reproductive period of the freshwater mussel Potomida littoralis (Unionidae)

Table II. Observations on the brooding behaviour of Potomida littoralis. The number of gravid

mussels when present is given in parenthesis after the sample size.

Live glochidia: data from

ARAUJO, BRAGADO AND RAMOS (2000). Abbreviations. ?: number not specified.

Tabla IT. Observaciones sobre el comportamiento de cría de Potomida littoralis. El número de ejempla-

res reproductores se da entre paréntesis tras el tamaño de muestra. Gloquidios vivos: datos tomados de

Araujo, Bragado y Ramos (2000). Abreviaturas. ?: número sin especificar.

Month Live Gravid Sample size No. of sampling

glochidia— mussels locality (from Table 1)

January No!

February No data No 2 18

March Ves? No 21 11

April Yes No 6 14

No 9 1

Yes 10 (3) 1

May Yes

June Yes Yes 3 (1) 15

Yes 36 (9) 1

Yes “several” (4) 2

Yes 6 (2) 1

No 20 1)

No 18 6

July Yes No 33 4

Yes 1 (1) 10

No 7 14

Yes "several hundred” (?) 12, at 0-0.5m

sub-sample: 47 (14) depth

No 5 5

No 3 6

No 4 7

No 2 8

August Yes Yes “many” (2) at0-0.5 m, “some” 12

(all ?) at 5-6 m depth

No 6 21

Yes more than 1 (1) 22

Yes 8 (4) 23

Yes a 24 and 25

No 18

Yes 7 at0-0.5 m, ? (all a 12

at 6 m depth

Yes 9 (2) 20

No 2 14

September Yes Yes sub-sample: 35 (5) 12

No 42 12

No 14 13

No 1 19

No 10 9

No 5 14

October Ves? Yes 3 (1) 23

No 17 3

No 19 17

No 1 16

November No data No 2 14

No 1 18

December No

four dates in 1998; 2 from 6.3.1997; * until 9.10,1997

Date

26.2.2001

IIS

16.4.1991

23.4.2000

29.4.2001

6.6.1990

10/11.6.2000

23.6.2000

30.6.1991

AS9l

11.7.1996

MO9O

18.17.1916

21.1.1985

23.1.1985

24.1.1985

26.17.1985

2.8.1916

3.8.19178

3.8.1978

5.8.1918

8/10.8.1978

4.8.1988

8.8.1916

23.8.1978

28.8.1990

1/8.9.1915

8.9.1915

8.9.2000

11.9.2002

15.9.1914

18.9.1990

October 1912

5.10.1989

71.10.1988

24.10.1989

6.11.1990

Reference

MNCN FW-681

Haas, 1916; SMF 003052

MNCN FW-691

this study, see Table Il

this study, see Table ll

MNCN FW-640

this study, see Table 11

this study

this study, see Table l1l

Nagel and Badino, 2001

MNCN FW-1157

MNCN FW-645

Haas, 1917, 1920; SMF 003054;

this study

Nesemann and Nagel, 1989

Neserann and Nagel, 1989

Haas, 1917; Haas, 1920

VO-78/12

vO-78/12a

VO-78/20

VO-78/28; VO-78/29

MNCN FW-1295

Haas, 1917

VO-78/49

MNCN FW-652

SMF 323165; this study; (Haas, 1916)

Haas, 1916; SMF 003053

MNCN FW-1500

MNCN FW-1543

SMF 003049

MNCN FW-654

Prashad, 1919

Nagel and Badino, 2001

MNCN FW-498

MNCN FW-664

MNCN FW-665

Iberus, 22 (2), 2004

Table III. Observations of the reproductive status of Potomida littoralis in the river Ognon near

Cirey-les-Bellevaux, France. From a total of 37 individually marked animals only those are listed

that could be inspected at least two times. The remainding ones were not gravid on April 23, 2000

(1 specimen) or on June 10 - 11, 2000 (12 specimens). Length of specimens as measured on day

of first record. No intensive sampling on June 23, 2000.

Tabla 111. Observaciones del estatus reproductor de Potomida littoralis en el río Ognon cerca de Cirey-

les-Bellevaux, Francia. De un total de 37 especímenes marcados sólo se incluyen aquellos que se observa-

ron al menos dos veces. Los restantes no estaban en estado reproductor el 23 de abril de 2000 (1 ejem-

plar) o el 10-11 de junio de 2000 (12 ejemplares). Se da la longitud tomada el día de la primera obser-

vación. No se realizó un muestreo intenso el 23 de junio de 2000.

Gravid

2000

April 23 June 10/11 June 23

Length

(mm) 2001

21.8

30.2

39.0

39.4

40.0 ; i

41.0 : +

41.8 +

44.6 ep ap

45.6 : 2

46.6 o

46.1 mi

46.8 ;

April 29

Length Gravid

(mm) 2000 2001

April 23 June 10/11 June 23 April 29

47.0 - a

41.1 d+

48.1 ;

48.2 +

49,3 + +

50,2 .

915 - +

dl : ap

54.] : ; ;

30 : = 5 +

58.1 : a

65.9 | z 1

Explanation of symbols: + gravid, - not gravid, F dead, no entry: not found.

the gravidity period and had already

expelled the content of the inner gills

thus giving the appearance of an ecto-

branchous state.

Based on the observations of HAAS

(1920, 1924), NAGEL (1988, p. 3, fig. 1)

hypothesized that Potomida littoralis is a

summer breeder, i.e. tachytictic like the

European species of the genus Unio. The

observations from the river Ognon

(Table II) fit well into this pattern.

Recently, ARAUJO, BRAGADO AND RAMOS

(2000) reported on live glochidia (valves

with soft body) in drift net samples

from early March to beginning of

October, 1997, in the Canal Imperial de

Aragón that takes water from the Ebro

(sampling localities near the villages of

Grisén and Alagón, between Gallur and

Zaragoza, see Table II). These data

confirm the tachitixis but also give evi-

dence of a very long reproductive

period.

No gravid mussels were found in

France in the 64 specimens sampled in

July and October. Several explanations

are possible for this, for example, lati-

tude. It may be safe to assume that the

reproductive season for P. littoralis is

abbreviated at higher latitudes where

periods with water temperatures

favourable for the growth of food parti-

cles (bacteria, algae) are short. There is

evidence that at least some unionoid

species allocate only surplus energy to

reproduction (HOCHWALD, 2001 and ref-

erences therein). Alternative explana-

tions are population specific and would

point to a high parasite load and natural

or anthropogenic habitat disturbances.

All these factors can induce individuals

or entire populations to suppress repro-

duction. This latter phenomenon was

previously described for some species

(Margaritifera margaritifera: BAUER, 1987;

Ross, 1992; Unio crassus: HOCHWALD,

NAGEL: Reproductive period of the freshwater mussel Potomida littoralis (Unionidae)

2001; Hyridella depressa: WALKER, BYRNE,

HICKEY AND ROPER, 2001).

The apparently very long reproduc-

tive period in the southern locations

makes consecutive breeding (more than

one brood in the reproductive period) of

the species quite probable, but presently

there is no direct evidence for this. This

form of reproductive behaviour,

however, seems to be widespread

among unioniod mussels (HEARD, 1998).

A mean of 30% gravid mussels was

found in the samples from the Ognon,

one sample from the Ebro and the

pooled Orontes samples (Table Il.

Assuming a sex ratio of 1:1 would mean

that one-third of the females did not

actively participate in reproduction at

the times of sampling. This pattern indi-

cates unsynchronised breeding within a

population.

From the above it is concluded that

Potomida littoralis is a tachytictic brooder

BIBLIOGRAPHY

ARAUJO, R. BRAGADO, D. AND RAMOS, M. A.,

2000. Occurrence of glochidia of the endan-

gered Margaritifera auricularia (Spengler, 1793)

and other mussel species (Bivalvia:

Unionoida) in drift and on fishes in an ancient

channel of the Ebro River, Spain. Archiv fiir

Hydrobiologie, 148: 147-160.

ARAUJO, R. AND RAMOS, M. A. , 2001. Life-his-

tory data on the virtually unknown Margar-

itifera auricularia. In Bauer, G. and Wáchtler,

K. (Eds. ): Ecology and Evolutionary Biology

of the Freshwater Mussels Unionoida, Eco-

logical Studies, 145: 143-152. Springer Verlag

Berlin, Heidelberg.

BAUER, G., 1987. Reproductive strategy of the

freshwater pearl mussel Margaritifera mar-

garitifera. Journal of Animal Ecology, 56: 691-

704.

BAUER, G., 1988. Threats to the freshwater pearl

mussel Margaritifera margaritifera L. in Cen-

tral Europe. Biological Conservation, 45: 239-

23.

BAUER, G. , 2001. Environmental relationships

of naiads: threats, impact on the ecosystem,

indicator function. In Bauer, G. and Wachtler,

K. (Eds. ): Ecology and Evolutionary Biology

of the Freshwater Mussels Unionoida, Eco-

logical Studies, 145: 311-315. Springer Verlag,

Berlin Heidelberg.

with probably consecutive unsynchro-

nised broods. The reproductive period

can span from March to October. More

data are needed to determine if the

length of the reproductive period

follows a latitudinal cline.

ACKNOWLEDGEMENTS

R. Araujo and R. Kinzelbach pro-

vided information on collection material

and E.M. Nagel assisted the field sam-

plings. The manuscript benefited from

comments provided by a referee and by

R. Araujo and J. Plant, the latter who

edited my English. To all of them 1

remain most grateful. The study of the

collection in Madrid was supported by

the Museo Nacional de Ciencias Natu-

rales and by a grant from the European

Commission's Human Potential Pro-

gramme.

BOGAN, A. E. , 1993. Freshwater bivalve ex-

tinctions (Mollusca: Unionoida): a search for

causes. American Zoologíst, 33: 599-609.

FALKNER, G. , BANK, R. A. AND VON

PROSCHwItz, T. , 2001. CLECOM-PRO-

JECT. Check list of the non-marine species-

group taxa of the states of Northern, At-

lantic and Central Europe (CLECOM J].. -

Heldia, 4: 1-76.

FALKNER, G. , RIPKEN, TH. E. J. AND FALKNER,

M. 2002. Mollusques continentaux de France.

Liste de Référence annotee et Bibliographie.

Patrimoines naturelles, 52: 350 pp.

HAas, F. , 1916. Sobre una concha fluvial in-

teresante (“Margaritana auricularia” Spglr. )

y su existencia en España. Boletin de la Sociedad

Aragonesa de Ciencias Naturales, 15:33-44, plate

TL.

Haas, F., 1917. Estudios sobre las náyades del

Ebro. Boletin de la Sociedad Aragonesa de Cien-

cias Naturales, 16:71-82.

Haas, F., 1920. Die Gattung Rhombunio, ihre

Anatomie und Stellung im System. Sencken-

bergiana, 2: 70-80.

Haas, F., 1924. Anatomische Untersuchungen

an europáischen Najaden. 1. Archiv fir Mol-

luskenkunde, 56: 66-82, plates IV + V.

Haas, F., 1969. Superfamilia Unionacea. Das

Tierreich, 88: i-x, 1-663. De Gruyter, Berlin

Iberus, 22 (2), 2004

HEARD, W. H., 1998. Brooding patterns in fresh-

water mussels. Malacological Review, suppl. 7

(Bivalvia 1): 105-121.

HOCHWwALD, $. , 2001. Plasticity of life-history

traits in Unio crassus. In Bauer, G. and

Wáchtler, K. (Eds. ): Ecology and Evolu-

tionary Biology of the Freshwater Mussels

Unionoida, Ecological Studies, 145: 127-141.

Springer Verlag, Berlin Heidelberg.

NAGEL, K. -O. , 1988. Anatomische, morphologí-

sche und biochemische Untersuchungen zur Tax-

onomie und Systematik der europáischen

Unionacea (Mollusca: Bivalvia). Ph. D. thesis,

University of Kassel (Germany) . ii + 101 pp.

NAGEL, K. -O. AND BADINO, G. , 2001. Popula-

tion genetics and systematics of European

Unionoidea. In Bauer, G. and Wáchtler, K.

(Eds. ): Ecology and Evolutionary Biology of

the Freshwater Mussels Unionoida, Ecologi-

cal Studies, 145: 51-80. Springer Verlag, Berlin

Heidelberg.

NESEMANN, H. AND NAGEL, K. -0. , 1989. Die

Flufímuscheln (Bivalvia: Unionacea) im

Einzugsgebiet der Loire (Zentralfrankreich)

- eine erste Bestandserfassung. Mitteilungen

der deutschen malakozoologischen Gesellschaft,

44/45: 1-15.

PRASHAD, B.,, 1919. On the generic position of

some Asiatic Unionidae. Records of the Indian

Museum, 16: 403-411.

Ross, H. C.G., 1992. The reproductive biology

of the freshwater pearl mussel Margaritifera

margaritifera (L. ) in Co Donegal. The Irish

Naturalists” Journal, 24: 43-50.

WALKER, K. F., BYRNE, M. , HICKEY, C. W. AND

ROPER, D.S. 2001. Freshwater Mussels (Hyri-

idae) of Australasia. In Bauer, G. and

Waáchtler, K. (Eds. ): Ecology and Evolu-

tionary Biology of the Freshwater Mussels

Unionoida, Ecological Studies, 145: 5-31.

Springer Verlag, Berlin Heidelberg.

YOUNG, M. R.,, COSGROVE, P. J. AND HASTIE, L.

C. 2001. The extent of, and causes for, the de-

cline of a highly threatened naiad: Margari-

tifera margaritifera. In Bauer, G. and Wáchtler,

K. (Eds. ): Ecology and Evolutionary Biology

of the Freshwater Mussels Unionoida, Eco-

logical Studies, 145: 337-357. Springer Verlag,

Berlin Heidelberg.

O Sociedad Española de Malacología Iberus, 22 (2): 9-17, 2004

La malacofauna de la Sierra de Alcaraz (Albacete, España)

The molluscan fauna of the Alcaraz mountains (Albacete, Spain)

Alberto MARTÍNEZ-ORTÍ*, María Teresa APARICIO** y Fernando ROBLES***

Recibido el 3-IV-2003. Aceptado el 23-I11-2004

RESUMEN

Se realiza el primer estudio de la malacofauna continental de la sierra de Alcaraz (Alba-

cete, España). Los datos previos son escasos, dispersos y de muy diversos autores. Se han

recogido muestras en 29 localidades del área de estudio y se han hallado un total de 50

especies de gasterópodos, de las cuales 47 son terrestres y 3 dulceacuícolas. Se citan por

primera vez 22 especies en la provincia de Albacete y 10 en la Comunidad de Castilla-La

Mancha.

ABSTRACT

The first complete study of non-marine molluscs of the Alcaraz mountains (Albacete, Spain)

has been carried out. Previous data are scarce, disperse and from many diverse authors.

Samples have been collected in 29 localities of the study area and a total of 50 species of

gastropods have been found, 47 corresponding to land species and 3 to freshwater spe-

cies. Twenty-wo species are recorded for the first time for the province of Albacete and 10

for the “Comunidad de Castilla-La Mancha”.

PALABRAS CLAVE: moluscos, terrestres, dulceacuícolas, Alcaraz, Albacete, España.

KEY WORDS: molluscs, land, freshwater, Alcaraz, Albacete, Spain.

INTRODUCCIÓN

La sierra de Alcaraz, con una exten-

sión de 156.000 hectáreas, se encuentra

en el SW de la provincia de Albacete y

enlaza con las sierras andaluzas de

Cazorla y Segura. Junto a la cabecera del

río Mundo forma parte del “Parque

Natural de la Sierra de Alcaraz y Alto

del Segura” de la Comunidad Autó-

noma de Castilla-La Mancha. Hasta

ahora se han realizado algunos inventa-

rios de fauna (ANDÚJAR TOMÁS Y GÓMEZ

DE (GUEVARA, 1985; VIDAL-ABARCA,

SUÁREZ, MILLÁN, GÓMEZ, ORTEGA,

VELASCO Y RAMÍREZ-DÍíAz, 1991) y de

flora (HERRANZ Y GÓMEZ CAMPO, 1986;

GUERRERA MONTES, ROs EsPíN, HERAS

IBÁÑEZ, GARCÍA ZAMORA Y JIMÉNEZ

MARTÍNEZ, 1989) de esta sierra, pero se

poseen escasos datos sobre su malaco-

fauna.

Esta falta de información se extiende

a la mayor parte de la provincia de

Albacete, en la cual predominan las citas

aisladas de diversos autores, destacando

* Museu Valencia d'Historia Natural. Passeig de la Petxina, 15. 46008 Valencia (España).

alberto.martinezOPuv.es.

** Museo Nacional de Ciencias Naturales de Madrid. C.S.I.C. c/ José Gutiérrez Abascal, 2. 28006 Madrid.

menta2a2mncn.csic.es.

“Instituto “Cavanilles”de Biodiversidad y Biología Evolutiva y Departamento de Geología de la Universitat

de Valencia. 46100 Burjassot (Valencia). roblesfPuv.es.

Iberus, 22 (2), 2004

Río Alcaraz 8

QRO

Reolid

Río Salobre de ich

ESPAÑA 12 En Ñ

Ario

10 Dr ae

Paterna. o

Bogarra

29 “Riópar

pio yndS

27.

Nacimiento

Figura 1. A. Situación geográfica del área de estudio (gris: provincia de Albacete; negro: sierra de

Alcaraz). B. Localización de las localidades de muestreo en la sierra de Alcaraz (Albacete).

Figure 1. A. Geograpbical situation of the study area (grey: Albacete province; black: Alcaraz moun-

tains). B. Location of the sampled localities in the Alcaraz mountains (Albacete).

entre ellas la descripción de una nueva

especie, Helix semipicta, por HIDALGO

(1870), cuya localidad tipo es Alcaraz.

En los últimos 30 años han aumen-

tado las referencias sobre la malaco-

fauna de la provincia de Albacete.

GASULL (1975) cita varias especies, y

RAMOS Y APARICIO (1985) estudian los

gasterópodos terrestres y dulceacuícolas

de las Lagunas de Ruidera, encontrando

un total de 19 especies. ROBLES,

BORREDA Y COLLADO (1991) estudian los

moluscos continentales de la región de

Almansa, y citan 43 especies. BORREDA Y

COLLADO (1991) y BORREDA, COLLADO,

BLASCO Y ESPÍN (1991) estudian los pul-

monados desnudos de la provincia,

citando un total de 11 especies. Los

moluscos dulceacuícolas también han

sido objeto de estudio, sobre todo en el

río Júcar: JIMÉNEZ Y MARTÍNEZ-LÓPEZ

(1988) determinan 9 especies, MARTÍ-

NEZ-LÓPEZ, PUJANTE Y TAPIA (1993) siete,

y RUEDA, HERNÁNDEZ.Y TAPIA (2001)

nueve especies. Además, VIDAL-ABARCA

ET AL. (1991) realizan un estudio limno-

lógico de la cuenca del río Mundo,

donde citan 11 especies.

En su Tesis Doctoral, PUENTE (1994)

realiza una revisión de los Helicoidea de

la Península Ibérica y cita algunas espe-

cies procedentes del área de estudio.

BORREDA (1996) caracteriza las babosas

del este de la Península Ibérica y cita

varias especies en la provincia de Alba-

cete, algunas de ellas en el área de

muestreo del presente trabajo. Por

último, GÓMEZ Y DANTART (1996) dan a

conocer nuevas localidades de especies

del Orden Orthurethra recolectadas en

Castilla-La Mancha y, algunas de ellas,

en la provincia de Albacete.

En este trabajo se da a conocer el

primer inventario detallado de las espe-

cies de moluscos terrestres y dulceacuí-

colas de la sierra de Alcaraz, lo que

permite ampliar el conocimiento de este

filo en la provincia de Albacete y en la

Comunidad Autónoma de Castilla-La

Mancha.

MARTÍNEZ-ORTÍ ET AL.: La malacofauna de la sierra de Alcaraz (Albacete, España)

Tabla I. Listado de las localidades de muestreo en la sierra de Alcaraz (Albacete, España). Abrevia-

turas: N, número de la localidad de muestreo.

Table I. List of the sampled localities in Alcaraz mountains (Albacete, Spain). Abbreviations: N,

number of the sampling locality.

U.T.M. Altitud Hábitat

305WH4683 1100 Carrascal.

305WH4882 1040 Nogales/Ribera

305WH5180 1140 Chopera/ Carrascal

305WH5379 1180 Pinada/Quejigal/Juncal

305WH5679 1300 Pinada

305SWH6077 1240 Ribera

305WH6474 900 Retamar/Ruderal

305WH4379 980 Ribera/ Chopera

305WH4376 1100 Carrascal

305WH4573 1220 Arroyo

305WH3774 840 Ribera/Sauceda

305WH3672 1000 Matorral/ Carrascal

305WH3971 940 Ribera

305WH4170 980 Pinada

305WH4367 1100 Ribera/Chopera

305WH4665 1380 Carrascal/Pinada

305WH4866 1798 Pinada/Veget. pratense

305WH4968 1600 Pinada/Carrascal

30SWH5070 1440 Pinada/Carrascal

305WH4363 1300 Pinada

305WH4362 1200 Pinada/Ribera

305WH4764 1180 Pinada/Carrascal / Matorral

305WH5165 1380 Roquedo/Arroyo

30SWH5970 1250 Pinada/Roquedo

305WH5873 1040 Ribera

305WH5974 1080 Pinada/Matorral

305WH4956 1040 Ribera/Roquedo

305WH5589 1000 Ribera

305WH5061 980 Ribera

ÑN Localidad Fecha

1 Ermita Cortés. Depósito de agua 10.10.96

2 LaHoz. Río Cortes 10.10.96

3 Peñascosas. Camping río del Arquillo 10.10.96

4 Las Baguadas 11.10.96

5 Casas de Carboneras 11.10.96

6 Nacimiento del río del Vidrio 11.10.96

7 Rambla de la Dehesa del Val 11.10.96

8 Río de Alcaraz. Molino de Potrera 12.10.96

9 Vianos. Cabecera del Bco. de los Quiñones 12.10.96

10 Carretera de Vianos, km 158,5 12.10.96

11 Reolid. Camino al Balneario. Río Salobre 15.03.97

12 Cerro de los Pizorrosos del Aljibe 15.03.97

13 Salobre. Río Salobre 15.03.97

14 La Herrería 15.03.97

15 Zapateros, a 500 m 16.03.97

16 Subida al Pico Almenaras 16.03.97

17 Pico Almenaras 16.03.97

18 Puerto del Mosquito 16.03.97

19 Cruce de carreteras 17.03.97

20 Puerto de Crucetas 17.03.97

21 Arroyo de las Fábricas 17.03.97

22 Cortijo de Vilutia 17.03.97

23 Las Espineras 17.03.97

24 Entre Batán del Puerto y Río Madera 18.03.97

25 Río Motilla 18.03.97

26 La Casa Nueva 18.03.97

27 Nacimiento Río Mundo 18.03.97

21.04.97

28 Río Arquillo (col. R. Carr) 02.06.02

29 Riópar (col. R. Carr) 01.06.02

MATERIAL Y MÉTODOS

El área de estudio engloba la sierra

de Alcaraz (Fig. 1), donde se han selec-

cionado 27 localidades extendidas por la

mayor parte de su área geográfica (Fig.

1b; Tabla I). La duración mínima de

cada muestreo ha sido de una hora y se

ha muestreado en los diferentes ambien-

tes presentes en cada localidad. Para la

localización de las estaciones se han uti-

lizado los mapas topográficos a escala

1:50.000. Cada estación se indica por sus

coordenadas UTM con precisión de 1x1

km, así como por su altitud y su hábitat

(Tabla 1). De la colección Ron Carr se

han estudiado muestras de 3 localida-

des; dos de ellas, 28 y 29, no figuraban

entre nuestras localidades de muestreo,

por lo que se han incorporado al listado,

mientras que la tercera corresponde a

nuestra localidad 27.

Iberus, 22 (2), 2004

Tabla II. Listado de especies de moluscos continentales de la Sierra de Alcaraz (Albacete, España)

(primer número: ejemplares recolectados vivos/segundo número: conchas).

Table II. List of the non-marine molluscs of the Alcaraz mountains (Albacete, Spain) (first number:

alive collected specimens/ second number: shells).

ESPECIES

LOCALIDADES

1234556785900 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29

Potamopyrgus antipodarum (/. E. Gray, 1843) 215 50 1N

Stagnicola sp. 01

Ancylus fuviatilis O. F. Múller, 1774 2/0

Carychium (Saraphia) tridentatum (Risso, 1826) 2/0

Oxyloma (Oxyloma) elegans elegans (Risso, 1826) 2/3

Cochlicopa lubrica (0. F. Múller, 1774)

Lauria (Lauria) cylindracea (Da Costa, 1778)

Vallonia costata (0. F. Múller, 1774)

Vallonia enniensis (Gredler, 1856)

Vallonia pulchella (0. F. Múller, 1774)

Acanthinula aculeata (0. F. Múller, 1774)

Pyramidula pusilla (Vallot, 1801)

Granopupa granum (Draparnaud, 1801)

Chondrina granatensis Alonso, 1974

Truncatellina callicratis (Scacchi, 1833)

Truncatellina claustralis (Gredler, 1856)

Jaminia quadridens (0. F. Miller, 1774) 0W4 0/3

Merdigera obscura (0. F. Múller, 1774) 0/4 0/4

Cecilioides (Cecilioides) acicula (0. F. Múller, 1774) 2/0

Ferussacia (Ferussacia) folliculus (Gmelin, 1791) 17

Rumina decollata (Linnaeus, 1758) 01

Paralaoma servilis (Shuttleworth, 1852) 01M

Discus rotundatus (0. F. Múller, 1774)

Vitrea (Crystallus) contracta (Westerlund, 1871) 1/0

Euconulus (Euconulus) fulvus (0. F. Miller, 1774)

Oxychilus (Oxychilus) drapamaudi draparnaudi (H. Beck, 1837)

Aegopinella nitidula (Draparnaud, 1805)

Milax (Milax) gagates (Draparnaud, 1801) 3

Vitrina pellucida pellucida (0. F. Múller, 1774)

0ligolimax annutaris (Studer, 1820) 11

Lehmannia valentiana (A. Férussac, 1823) 1 2

Deroceras (Deroceras) laeve (0. F. Múller, 1774) 3

Deroceras nitidurn (Morelet, 1845) 2

Deroceras (Deroceras) reticulatum (0. F. Múller, 1774) ES 2

Sphincterochila (Cariosula) baetica (Rossmássler, 1854)

Arion (Arion) lusitanicus J. Mabille, 1868 1 2

Cochlicella (Prietocella) barbara (Linnaeus, 1758) 0M

Monacha (Monacha) cartusiana (0. F. Múller, 1774) 111

Xerocrassa subrogata (Rossmássler in Pfeiffer, 1853) 1 15 1/8

Xerocrassa penchinati (Bourguignat, 1868)

Xerotricha conspurcata (Draparnaud, 1801) 2/4

Helicella cistorum (Morelet, 1845)

Candidula gigaxii (Pfeiffer, 1850)

Cernuella (Cernuella) virgata (Da Costa, 1778) 3/4

Xerosecta (Xerosecta) cespitum (Draparnaud, 1801) 5/0 3/1 0/4

Microxeromagna lowei (Potiez y Michaud, 1838) 21

Cepaea (Cepaea) nemoralis nemoralis (L., 1758) 0/9

Cornu aspersum (0. F. Múller, 1774) 111 2/0

Iberus qualtierianus (L, 1758) m. quiraoanus (Rossmássler, 1854)

Theba pisana (0. F. Miller, 1774)

2/0 1/0

11021 0/1

31 1/0

0/21

012 01

50/0 5/0

3/0

(Mv 10

0 0/4

2160 08

Mm 34

1/0

0/6

1/0

0/2

2/4 0/2

114 01

0/5

01 0/1. 0/2

0/5 0/2

01 0/3

8/0

2/0 0/5

1/14

110 1/0

5/5 2/0 0/4 04

03 9

0/2 0/1 01

09 11 01 1/0

0/3

1/520/2

1/0 6/0

3/0

2/0

0/2 0/2 0/120/6 01

2/0

21 1002

0/15

14/010/3

0/2

0/3

02010307 01

01 01 1/3 3/8 111 011 0/8

1/0 0/2

0/15

014 01

0 1/0

112 0102

4/0

1/0 1/0

5/0

2/6

2/0

0/5

1/0

4/0

0/20

MARTÍNEZ-ORTÍ ET AL.: La malacofauna de la sierra de Alcaraz (Albacete, España)

Figuras 2-8. Algunas especies interesantes de moluscos recolectadas en la sierra de Alcaraz (Albacete).

2. Chondrina granatensis (localidad 24) (diámetro, 2,7 mm). 3. Aegopinella nitidula (loc. 10) (diá-

metro, 7,1 mm). 4. Sphincterochila baetica (loc. 9) (diámetro, 20,05 mm). 5. Xerocrassa penchinati

(loc. 20) (diámetro, 4,2 mm). 6. Candidula gigaxii (loc. 13) (diámetro, 7,8 mm). 7. Helicella cisto-

rum (loc. 12) (diámetro, 11,1 mm). 8. /berus gualtierianus guiraoanus (loc. 21) (diámetro, 24,85 mm).

Figures 2-8. Some interesting species of molluscs collected in the Alcaraz mountains (Albacete). 2. Chon-

drina granatensis (loc. 24) (diameter 2.7 mm). 3. Aegopinella nitidula (loc. 10) (diameter 7.1 mm). 4.

Sphincterochila baetica (loc. 9) (diameter 20.05 mm). 5. Xerocrassa penchinati (loc. 20) (diameter 4.2

mm). 6. Candidula gigaxii (loc. 13) (diameter 7.8 mm). 7. Helicella cistorum (loc. 12) (diameter 11.1

mm). 8. Iberus gualtierianus guiraoanus (loc. 21) (diameter 24.85 mm).

nido en cuenta los trabajos de ALTO-

NAGA (1988), GÓMEZ (1988), FECHTER Y

FALKNER (1993), PUENTE (1994), Bo-

El listado de las especies determina-

das se muestra en la Tabla II. Se han rea-

lizado estudios anatómicos del aparato

reproductor de aquellas especies cuyos

caracteres conquiliológicos son insufi-

cientes para identificarlas correcta-

mente. Para la determinación taxonó-

mica y posición sistemática se han te-

RREDA (1996), GLOER Y MEIER-BROOK

(1998), MARTÍNEZ-ORTÍ (1999), BANK,

FALKNER, NORDSIECK Y RIPKEN (2001) y

FALKNER, RIPKEN Y FALKNER (2002), en-

tre otros. Todo el material recolectado se

Iberus, 22 (2), 2004

encuentra depositado en el Museu Va-

lencia d'Historia Natural.

RESULTADOS Y DISCUSIÓN

Se han hallado un total de 50 especies

de moluscos, pertenecientes a 14 superfa-

milias y 27 familias, de las cuales 47 corres-

ponden a gasterópodos terrestres y tres a

dulceacuícolas. Las especies de moluscos

encontradas en el área de muestreo apa-

recen relacionadas en la Tabla II.

Comentarios taxonómicos: La taxono-

mía del género Stagnicola Jeffreys, 1830

ha sido revisada recientemente por

diversos autores (BARGUES, VIGO,

HORAK, DVORAK, PATZNER, POINTIER,

JACKIEWICZ, MEIER-BROOK Y MAS-COMA,

2001, con referencias; FALKNER ET AL.,

2002, con referencias). En la actualidad,

se acepta que este género está represen-

tado en Europa por cinco especies, de las

que sólo una, S. fuscus (Pfeiffer, 1821), ha

sido caracterizada en España, mediante

análisis molecular y estudios anatómi-

cos. La ausencia de partes blandas en el

único ejemplar recogido (una concha

juvenil), impide su asignación a una

especie concreta del género Stagnicola.

ROBLES, BORREDA Y COLLADO (1991)

citan Carychium sp. en la región de

Almansa. La revisión de este material

permite asignarlo a C. tridentatum

(Risso, 1826).

Helix semipicta Hidalgo, 1870, cuya

localidad tipo es la sierra de Alcaraz, es

un sinónimo posterior de Helicella cisto-

rum (Morelet, 1845) (MARTÍNEZ-ORTÍ Y

APARICIO, 2003). Esta última especie es

un endemismo ibérico (PUENTE, 1994)

que se extiende por el Alemtejo (locali-

dad tipo) y el este del Algarve (Portu-

gal) y por las provincias españolas de

Badajoz y Cáceres, suroeste de Madrid y

de Toledo, Ciudad Real, Córdoba, Jaén,

Huelva y Sevilla. Por lo tanto, su

hallazgo constituye la primera cita en la

provincia de Albacete, siendo además,

la más oriental para esta especie (MARTÍ-

NEZ-ORTÍ Y APARICIO, 2003).

La composición-específica del género

Iberus Montfort, 1810 es controvertida.

Los autores más recientes (PUENTE, 1994;

ARRÉBOLA, 1995) consideran que existe

una sola especie muy polimórfica, [. gual-

tierianus (Linnaeus, 1758), criterio que se-

guimos aquí. No obstante, hay que tener

en cuenta que otros autores (APARICIO,

1983; APARICIO Y RAMOS, 1988) reconocen

la existencia de varias especies bien defi-

nidas dentro del género, una de las cuales

sería [. guiraoanus, si bien APARICIO Y RA-

MOS (1988) plantean la necesidad de nue-

vos estudios para resolver los problemas

taxonómicos planteados. Actualmente se

están realizando análisis moleculares (B.

GÓMEZ, com. pers.), que esperamos con-

tribuyan a solventar el problema. l. guira-

oanus fue descrito originalmente de Cas-

tellón por ROSssMASSLER (1854), donde no

ha vuelto a ser encontrado, y es conocido

de las provincias de Granada y Jaén (var.

angustata Rossmássler, 1854) (GARCÍA

SAN NICOLÁS, 1957; ALONSO, 1975; APARI-

CIO, 1983).

ORTIZ DE ZÁRATE (1991) cita Iberus

alcarazanus (Guirao en Rossmássler,

1854) en la sierra de Alcaraz, pensamos

que erróneamente, ya que el morfotipo

alcarazanus es más globoso y no presenta

ombligo. La posesión de un ombligo

bien conformado es la principal caracte-

rística del morfotipo guiraoanus.

Comentarios biogeográficos: Desde el

punto de vista biogeográfico podemos

agrupar las especies halladas en los si-

guientes conjuntos, teniendo en cuenta

sus áreas de distribución: una especie in-

troducida, de origen indo-australiano:

Potamopyrgus antipodarum,; siete especies

holárticas: Oxyloma elegans elegans, Cochli-

copa lubrica, Vallonia costata, V. pulchella,

Euconulus fulvus, Vitrina pellucida pellucida

y Deroceras laeve; dos paleárticas: Acanthi-

nula aculeata y Ancylus fluviatilis; cuatro

especies de amplia distribución europea:

Carychium tridentatum, Merdigera obscura,

Deroceras reticulatum y Aegopinella

nitidula; otras cuatro especies de amplia

distribución europea, aunque más res-

tringida: Vallonia enniensis, Discus rotun-

datus, Oligolimax annularis y Cepaea nemo-

ralis nemoralis; doce especies de distribu-

ción europea occidental y mediterránea:

Lauria cylindracea, Pyramidula pusilla,

MARTÍNEZ-ORTÍ ET AL.: La malacofauna de la sierra de Alcaraz (Albacete, España)

Truncatellina callicratis, T. claustralis, Jami-

nia quadridens, Cecilioides acicula, Oxychi-

lus draparnaudi draparnaudi, Monacha car-

tusiana, Cernuella virgata, Cornu aspersum,

Milax gagates y Candidula gigaxii; cuatro

especies preferentemente mediterráneas:

Ferussacia folliculus, Rumina decollata, Pa-

ralaoma servilis y Microxeromagna lowet;

otras cuatro especies preferentemente

mediterráneas que alcanzan la costa

atlántica occidental: Granopupa granum,

Cochlicella barbara, Xerotricha conspurcata y

Theba pisana pisana; una especie ibérica

cuya área de distribución se prolonga por

el SE de Francia: Xerosecta cespitum; una

especie iberomagrebí: Sphincterochila bae-

tica; ocho endemismos ibéricos: Chon-

drina granatensis, Xerocrassa subrogata, X.

penchinati, Helicella cistorum, Iberus gualtie-

rianus morfotipo guiraoanus, Lehmannia

valentiana, Deroceras nitidum y Arion lusi-

tanicus (FECHTER Y FALKNER, 1993; ALTO-

NAGA, GÓMEZ, MARTÍN, PRIETO, PUENTE y

RALLO, 1994; PUENTE, 1994; BORREDA,

1996). Alguna de ellas, como Lehmannia

valentiana y Arion lusitanicus o Potamopyr-

gus antipodarum, que originariamente

presentaban una distribución natural ibé-

rica (las dos primeras) o de Nueva Ze-

landa (la última), se han extendido am-

pliamente por varios continentes (FECH-

TER Y FALKNER, 1993; BORREDA, 1996).

Esta fauna malacológica presenta

una geonemia predominantemente

mediterránea con clara influencia conti-

nental, como corresponde al área geo-

gráfica estudiada. Cuarenta y seis de las

50 especies determinadas son comunes

a la Comunidad Valenciana. De las

cuatro restantes, Chondrina granatensis e

Iberus gualtierianus morfotipo guiraoanus

son especies de distribución meridional,

que alcanzan en la sierra de Alcaraz el

límite norte de su área de distribución,

donde se han encontrado por primera

vez. Helicella cistorum es un endemismo

ibérico occidental, correspondiendo la

nueva localidad, cerro de Los Pizarro-

sos, al punto más oriental conocido. Por

último, Candidula gigaxt presenta distri-

bución discontinua en la Península

Ibérica y es conocida en algunos puntos

de la provincia de Jaén cercanos a la

sierra de Alcaraz.

Conservación:La mayoría de las es-

pecies halladas en la sierra de Alcaraz

carecen actualmente de medidas de pro-

tección a nivel autonómico, estatal o eu-

ropeo. En general, se trata de especies

triviales que presentan una amplia dis-

tribución peninsular. Sin embargo, Ibe-

rus gutraoanus ha sido incluido en el Ca-

tálogo Regional de Especies Amenaza-

das de Castilla-La Mancha (Decreto

33/1998 de 5 de Mayo), con la categoría

de “interés especial”. El grupo de exper-

tos de la Sociedad Española de Malaco-

logía no propone ningún taxón de los

hallados en esta región para su inclu-

sión en el Catálogo Nacional de Espe-

cies Amenazadas (ALONSO, ALTONAGA,

ÁLVAREZ, ARAUJO, ARCONADA, ARRÉ-

BOLA, BECH, BROS, CASTILLEJO, GÓMEZ,

IBÁÑEZ, LUQUE, MARTÍNEZ-ORTÍ, MO-

RENO, PRIETO, PUENTE, PUJANTE, KRO-

BLES, ROLÁN Y TEMPLADO, 2001).

CONCLUSIONES

Se han hallado por primera vez

veintidós especies en la provincia de

Albacete, diez de las cuales (señaladas

con un asterisco) se citan por primera

vez en la Comunidad de Castilla-La

Mancha: Carychium tridentatum, Cochli-

copa lubrica, Lauria cylindracea, Vallonia

enniensis, V. pulchella, Acanthinula acule-

ata (*), Pyramidula pusilla, Truncatellina

claustralis (*), Chondrina granatensis (*),

Merdigera obscura, Paralaoma servilis (*),

Discus rotundatus (*), Aegopinella nitidula

(*), Euconulus fulvus (*), Oxychilus dra-

parnaudi draparnaudi, Vitrina pellucida

pellucida, Oligolimax annularis, Sphincte-

rochila baetica (*), Candidula gigaxii, Heli-

cella cistorum, Xerocrassa penchinati (*) e

Iberus gualtierianus morfotipo guiraoanus

(*). Estas nuevas citas amplían el conoci-

miento sobre la distribución de estas

especies en la Península Ibérica.

AGRADECIMIENTOS

A Vicent Borreda, Juan Domínguez y

Gloria Tapia por su colaboración en la

recogida de las muestras, y a V. Borreda,

Iberus, 22 (2), 2004

además, por confirmar algunas determi-

naciones de babosas. A la Sección de

Microscopía Electrónica del S. C. 1. E. de

la Universitat de Valencia por su ayuda en

BIBLIOGRAFÍA

ALONSO, M. R., 1975. Moluscos terrestres y dul-

ceacuícolas de la depresión de Granada (Es-

paña) y sus alrededores. Cuadernos de Cien-

cias Biológicas, 4 (2): 125-157.

ALONSO M. R., ALTONAGA, K., ÁLVAREZ, R.

M., ARAUJO, R., ARCONADA, B., ARRÉBOLA,

J. R., BECH, M., BROS, V., CASTILLEJO, J., GÓ-

MEZ, B. J., IBÁÑEZ, M., LUQUE, A., MaARTÍ-

NEZ-ORTÍ, A., MORENO, D., PRIETO, C.,

PUENTE, A. I., PUJANTE, M., ROBLES, F., Ro-

LÁN, E. Y TEMPLADO, J., 2001. Protección de

Moluscos en el Catálogo Nacional de Espe-

cies Amenazadas. Eds.: Gómez, B., Mo-

reno, D., Rolán, E., Araujo, R. y Alvarez, R.

M. Reseñas malacológicas (Sociedad Española

de Malacología), 11: 1-286.

ANDÚJAR TOMÁS, A. Y GÓMEZ DE GUEVARA,

R., 1985. Ropalóceros de la sierra de Alcaraz y

Calar del Mundo. Albacete. Instituto de Estu-

dios Albacetenses, CSIC, CECEL, Albacete,

191 pp.

ALTONAGA, K., 1988. Estudio anatómico y bioge-

ográfico de las familias Endodontidae, Euconu-

lidae, Zonitidae y Vitrinidae (Gastropoda: Pul-

monata: Stylommatophora) de la Península Ibé-

rica, con especial referencia al País Vasco y zonas

adyacentes. Universidad del País Vasco, Te-

sis doctoral (inédita), 529 pp.

ALTONAGA, K., GÓMEZ, B., MARTÍN, R., PRIETO,

C. E., PUENTE, A. I. Y RALLO, A., 1994. Estu-

dio faunístico y biogeográfico de los moluscos te-

rrestres del norte de la Península Ibérica. Parla-

mento Vasco, Vitoria, 503 pp.

APARICIO, M. T., 1983. The chromosomes of

eight species of the subfamily Helicinae (Gas-

tropoda, Pulmonata, Helicidae) from Spain.

Malacological Review, 16: 71-78.

APARICIO, M. T. Y RAMOS, M. A., 1988. A com-

parative study of the morphology of the pul-

monate snail Pseudotachea litturata (Pfeiffer)

and other species of Pseudotachea, Iberus and

Cepaea. Journal of Molluscan Studies, 54: 278-

294.

ARRÉBOLA, J. R., 1995. Caracoles terrestres (Gas-

tropoda, Stylommatophora) de Andalucía, con

especial referencia a las provincias de Sevilla y Cá-

diz. Universidad de Sevilla, Tesis Doctoral

(Inédita), 589 pp.

la utilización del microscopio electrónico

de barrido. Al Instituto de Estudios Alba-

cetenses por la financiación parcial de este

estudio mediante el proyecto 195/96.

ARRÉBOLA, J. R. Y GÓMEZ. B., 1988. Nuevas

aportaciones al conocimiento del género

Chondrina (Gastropoda, Pulmonata) en el sur

de la Península Ibérica, incluyendo la des-

cripción de Chondrina maginensis spec. nov.

Iberus, 16 (2):109-116.

BANK, R. A., FALKNER, G., NORDSIECK, H. Y RIpP-

KEN, TH. E. J., 2001. CLECOM-PROJECT.

First Update to Systematics and Nomencla-

ture of the CLECOM-Checklists, including

Corrigenda et Addenda to the printed Lists.

Heldia, 4 (1/2): 1-76, Supplement: A1-A6.

BARGUES, M. D., VIGO, M., HORAK, P., Dvo-

RAK, J., PATZNER, R. A., POINTIER, J. P., Jac-

KIEWICZ, M., MEIER-BROOK, C. Y MAS-COMA,

S., 2001. European Lymnaeidae (Mollusca:

Gastropoda), intermediate hosts of tremato-

diases, based on nuclear ribosomal DNA

ITS-2 sequences. Infection, Genetics and Evo-

lution 1: 85-107.

BORREDA, V., 1996. Pulmonados desnudos (Mo-

llusca: Gastropoda: Pulmonata) del Este de la

Península Ibérica. Tesis Doctoral (Inédita),

Universitat de Valencia, 475 pp.

BORREDA, V. Y COLLADO, M. A., 1991. Intro-

ducción al conocimiento de los Pulmonados

desnudos de la provincia de Albacete (Cas-

tilla-La Mancha, España). Actas de las Jorna-

das sobre el medio natural albacetense. Instituto

de Estudios Albacetenses: 377-384.

BORREDA, V., COLLADO, M. A., BLASCO, J. Y Es-

PÍN, J. S., 1991. Nuevos datos sobre los pul-

monados desnudos (Mollusca, Gastropoda)

de la provincia de Albacete (Castilla-La Man-

cha, España). Instituto Estudios Albacetenses.

Al-Basit, 29: 1-17.

FALKNER, G., RIPKEN, TH. E. J. Y FALKNER, M.,

2002. Mollusques continentaux de France. Liste

de Référence annotée et Bibliographie. Muséum

national d'Histoire naturelle, Patrimoines

Naturels, n* 52, 350 pp.

FECHTER, R. Y FALKNER, G., 1993. Guía de la na-

turaleza Blume: Moluscos. Ed. Gaybán Grafic

SA, Barcelona, 287 pp.

GARCÍA SAN NICOLÁS, E., 1957. Estudios sobre

la biología, la anatomía y la sistemática del

género Iberus Montfort, 1810. Boletín de la

Real Sociedad Española de Historia Natural (Sec-

ción Biología), 55: 199-390.

MARTÍNEZ-ORTÍ £7 AL.: La malacofauna de la sierra de Alcaraz (Albacete, España)

GASULL, L., 1975. Fauna malacológica terrestre

del sudeste ibérico. Boletín de la Sociedad His-

toria Natural de Baleares, 20: 149 pp.

GLOEÉR, P. Y MEIER-BROOK, C., 1998. Stsswas-

sermollusken. Ein Bestimmunggsschlússel fir

die Bundersrepublik. Deutschland, 12 Ed.

Deutscher Jugendbund fúr Naturbeobach-

tung, 136 pp.

Gómez, B., 1988. Estudio sistemático y biogeo-

gráfico de los moluscos terrestres del Suborden

Orthurethra (Gastropoda: Pulmonata: Stylom-

matophora) del País Vasco y regiones adyacen-

tes, y catálogo de las especies ibéricas. Universi-

dad del País Vasco, Tesis Doctoral (inédita),

424 pp.

GÓMEz, B. J. Y DANTART, L., 1996. Moluscos te-

rrestres ibéricos del suborden Orthurethra.

Nuevos datos de distribución. Iberus, 14 (1):

103-107.

GUERRERA MONTES, J., ROs EspPín, R., HERAS

IBÁÑEZ, J. DE LAS, GARCÍA ZAMORA, P. Y Ji-

MÉNEZ MARTÍNEZ, N., 1989. Estudio de la

flora briofítica de la sierra de Alcaraz (Alba-

cete) como base para una evaluación fito-

biológica del territorio. Instituto de Estudios

Albacetenses, serie I: Ensayos Históricos y Cien-

tíficos, 39: 1-113.

HERRANZ, J. M. Y GÓMEZ CAMPO, C., 1986. Con-

tribución al conocimiento de la flora y vegetación

de la comarca de Alcaraz (Albacete). Caja de

Ahorros de Albacete, 279 pp.

HIDALGO, J. G., 1870. Description de trois espe-

ces nouvelles d'Helix d'Espagne. Journal de

Conchyliologie, 298-299.

JIMÉNEZ, J. Y MARTÍNEZ-LÓPEZ, F., 1988. Distri-

bución y composición específica de la mala-

cofauna del río Júcar. Limnetica, 4: 9-18.

MARTÍNEZ-LÓPEZ, F., PUJANTE, A. M. Y TAPIA,

G., 1993. Sobre la malacofauna del río Júcar:

importancia de los Gastropoda como hos-

pedadores intermediarios de Trematodos

Digenea. Revista de Estudios Albacetenses, Al-

Basit, 32: 169-211.

MARTÍNEZ-ORTÍ, A., 1999. Moluscos terrestres

testáceos de la Comunidad Valenciana. Univer-

sitat de Valencia, Tesis doctoral (inédita), 743

O A. Y APARICIO, M. T., 2003.

Taxonomical clarification of Helix semipicta

Hidalgo, 1870 (Gastropoda, Pulmonata: Hy-

gromiidae). Journal of Conchology, 38 (2): 125-

129.

PUENTE, A. IL, 1994. Estudio taxonómico y bioge-

ográfico de la superfamilia Helicoidea Rafinesque,

1815 (Gastropoda: Pulmonata: Stylommatop-

hora) de la Península Ibérica e Islas Baleares.

Universidad del País Vasco, Tesis Doctoral

(Inédita), 970 pp.

RAMOS, M. A. Y APARICIO, M. T., 1985. Gaste-

rópodos terrestres y dulceacuícolas de las

Lagunas de Ruidera (España). Iberus, 5: 113-

128:

ROBLES, F., BORREDA, V. Y COLLADO, M. A,,

1991. Gasterópodos terrestres y dulceacuí-

colas de la región de Almansa. Jornadas sobre

el Medio Natural Albacetense, Instituto Estu-

dios Albacetenses, 385-393.

RUEDA, J., HERNÁNDEZ, R Y TAPIA, G., 2001.

Biodiversidad, caracterización de los inver-

tebrados y calidad biológica del río Júcar a

su paso por la provincia de Albacete. Sabuco,

1: 7-42.

VIDAL-ABARCA, C., SUÁREZ, M. L., MILLÁN, A.,

GÓMEZ, R., ORTEGA, M., VELASCO, J. Y RA-

MÍREZ-Díaz, L., 1991. Estudio limnológico

de la cuenca del río Mundo (río Segura). Jor-

nadas sobre el Medio Natural Albacetense, Ins-

tituto Estudios Albacetenses, 339-357.

oe sosdlA) Ba: É SE > GON

4%,

nia A Y de HARO) " e '

Ni A 18

UI ER

| bed 1 st

rial JE

ps NO

O Sociedad Española de Malacología Iberus, 22 (2): 19-31, 2004

Los moluscos de las aguas continentales de la provincia de

Huelva (SO España)

Freshwater Molluscs of Huelva Province (SW Spain)

Juan Carlos PÉREZ-QUINTERO*, Miguel BECH TABERNER** y José

Luis HUERTAS DIONISIO***

Recibido el 25-VI-2003. Aceptado el 19-V-2004

RESUMEN

En este trabajo se estudia la faunística y distribución de los moluscos de las aguas conti-

nentales de la provincia de Huelva (SO de España). Se han encontrado 29 especies de

gasterópodos y bivalvos repartidos en cinco cuencas hidrográficas, siendo las especies

de mayor abundancia Physella acuta, Ancylus fluviatilis, Planorbarius metidjensis, Radix

balthica (= R. peregra) (Gastropoda) y Pisidium casertanum (Bivalvia); todas las especies

se referencian en el entramado provincial de cuadrículas UTM 10 x 10 km.

ABSTRACT

In this work we study the faunistic and distribution of freshwater molluscs of Huelva (SW

Spain). We have found 29 species of gastropods and bivalves in five hydrographic

basins, the most abundant species were Physella acuta, Ancylus fluviatilis, Planorbarivs

metidjensis, Radix balthica (= R. peregra) [(Gastropoda)] and Pisidium casertanum (Bival-

via); all the species are refered to the 10 x 10 km UTM grid.

PALABRAS CLAVE: Moluscos, Gasterópodos, Bivalvos, Huelva, Andalucía, España, Distribución, Ríos.

KEY WORDS: Molluscs, Gastropoda, Bivalvia, Huelva, Andalucia, Spain, Distribution, Rivers

INTRODUCCIÓN

La malacofauna continental de la

provincia de Huelva ha sido, en general,

poco estudiada; mientras que existe un

catálogo bastante preciso de las especies

terrestres (MUÑOZ, 1992), de la faunística

dulceacuícola sólo se conocen aproxima-

ciones muy generales o localizadas

(ORTIZ DE ZÁRATE y ORTIZ DE ZÁRATE,

1961; BIGOT y MARAZANOF, 1965; MARA-

ZANOF, 1966; GASULL, 1985; PÉREZ-QUIN-

TERO, 1988), habiéndose citado un

máximo de 14 especies en el entorno

provincial de las que la mayoría han sido

descritas en el entorno de las marismas

del Guadalquivir y Coto de Doñana.

Recientemente, MONTES DEL OLMO

(1993), estudiando el efecto de la intro-

ducción de Procambarus clarkii (GIRARD,

1852) en Doñana, encuentra sólo cinco

especies en el entorno del Parque Nacio-

nal, no existiendo por tanto confirma-

ción de la presencia actual de algunas de

las descritas previamente en la zona.

Tras un detallado estudio de los cursos

* Departamento de Biología Ambiental y Salud Pública, Universidad de Huelva. Avda. Fuerzas Armadas s/n,

21071 Huelva (España). E-mail: jcperezQuhu.es

** C/ Córcega, 404. 08037 Barcelona

** Ayda. Andalucía, 5. 21004 Huelva

Iberus, 22 (2), 2004

Figura 1. Distribución de las cinco cuencas hidrográficas de la provincia de Huelva: I Guadiana, II

Piedras, IM Odiel, IV Tinto, V Guadalquivir.

Figure 1. Distribution of the five hydrographic basins of the province of Huelva: I Guadiana, II Piedras,

111 Odiel, IV Tinto, V Guadalquivir.

de agua de la provincia, en este trabajo

se amplía el listado de especies a 29 (21

Gasterópodos y 8 Bivalvos), aportando

datos sobre la distribución actual de las

mismas en las distintas cuencas hidro-

gráficas de la red fluvial provincial.

MATERIAL Y MÉTODOS

Se han prospectado 251 localidades,

habiéndose encontrado moluscos en 228

de ellas (91,0 % de frecuencia de

captura): 64 en la cuenca del río Guadal-

quivir, 40 en la del río Tinto, 48 en la del

río Odiel y 76 en la de los ríos Guadiana

y Piedras (Fig. 1), que han sido muestre-

adas entre los años 1999 y 2002, todas

ellas en, al menos, dos ocasiones.

En cada estación se registraba la coor-

denada UTM y la altitud, ambas

mediante GPS (ver Anexo); se anotaba,

igualmente, la profundidad, densidad y

situación espacial de los individuos, así

como una estima de la velocidad de la

corriente y la granulometría del sustrato.

Para cada especie se ha calculado su

cobertura como porcentaje de presencia

referido a las 127 cuadrículas UTM 10x10

km de la provincia, y su abundancia

como porcentaje de presencia referido a

las 228 estaciones en que se han encon-

trado moluscos; en ambos casos siempre

se computan los individuos vivos, no sus

restos. En cada cuenca se llevaron a cabo

dos aproximaciones a su diversidad: con-

signando el número de especies, géneros

y familias (riqueza taxonómica), y calcu-

lando su diversidad específica según la

expresión de SHANNON-WEAVER: H= -)2,

p¡ In p¡ (MAGURRAN, 1988); a efectos de

cálculo, y considerando la escasa superfi-

cie de la cuenca del río Piedras y su pro-

ximidad geográfica con la del río Gua-

diana, se creyó conveniente asociar la

primera con la segunda.

Los moluscos se colectaron barriendo

el fondo de los cauces con redes de 0,3 mm

de luz, y capturándolos manualmente en

rocas, plantas y otros sustratos sumergi-

dos. Tras la captura y lavado de muestras

los ejemplares fueron conservados en

alcohol al 70% y posteriormente determi-

nados a nivel de especie siguiendo las indi-

caciones de manuales clásicos (ADAM,

1960; ELLIS, 1978) y actuales (GLOER y

MEIER-BROOK, 2003). En algunas especies

de difícil determinación a partir de la mor-

PÉREZ-QUINTERO 7 4£.: Moluscos de aguas continentales de Huelva (SO España)

fología de su concha, se recurrió al análi-

sis de su aparato reproductor.

RESULTADOS

En el análisis de las cuencas y sub-

cuencas estudiadas se han encontrado 25

géneros y 29 especies, de las que 27 son

autóctonas (93,1 %) y 2 introducidas (Pota-

mopyrgus antipodarum (J.E. Gray 1843) y

Corbicula fluminea (O.F. Múller 1774)), apa-

reciendo como primeras citas para la pro-

vincia: Bithynia (Bithynia) tentaculata (Lin-

naeus 1758), Potamopyrgus antipodarum,

Hydrobía (Hydrobia) acuta (Draparnaud

1805), Mercuria emiliana (Paladilhe 1869),

Islamia minuta (Draparnaud 1805), Stagni-

cola palustris (O.F. Múller 1774), Radix aff.

auricularia (Linnaeus 1758), Radix balthica

Linnaeus 1758, Bulinus (Isidora) truncatus

contortus (Michaud 1829), Planorbarius

metidjensis (Forbes 1838), Ferrisia (Pet-

tancylus) clessiniana (Jickeli 1882), Planor-

bis (Planorbis) carinatus O.F. Múller 1774,

Gyraulus (Gyraulus) albus (O.F. Múller

1774), Hippeutis complanatus (Linnaeus

1758), Myosotella myosotis (Draparnaud

1801), Potomida littoralis (Cuvier 1798),

Musculium (Musculium) lacustre (O.F.

Miller 1774), Pisidium (Euglessa) caserta-

num (Poli 1791), Pisidium (Euglessa) perso-

natum Malm 1855 y Pisidium (Cingulipisi-

dium) milium Held 1836. El número de

especies por curso de agua ha oscilado

entre 1 y 9 (= 2,6, sd= 1,5); la cuenca de

los ríos Guadiana-Piedras es la más diversa

(23 especies, H= 3,75), la menos diversa la

del río Tinto (12 especies, H= 2,19).

DISCUSIÓN

Bithynia (Bithynia) tentaculata (Fig. 2,

Cobertura (C)= 2,3%, Abundancia (A)=

1,3%) es una especie holártica (BOETERS,

1998), residente en Huelva en cauces de

poca profundidad sobre sustratos

rocosos o macrófitas de la cuenca del

Guadiana; una concha vacía en la rivera

de Nicoba, cuenca del río Tinto, atesti-

gua su presencia en la zona, aunque el

grado de erosión de la misma y su

ausencia en el resto de estaciones de la

cuenca parecen indicar que, probable-

mente, la especie no se encuentre actual-

mente en el entorno de dicho río.

Potamopyrgus antipodarum (Fig. 2, C=

0,7%, A= 0,8%) es una especie originaria

de Nueva Zelanda (PONDER, 1988), aunque

ANISTRATENKO (1997) sugiere que indivi-

duos de este género han aparecido, a

comienzos del siglo XX, en depósitos cua-

ternarios de Lituania, pudiendo existir en

la actualidad más de una especie en

Europa. En la provincia se encuentra muy

localizada en tramos de cabecera de la

cuenca del río Odiel, asociada siempre a

sustratos rocosos y macrófitas sumergi-

das entre 5 y 30 centímetros de profundi-

dad, alcanzando densidades de más de

100 individuos por metro cuadrado.

Hydrobia (Hydrobia) acuta (Fig. 2, C=

1,5%, A= 0,8%) es una especie residente

en las costas de Europa occidental en

entornos litorales con salinidad superior

al 10%o (WILKE, ROLAN y Davis, 2000).

En la provincia se ha encontrado en cris-

talizadores de salinas de extracción in-

dustrial situadas en marismas mareales

del Paraje Natural Marismas del Odiel.

Peringia ulvae (Pennant 1777) (Figura

2, C= 0,7%, A= 0,4%) se distribuye a lo

largo de las costas de Europa occidental,

en el sedimento litoral, con un intervalo

de salinidad de 4-33%o (KERNEY, 1999).

En la provincia se encuentra en maris-

mas mareales con influencia continental

en la desembocadura del río Piedras,

entre sedimentos y sustratos rocosos.

Mercuria emiliana (Fig. 2, C= 2,3%,

A= 1,3%) ha sido descrita en localidades

de Granada, Alicante, Tarragona y

Mallorca (BOETERS, 1988), recientemente

ha sido encontrada en riveras de

Almería (BAYO MONTORO, comunicación

personal). En Huelva se encuentra en la

cuenca baja del río Guadiana, en sustra-

tos rocosos de cursos de agua dulce.

Islamia minuta (Fig. 2, C= 5,5%, A=

6,1%) es una especie abundante en cursos

de agua leníticos o lóticos, en entornos

calcícolas, de la cabecera de las cuencas

de los ríos Guadiana, Odiel y Guadalqui-

vir. Esta especie fue nominada original-

mente como Valvata globulina, familia Val-

vatidae, posteriormente BINDER (1967a,

1967b) demuestra que la ornamentación

Iberus, 22 (2), 2004

de su protoconcha es propia de la familia

Hydrobiidae, habiendo sido ubicada

dentro del género Neohoratia Schútt 1961

por BOETERS (1998), TURNER, KUIPER,

THEw, BERNASCONI, RÚETSCHI, WUTH-

RICH y GOSTELI (1998) y FALKNER, RIPKEN

y FALKNER (2002), mientras que el comité

del proyecto CLECOM (2002) la considera

sinónima del género Islamia Radoman

1974, considerando como nombre válido

para la especie el de Islamia minuta.

El género Melanopsis Férussac 1807 es

muy polimórfico cariotípica (BARSIENE,

TAPIA y BARSYTE, 1998) y conquiológica-

mente (GLAUBRECHT, 1996; PUJANTE, TA-

PIA y MARTÍNEZ, 1998). Melanopsis prae-

morsa (Fig. 2, C= 2,3%, A= 1,7%) se carac-

teriza por tener morfotipo liso sin quillas

en la teloconcha (PUJANTE, comunicación

personal), distribuyéndose en la Penín-

sula Ibérica por Andalucía centro-occi-

dental (GÓMEZ, MORENO, ROLÁN,

ARAUJO y ÁLVAREZ, 2001); en la provincia

de Huelva se encuentra únicamente en la

cuenca del río Guadalquivir, en cursos de

cabecera de la rivera de Huelva, en me-

dios leníticos e incluso en acequias.

Actualmente los únicos criterios

fiables para la correcta determinación de

Stagnicola palustris (Fig. 2, C= 0,7%, A=

0,4%) se basan en el estudio morfológico

del aparato reproductor y en el análisis

de la sección terminal de la próstata

(GLOER y MEIER-BROOK, 2003), siendo la

presente la primera confirmación de la

presencia de la especie en España basada

en el estudio de la anatomía de su

aparato reproductor (MARTÍNEZ ORTÍ,

comunicación personal). En la provincia

de Huelva se encuentra en una sola loca-

lidad de cabecera de la cuenca del Odiel,

en un entorno de intenso manejo

humano del Parque Natural de la Sierra

de Aracena y Picos de Aroche, en aguas

leníticas y profundidad entre 20 y 100 cm,

menos frecuentemente fuera del agua.

Galba truncatula (Fig. 2, C= 12,5%, A=

8,37) es una especie holártica de carácter

anfibio que se distribuye entre zonas de

cabecera y marismas mareales (KERNEY,

1999). Se ha encontrado en todas las

cuencas analizadas, siendo frecuente que

aparezca asociada a rocas emergidas a 10-

30 cm del borde del agua.

Radix aff. auricularia (Fig. 2, C= 2,3%,

A= 1,7%) reside en aguas leníticas hasta

con un 6% de salinidad (GIROD,

BIANCHI y MARIANLI, 1980), habiendo

sido encontrada en la provincia en la

subcuenca de la rivera de Huelva

(cuenca del Guadalquivir) en sustratos

rocosos de distintas superficies, y en

entornos de marisma dulce de la cuenca

del río Piedras en fondos de cauces con

sustratos de limos y arcillas.

La sistemática de los Lymnaeidae

europeos ha sido revisada reciente-

mente desde el punto de vista molecular

(BARGUES, VIGO, HORAK, DVORAK,

PATZNER, POINTIER, JACKIEWICZ, MEIER-

BROOK y MAsS-COMA, 2001) y de priori-

dad de nomenclatura (FALKNER ET AL.,

2002), concluyendo en ambos casos que

la especie Radix peregra (Muller 1774) es

sinónima de Radix balthica (Linnaeus

1758). Esta es una especie eurícora que

puede residir desde el crenon hasta en

aguas mesohalinas con salinidad del

14%o (GIROD ET AL., 1980); en Huelva

(Figura 2, C= 14,9%, A= 17,9%) aparece

en todas las cuencas muestreadas, desde

en aguas muy limpias hasta ligeramente

eutrofizadas, incluso ha sido encontrada

fuera del agua en fuentes y acequias.

Physella (Costatella) acuta (Drapar-

naud 1805) se distribuye en Norteamé-

rica, Europa y África, siendo uno de los

gasterópodos más abundantes de la

Península Ibérica (VIDAL-ABARCA y

SUÁREZ, 1985); es una especie de

amplios requerimientos ecológicos que

se encuentra indistintamente en cabece-

ras y desembocaduras, asociada a

medios lóticos o leníticos desde O hasta

2 m de profundidad (MOUTHON y

Duboss, 2001). Es la especie más amplia-

mente representada en la provincia de

Huelva (Fig. 2, C= 61,4%, A= 67,9%),

encontrándose en todas las cuencas en

aguas limpias o muy eutrofizadas.

Bulinus truncatus contortus (Fig. 2, C=

2,3%, A= 1,7%) se distribuye a lo largo

de la región mediterránea, Portugal y

localidades del sudeste de Asia y Africa

(BROWN, 1980); en Huelva se localiza en

entornos lagunares de la cuenca del

Odiel, encharcamientos someros tempo-

rales del Parque Nacional de Doñana y

PÉREZ-QUINTERO ET AL.: Moluscos de aguas continentales de Huelva (SO España)

Bithynia tentaculata Potamopyrgus antipodarum

pe JU eN

¿6000 00.

(1111000 7

j o QB

al EAN

0 ll e el al s

Ol /

PB Ú

LS —

QAN |]

Radix balthica

yl NN ATT

dl 1% dqbdsgdeas

4 lA f |

: B f iaa

Ancylus fluviatilis

Hippeutis complanatus

Gyraulus laevis

Gyraulus albus

Figura 2. Mapas de distribución. Los círculos blancos indican la presencia de conchas o valvas

vacías.

Figure 2. Distribution maps. White circles are empty shells or valves.

Iberus, 22 (2), 2004

pequeños tributarios del río Piedras,

siempre asociada a macrófitos o superfi-

cies cubiertas por películas de algas.

Planorbarius metidjensis (Fig. 2, C=

28,3%, A= 24,5%) se distribuye por la

Península Ibérica y norte de África

(VIDAL-ABARCA y SUÁREZ, 1985); es una

especie tolerante a condiciones micro-

ambientales, lenitófila, muy depen-

diente del sustrato vegetal y residente

en aguas de diferente dureza y salinidad

(MEDEIROS y SIMOES, 1987). En la pro-

vincia se encuentra en todas las cuencas

analizadas, localizándose siempre en

aguas leníticas asociada a sustratos

rocosos de gran superficie y, en menor

medida, sobre macrófitos.

La presencia de Ferrisia clessiniana

(Fig. 2, C= 9,4%, A= 7,0%) fue descrita

por primera vez en la Península Ibérica,

como Ferrisia wautterí (Mirolli, 1960), por

ALTABA, TRAVESET, BOGUÑÁ y BECH

(1985); es una especie muy polimórfica,

residente en aguas calmas o con flujo

moderado (WAUTIER, 1977). Se ha encon-

trado en aguas leníticas de las cuencas

de los ríos Guadiana, Odiel, Tinto y Gua-

dalquivir, sobre vegetación o, más fre-

cuentemente, sustratos rocosos de dis-

tinta superficie.

Planorbis carinatus (Fig. 2, C= 3,9%,

A= 1,7%) se distribuye por Europa y

oeste de Siberia (VIDAL-ABARCA y

SUÁREZ, 1985). En Huelva se ha encon-

trado en humedales del entorno del

Parque Nacional de Doñana y en las

cuencas de los ríos Tinto y Guadalqui-

vir, siempre asociado a macrófitos o sus-

tratos inertes de pequeña superficie.

Las especies del género Gyraulus

Charpentier 1837 que se encuentran en

la provincia son de carácter lenitófilo y

se asocian a limos y rocas de gran super-

ficie. Gyraulus albus (Fig. 2, C= 9,4%, A=

6,1%) se encuentra en las cuencas de los

ríos Guadiana, Odiel y Tinto desde

cabecera hasta desembocadura; Gyrau-

lus laevis (Alder 1838) (Fig. 2, C= 3,9%,

A= 3,9%) en las cuencas de Guadiana,

Odiel y Guadalquivir.

Hippeutis complanatus (Fig. 2, C=

1,5%, A= 0,8%) es una especie particu-

larmente común en aguas eutrofizadas

(CosTIL y CLEMENT, 1996). En Huelva se

encuentra en entornos leníticos y, en la

subcuenca del río Chanza, muy eutrofi-

zados, sobre macrófitos o sustratos

rocosos de la cuenca del Guadiana.

Ancylus fluviatilis O.F Múller 1774

(Fig. 2, C= 50,3%, A= 61,8%) es una es-

pecie paleártica muy difundida en la Pe-

nínsula Ibérica (VIDAL-ABARCA y SUÁ-

REZ, 1985), litófila y de marcado carácter

reófilo (GELDIAY, 1956), que muestra un

acusado polimorfismo en relación al ca-

rácter lótico o lenítico de los cursos de

agua donde reside (ANGELIER, 2002). En

la provincia de Huelva es la segunda es-

pecie en cobertura y abundancia, distri-

buyéndose desde cabecera hasta desem-

bocadura en corrientes lóticas o leníticas

de todas las cuencas analizadas; se ha

encontrado una elevada correlación pre-

sencia-ausencia entre esta especie y Phy-

sella acuta (correlación por rangos de

SPEARMAN, |R=0,829, p= 0,0001).

Myosotella myosotis (Fig. 3, C= 0,7%,

A= 0,8%) es una especie halófila atlán-

tico-mediterránea presente en costas,

marismas y salinas, que vive principal-

mente fuera del agua (KERNEY, 1999). En

la provincia se encuentra en marismas

mareales y desembocaduras de ríos,

sobre vegetación emergida o en limos y

barros de fondo o superficie de la

cuenca del río Piedras.

Potomida littoralis (Fig. 3, C= 0,7%,

A= 0,4%) es una especie con grandes

problemas de conservación derivados

de distribuciones muy localizadas y de

competencia con especies introducidas

(GÓMEZ ET AL., 2001), habiendo sido

propuesta por ello para su inclusión en

los listados de especies protegidas por el

Convenio de Berna (BOUCHET, FALKNER

y SEDDON, 1999). Se encuentra en aguas

leníticas sobre sedimentos finos de la

cuenca del Guadiana.

Unio pictorum (Fig. 3, C= 5,5%, A=

5,2%) es una especie ampliamente distri-

buida en la región paleártica (ELLIS,

1978) y muy abundante en la Península

Ibérica (VIDAL-ABARCA y SUÁREZ, 1985).

Se han encontrado ejemplares vivos en

las cuencas de los ríos Odiel y Guadiana,

siempre sobre lechos de limos y arcillas y

entre sustratos rocosos de pequeña su-

perficie (< 10 cm de diámetro mayor), y

PÉREZ-QUINTERO ET AL.: Moluscos de aguas continentales de Huelva (SO España)

++ AL

y +

AA Ha A.

Aaa:

AH E

— dale Los

+ S

Musculium lacustre

Corbicula fluminea

fon /

PC AH PC fa ee

LO TAO 74 O e

Ó 000 do £ '2

MOR o 2 LO DE E

MARES a COOP Dar

oe. $0 08 AH 0B

[a TT HIOON 4

El O |

Uno pictorum Anodonta cygnea

PC / Ele QC PC Mar “as

LL . TH

(e e 7 ADOS

/ LB ATA] HL 0B

aL ++ arg

PRA

QAN /

Pisidium casertanum O Pisidium personatum

O Pisidium milium

Figura 3. Mapas de distribución. Los círculos blancos indican la presencia de valvas vacías.

Figure 3. Distribution maps. White circles are empty shells or valves.

únicamente valvas vacías en la cuencas

de los ríos Guadalquivir y Tinto.

El amplio conocimiento de la biolo-

gía de Anodonta cygnea (Fig. 3, C= 2,3%,

A= 1,7) en relación con su reproducción

(GALHANO y FERREIRA, 1983), con su

papel como bioacumulador de metales

pesados (GUNDACKER, 2000) o con su

particular polimorfismo en relación a la

profundidad (MULLER y PATZNER, 1996),

muestra a este bivalvo como una de las

especies de moluscos dulceacuícolas

más estudiada de nuestro entorno pale-

ártico. Se han encontrado ejemplares

vivos únicamente en la cuenca del río

Guadiana, siempre enterrados en sedi-

mentos finos alternados con sustratos

rocosos de pequeña superficie.

La primera cita de Corbicula fluminea

(Fig. 3, C= 11,0%, A= 3,0%) en la Penín-

sula Ibérica es de MOUTHON (1981) en Por-

tugal; posteriormente PÉREZ-QUINTERO

(1990) la cita por primera vez en España,

asociada a la cuenca del río Guadiana en

la provincia de Huelva, y ARAUJO,

MORENO y RAMOS (1993) amplían su.dis-

tribución en el entorno europeo. Su enorme

potencial biológico, dispersivo y de resis-

tencia a las fluctuaciones del medio (ELDER

y COLLINS, 1991) hacen de esta especie

invasora una formidable competidora,

desplazando y abocando a extinciones

locales a la fauna autóctona, originando

igualmente serios problemas en sistemas

de conducción de aguas asociados a rega-

díos o consumo humano (STITES, BENKE y

GUILLESPIE, 1995), como sucede en la zona

regable del Chanza, Lepe, y en las con-

ducciones de agua potable de la ciudad

de Huelva (obs. pers.). Se encuentra en las

cuencas de los ríos Guadiana y Piedras,

en corrientes lentas y sustratos de sedi-

mentos finos, alcanzando densidades

superiores a 200 ind. /m2 en algunas zonas.

Musculium lacustre (Fig. 3, C= 2,3%,

A= 1,3%) ha sido encontrada a escasa

profundidad en corrientes lentas y entre

sedimentos finos, con dos intervalos alti-

tudinales bien diferenciados: 405 metros

(en una acequia de la cuenca del Guadal-

quivir) y 88 metros (en un humedal

cercano al Parque Nacional de Doñana).

De las tres especies del género Pisidium

Pfeiffer 1821 encontradas en la provincia

de Huelva, sólo Pisidium casertanum (Fig.

3, C= 7,8%, A= 8,7%) y Pisidium persona-

Iberus, 22 (2), 2004

tum (Fig. 3, C= 4,7%, A= 3,5%) registran

amplia distribución en la Península Ibérica,

mientras que Pisidium milium (Fig. 3) se

localiza en entornos concretos de la cuenca

mediterránea (KuIPER, 1961; VIDAL-

ABARCA y SUÁREZ, 1985). Todas ellas

residen en aguas de flujo lento, entre sedi-

mentos finos y a escasa profundidad (15-

85 cm), P. casertanum entre 116-740 m de

altitud, P. personatum entre 360-640 m,

ambas en la cuenca de los ríos Guadiana,

Odiel y Guadalquivir, y P. milium, de la

que no se han encontrado individuos vivos

sino solamente valvas, a 160 m de altitud

en la cuenca del Guadiana.

AGRADECIMIENTOS

Nuestros más sinceros agradeci-

mientos a la Dra. Ana M* Pujante Mora,

del Departamento de Biología Animal

BIBLIOGRAFÍA

ADAM, W., 1960. Faune de Belgique. Mollusques.

Tome 1, Mollusques terrestres et dulcicoles. Ins-

titut Royal des Sciences Naturelles de Belgi-

que, 402 pp.

ALTABA, C. R., TRAVESET, A., BOGUÑÁ, E. y

BEcH, M., 1985. Sobre la presencia de Ferri-

sia i Acroloxus (Gastropoda: Basommato-

phora) als paisos catalans. Butlletí de la Insti-

tució Catalana d Historia Natural, 52 (sección

Zoología, 6): 61-71.

ANGELIER, E., 2002. Ecología de las aguas co-

rrientes. Editorial Acribia, Zaragoza, 219 pp.

ANISTRATENKO, V. V., 1997. Is the european oc-

currence of Potamopyrgus (Gatropoda: Ta-

teidae) anthropochorous or autochthonous?.

Heldia, 4 (5): 141-142.

ARAUJO, R., MORENO, D. y RAMOS, M. A., 1993.

The asiactic clam Corbicula fluminea (Múller)

(Bivalvia: Corbiculidae) in Europe. Ameri-

can Malacological Bulletin, 10 (1): 39-49.

BARGUES, M. D., VIGO, M., HORAK, P., Dvo-

RAK, J., PATZNER, R. A., POINTIER, J. P., JAC-

KIEWICZ, M., MEIER-BROOK, C. y MAS-COMA,

S., 2001. European Lymnaeidae (Mollusca:

Gastropoda), intermediate hosts of tremato-

diases, based on nuclear ribosomal DNA

ITS-2 sequences. Infection, Genetics and Evo-

lution, 1: 85-107.

BARSIENE, J., TAPIA, G. y BARSYTE, D., 1998.

Chromosomes of Melanopsis dufouri Férussac

1823 (Caenogastropoda: Melanopsidae). Jour-

nal of Molluscan Studies, 64: 309-318.

de la Universitat de Valencia, por sus

consejos sobre el género Melanopsis en

general y la determinación de Melanop-

sis praemorsa; al Dr. Alberto Martínez-

Ortí, conservador de moluscos del

Museu Valencia d Historia Natural,

Valencia, por la determinación de Stag-

nicola palustris basada en el estudio de la

anatomía de su aparato reproductor; a

María del Mar Bayo Montoya, del

Departamento de Biología Vegetal y

Ecología de la Universidad de Almería,

por el envío de ejemplares de Mercuria

emiliana de su zona de estudio; a Javitxu,

Pepe, Laura y Poli por sus facilidades

para muestrear el Parque Nacional de

Doñana; a Miguel Angel Bravo, de la

Estación Biológica de Doñana, por facili-

tarnos gentilmente información biblio-

gráfica de los moluscos de la zona. A

Pilar y Carlos, por aguantar tantos

“caracoles”.

BINDER, E., 1967a. Position systématique de

Valvata minuta Drap., Valvata globulina Pala-

dilhe et d “autres petites especes attribuées au

genre Valvata. Atti Societa malacologica Ita-

liana, Milano, 105: 371-376.

BINDER, E., 1967b. La coquille embrionaire des

Valvatidae. Archiv fir Molluskenkunde, 96

(1/2): 21-24.

BIGOT, L. y MARAZANOEF, F., 1965. Considéra-

tions sur l'ecologie des invertébrés terres-

tres et aquatiques des marismas du Guadal-

quivir (Andalucia). Vie et Milieu. Serie C, 16

(1): 441-473.

BOETERS, H. D., 1988. Moitessieriidae und Hy-

drobiidae in Spanien und Portugal (Gastro-

poda: Prosobranchia). Archiv fiir Mollusken-

kunde, 118 (4/6): 181-261.

BOETERS, H. D., 1998. Mollusca: Gastropoda: Ris-

soidea. Súfswasserfauna von Mitteleuropa

5/1-2. Gustav Fischer, Stuttgart, 76 pp.

BOUCHET, P., FALKNER, G. y SEDDON, M. B., 1999.

List of protected land and freshwater molluscs

in the Bern Convention and European

Habitats Directive: are they relevant to con-

servation? Biological Conservation, 90: 21-31.

BROWN, D. S., 1980. Freshwater snails of Africa and

their medical importance. Taylor éz Francis,

London, 487 pp.

CLECOM-PROJECT 2002. Continental mollusca of the

western paleartic region (sections 1 € II of CLE-

COM): list of (sub) genera. Góteborg Natur-

historiska.

PÉREZ-QUINTERO £7 AL.: Moluscos de aguas continentales de Huelva (SO España)

CosTIL, K. y CLEMENT, B., 1996. Relationship bet-

ween fresh-water gastropods and plant com-

munities reflecting various trophic levels.

Hydrobiología, 321 (1): 7-16.

ELDER, J. F. y COLLINS, J. J., 1991. Freshwater mo-

lluscs as indicators of bioavailability and to-

xicity of metals in surface-water systems. Re-

views of Environmental Contamination and To-

xicology, 122: 37-79.

ELLIS, A. E., 1978. British Freshwater Bivalve Mo-

llusca. Synopses of the British Fauna, 11, 109

o G, RIPKEN, T. E. J. y FALKNER, M,,

2002. Mollusques continentaux de France. Liste

de Référence annotée et Bibliographie. Publica-

tions Scientifiques du Muséum National

d Histoire Naturelle, Paris, 350 pp.

GALHANO, M. H. y FERREIRA, M. T., 1983. The

reproductive cycle of Anodonta cygnea from

Mira Lagoon (Portugal). Publicacoes do Insti-

tuto de Zoologia “Dr. Augusto Nobre”, Facul-

dade de Ciencias do Porto, 179: 1-5.

GASULL, L., 1985. Fauna malacológica conti-

nental de la provincia de Huelva (España).

Miscelánea Zoológica, 9: 127-143.

GELDIAY, R., 1956. Studies on local populations

of freshwater limpet Ancylus fluviatilis Mú-

ller. Journal of Animal Ecology, 25: 389-402.

GIROD, A., BIANCHI, I. y MARIANI, M., 1980.

Gasteropodi, 1. En Rufo, S. (Ed. ): Guide per

il riconoscimento delle acque interne italiane,

7. Consiglio Nazionale delle Ricerche

AQ/1/44, Verona, 86 pp.

GLAUBRECHT, M., 1996. Evolutionsókologie und

Systematik am Beispiel von Siifi- und Brack-

wasserchnecken (Mollusca: Caenogastropoda:

Cerithioidea): Ontogenese-Strategien, palaónto-

logische Befunde und Historische Zoogeographte.

Backhuys Publishers, Leiden, 499 pp.

GLOER, P y MEIER-BROOK, C, 2003. Sússwasser-

mollusken. Deuster Jugendbund fúr Natur-

beobachtung, Hamburgo, 134 pp.

Gómez, B, MORENO, D., ROLÁN, E., ARAUJO, R.

y ALVAREZ, R. M., 2001. Protección de molus-

cos en el catálogo nacional de especies amenaza-

das. Sociedad Española de Malacología, Re-

señas Malacológicas XI, 286 pp.

GUNDACKER, C., 2000. Comparison of heavy

metal bioaccumulation in freshwater mo-

lluscs of urban river habitats in Vienna. En-

vironmental Pollution, 110: 61-71.

KERNEY, M., 1999. Atlas of the Land and Freshwater

Molluscs of Britain and Ireland. Harley Books,

England, 264 pp.

KUIPER, J. G. J., 1961. Contribution a la con-

naissance des speces du genre Pisidium vivant

en Espagne. Basteria, 25 (4/5): 54-67.

MAGURRAN, A, 1988. Ecological diversity and its

measurement. Princeton University Press, 192

EE:

MARAZANOF, F., 1966. Mollusques aquatiques

des marismas du Guadalquivir: dones éco-

logiques et biogéographiques. Annales de

Limnologte, 2 (3): 477-489.

MEDEIROS, L. y SIMOES, M., 1987. Contribuicao

para o estudo dos Gastropoda de agua doce

de Portugal II. Morfología e revisao siste-

mática de Planorbarius metidjensis (Forbes

1838). Garcia Orta, Série de Zoologia, 12 (1-2):

23-30.

MONTES DEL OLMO, C., 1993. Bases ecológicas

para la gestión integral del cangrejo rojo de la ma-

risma (Procambarus clarkii) en el Parque Nacio-

nal de Doñana. Tomo la Memoria. Departa-

mento Interuniversitario de Ecología, Uni-

versidad Autónoma de Madrid, 269 pp.

MOUTHON, J., 1981. Sur la présence en France

et au Portugal de Corbicula (Bivalvia, Corbi-

culidae) originaire d Asie. Basteria, 45: 109-

116.

MOUTHON, J. y DUBOS1S, P., 2001. Les peuple-

ments de mollusques de la zone littorale du

lac d'Annecy (Savoie, France). Annales de

Limnologte, 37 (4): 267-276.

MULLER, D. y PATZNER, R. A., 1996. Growth

and age structure of the swan mussel Ano-

donta cygnea (L) at different depths in lake

Mattsee (Salzburg, Austria). Hydrobiología,

341: 65-70.

MUÑOZ, B., 1992. Gasterópodos terrestres (Mo-

llusca, Gastropoda, Pulmonata) de Cáceres, Ba-

dajoz y Huelva. Tesis doctoral, Universidad

Complutense de Madrid, 303 pp.

ORTIZ DE ZÁRATE, A. y ORTIZ DE ZÁRATE, A.,

1961. Moluscos terrestres recogidos en la

provincia de Huelva. Boletín de la Real Socie-

dad Española de Historia Natural, 59: 169-190.

PÉREZ-QUINTERO, J. C., 1988. Faunística, distri-

bución y biometría de los moluscos dulcea-

cuícolas de la provincia de Huelva. Iberus, 8

(2): 203-209.

PÉREZ-QUINTERO, J. C., 1990. Primeros datos

sobre la presencia de Corbicula fluminea Mú-

ller (Bivalvia, Corbiculidae) en España. 1.

Biometría. Scientia Gerundensis, 16/1: 175-

182.

PONDER, W. F., 1988. Potamopyrgus antipoda-

rum, a molluscan coloniser of Europe and

Australia. Journal of Molluscan Studies, 54:

271-285.

PUJANTE, A. M., TAPIA, G. y MARTÍNEZ, F., 1998.

Los moluscos de los ríos de la Comunidad

Valenciana (España). Iberus, 16 (1): 1-19.

STITES, D. L., BENKES, A. C. y GUILLESPIE, D.

M., 1995. Population dynamics, growth and

production of the asiatic clam, Corbicula flu-

mínea, in a blackwater river. Canadian Jour-

nal of Fisheries and Aquatic Sciences, 52 (2):

425-437.

DY,

Iberus, 22 (2), 2004

TURNER, H, KUIPER, J. G. J., THEW, N., BERNAS-

-CONI, R., RÚETSCHI, J., WUTHRICH, M. y GOS-

TELL, M., 1998. Fauna Helvetica. Mollusca Atlas.

Centre suisse de cartographie de la faune-Sch-

weizerische Entomologische Gesellschaft,

Neuchátel, 527 pp.

VIDAL-ABARCA, C. y SUÁREZ, M. L., 1985. Lista

faunística y bibliográfica de los moluscos (Gas-

tropoda y Bivalvia) de las aguas continentales de

la Península Ibérica y Baleares. Asociación Es-

pañola de Limnología, 193 pp.

WAUTIER, J., 1977. Preliminary data on the geo-

graphical range of the freshwater limpet Fe-

rrisia wautieri. Malacología, 16 (1): 285-289.

WILKE, T., ROLÁN, E. y DAVIS, G. M., 2000. The

mudsnail genus Hydrobia s. s. in the nort-

hern Atlantic and western Mediterranean: a

phylogenetic hypothesis. Marine Biology, 137:

827-833.

PEREZ-QUINTERO ET AL.: Moluscos de aguas continentales de Huelva (SO España)

Anexo

Riveras muestreadas; en cada una de ellas aparece la referencia UTM de cada estación, la altura de

la misma en metros (A) y las especies encontradas (“individuos muertos). Junto al nombre de la

cuenca aparece la riqueza taxonómica de la misma, referenciada como (número de

especies/géneros/familias), y el índice de diversidad de Shannon-Weaver H.

Sampled rivers, with UTM references, high (A) and species found (* dead individuals). Between the

name of the basin, values of taxonomic richness (number of species/genera/lfamilies) and diversity index

(Shannon- Weaver H) are given.

Bt: Bithynia tentaculata; Pa: Potamopyrgus antipodarum; Ha: Hydrobia acuta; Pu: Peringia ulvae;

Me: Mercuria emiliana; Im: Islamia minuta; Mp: Melanopsis praemorsa; Sp: Stagnicola palustris; Gt:

Galba truncatula; Ra: Radix aff. auricularia; Rb: Radix balthica; Pt: Physella acuta; Bc: Bulinus

truncatus contortus; Pm: Planorbarius metidjensis, Ec: Ferrisia clessiniana; Pc: Planorbis carinatus;

Ga: Gyraulus albus; Gl: Gyraulus laevis; Hc: Hippeutis complanatus; Af. Ancylus fluviatilis, Mm:

Myosotella myosotis; Pl: Potomida littoralis, Up: Unio pictorum; Ac: Anodonta cygnea; Cf. Corbicula

fluminea; Ml: Musculium lacustre; Ps: Pisidium casertanum, Pp: Pisidium personatum, Pi: Pisidium

milium.

NOMBRE

Arroyo Fuente del Rey

Barranco Fuente el Castaño

Arroyo de la Parrilla

Barranco las Huertas

Arroyo Guijarra

Barranco del Lavadero

Barranco Navahermosa

Barranco del Toro

Barranco Fuente la Madrona

Arroyo Guijarra

Barranco Valdelamadera

Arroyo las Vegas

Arroyo Carboneras

Barranco del Ingenio

Charco de la Cuchara

Arroyo Corterrangel

Rivera de Hinojales

Arroyo Castañuelo

Rivera de Cala

Arroyo los Cocederos

Barranco Descansadero

Arroyo de Santa María

Arroyo de la Rocina

Arroyo de la Mayor

Arroyo de la Palmosa

Arroyo Madre del Avitor

Arroyo de la Rocina

Laguna del Moral

Laguna 12 del Martinazo

Charco del Navazo del Toro

Charca del Raposo

Laguna de Poli

UTM

295081798

295081298

295002514

295002613

2950B0999

295000801

29500401

295000809

295082196

295000901

295003009

295003010

2950B1599

29500201

295084299

295001101

295001906

295001202

295004305

2950B2590

295084497

2950B2024

295080816

2950B2818

295082517

295PB9115

295082412

295042100

295081868

295042299

295042297

295042198

CUENCA DEL GUADALQUIVIR (19/16/8) H= 3,49

A ESPECIES

7140 Pm, Af, Ps

680 Pm, Af

650 —Rb, Ps

640 Rh, Af

620 Rb, Pm, Ga, Af

620 Im, Rb, Pm, Ps

NOMBRE

Arroyo Borbolluela

Rivera de Cala

Rivera de Huelva

Rivera de Montemayor

Rivera de Cala

620 Im, Gt, Rb, Pm, Af, Ps Arroyo las Casetillas

570 Pm, Af

570 Pm

560 Af

535 Mp, Pt, Pm, Af

530 Mp, Rb, Af, Up, Pp

520 A

520 Pm,Ps

520 Pm

485 Af, Pp

465 Pi

460 Ra, Rb, PL, Af

455 Pt, Fc, Gl, Af, Up, Ac

450 Gt, Pt, Pm, Af

450 Pt, Pm

0 ll

00 nt

ZONE

ZO Bt

(RON

Ey lA

2,8 Pt

2,1 Pt

Barranco Maygalanes

Arroyo del Rey

Rivera de Hierro

Rivera de Huelva

Arroyo Alcarayón

Arroyo de la Cerrada

Arroyo de don Gil

Arroyo Barbacena

Arroyo del Avispero

Arroyo de la Cañada

Arroyo del Saltillo

Arroyo de Tejada

Arroyo del Algarbe

Arroyo de Calancha

Charca del Brezo

Caño las Gangas

Caño del Martinazo

Veta del Martinazo

Laguna de Santa Olalla

Laguna del Taraje

Charca de Wouter

Laguna Dulce

Lucio del Bolín

Charca Raya Pinar

Laguna Fría

UTM

2950C0704 430

295004702 430

2950C0803 420

295002009 405

Barranco Fuente del Puerto295QB2098 400

295085096 360

2950B2199 350

2950B2989 350

2950B2885 330

2950B2895 270

2950B2895 260

2950B2842 110

2950B1834

295PB9921

295083552

295080526

295081522

295081825

295083348

2950B3331

295081627

295042196

2950B2279

2950B1178

2950B1683

295042495

295042296

295042496

295042395

295042796

2950B2597

295042397

A ESPECIES

Pt, Fc, Gl, Af, Up, Ac

Gt, Pt, Af

Pt, Af, Up”, Ac”

Mp,Ra,Rb,Pt,Pm,Af Up,MI,Ps

Pm, Ga

Pt, Fc, Af

Im, Mp, Rb, Pt, Pm, Af

Pm, Ga, Af

Rb, Pt, Gl, Af, Up”

Pm, Ga, Af

Pt, Af, Up

Pt, Up”, Ac”

90 Pt

718 Pt

15 Gt, Pt, Pm

15 Pt, Fc, Gl

OSGUADE

54 Pt

50 Pt, Af Up"

50 Pt

40 Pt

2,1Pt

2,0Pt

2,0Gt, Pt, Bc, Pc

1,9Pt, Fc

1,9Pt

1,6Pt

1,5Pt

0,7Pt

NOMBRE

Rivera de la Adelfilla

Rivera de las Majadillas

Rivera delos Pinos

Arroyo Tamujoso

Rivera del Jarrama

Arroyo el Barrancoso

Rivera Cachán

Arroyo Bajohondo

Arroyo Gallego

Rivera Casa de Valverde

Arroyo del Pinar Serrano

Arroyo Tamujoso

Río Corumbel

Arroyo Gallinero

Rivera del Honueco

Arroyo de Trigueros

Arroyo de la Fuentidueña

Arroyo de Pozo Áncho

Arroyo de la Bárcena

Arroyo el Guijillo

NOMBRE

Rivera de Alájar

Rivera de Almonaster

Barranco la Umbría

Arroyo de Marimateos

Barranco de la Lana

Barranco los Casares

Rivera de las Molinillas

Barranco del Acebuche

Rivera de Santa Ana

Barranco el Cabezuelo

Barranco de Calabazares

Arroyo Plamencia

Barranco los Madroñeros

Rivera de Almonaster

Rivera de las Molinillas

Barranco la Cabra

Rivera del Villar

Río Vanegas

Rivera de Santa Ana

Arroyo del Tejarejo

Barranco el Moro

NOMBRE

Barranco la Higueruela

Rivera de Jabugo

Arroyo Sillo de Cumbres

Rivera de Jabugo

Barranco el Buho

Barranco la Urraca

Arroyo Gargallones

Barranco el Puerto

UTM

295081983

295081879

295080664

2950B1372

2950B2175

295080652

295081368

295080061

2950B1866

295081057

2950B0127

295081851

295082450

295081851

2950B1663

295PB9444

295082045

295PB9244

295080243

UTM

295080594

295PB9594

2950B2393

295081895

295PB9794

295080194

295081295

295PB9295

295080293

2950B0494

295PB9793

2950B0994

2950B0693

295PB9392

2950B0993

295PB9991

2950B0774

295081687

295080589

295081586

295PB9091

UTM

295PB9898

295080098

295001015

295080099

295PC9303

295080299

295PC8801

295PB9597

700

640

600

580

560

540

500

460

440

380

360

330

320

CUENCA DEL GUADIANA-PIEDRAS (23/19/9) H= 3,75

600

580

540

480

Iberus, 22 (2), 2004

CUENCA DEL TINTO (12/12/6) H= 2,19

ESPECIES

Pm, Ga, Af

NOMBRE

Arroyo de Pasadera

Arroyo la Peñuela

Arroyo del Arzobispo

Arroyo de Sapo Hondo

Arroyo de la Grulla

Arroyo Clarina

Arroyo Giraldo

Arroyo Lavapies

Arroyo Candón

Rivera de Nicoba

Arroyo del Puerco

Arroyo Lavapiés

Rivera de Nicoba

Arroyo del Puerco

Arroyo del Moro

Arroyo Candón

Estero Domingo Rubio

Arroyo de Freire

Rivera de Nicoba

CUENCA DEL ODIEL (15/13/5) H= 3,04

ESPECIES

Im, Sp, Pt

Pa, Im, Rb, Af, Pp

Pt, Pm, Ga, Af

Gt, Rb, Pm, Af, Ps

NOMBRE

Rivera Escalada

Rivera la Pelada

Río Odiel

Rivera del Vaho

Rivera de Santa Eulalia

Im, Gt, Rb, Pt, Pm, Gl, Af Rivera Escalada

Im, Af

Gt, Rb, Pt, Pm, Af

Rb, Af, Ps

Im, Rb, Af, Ps

Rb, Af, Ps

Rb, Pm, Gl, Af

Pa, Im, Rb, Af

Im, Af

Rb, Pm, Af, Ps, Pp

Pt, Af

Pt, Fc, Gl, Af

Gt, Rb, Af, Ps

Pt, Fc, Af

Rb, Af

ESPECIES

Af, Ps

Im, Gt, Rb, Pm, Af, Ps, Pp

Im, Rb, Af

Im, Rb, Pm, Af

Rb, Pm, Af, Ps, Pp

Rb, Af

Rb, Af, Ps, Pp

Rivera del Villar

Arroyo de la Burrilla

Rivera Olivargas

Barranco la Fresnera

Barranco del Tamujoso

Barranco de los Pinos

Barranco de los Ovejeros

Arroyo Galaperosa

Barranco de la Sepultura

Barranco de Juré

Arroyo de las Multas

Arroyo Carrasco

Arroyo Monte de la Osa

Arroyo Chapinero

Arroyo del Encinar

NOMBRE

Rivera del Chanza

Arroyo Arochete

Río Múrtigas

Rivera de los Ciries

Rivera Peramora

Rivera del Chanza

Arroyo Sillo

Rivera Piernaseca

UM A

295080130 83

2950B0344 80

295080938 70

2950B1140 65

295PB9621 59

295081144 50

2950B1643 45

2950B0439 45

295080040 35

295PB8939 35

295PB8439 25

2950B0437 25

295PB8737 25

295PB8439 22

295080336 15

295PB9935 9

295PB9834 7

295PB8820 7

295PB8427 4

295PB8432 4

UTM A

295PB9790 320

295PB8386 320

295081686 310

295PB0087 281

295080587 280

295PB9887 275

2950B0073 235

295PB5654 210

295PB9084 210

295PB7583 200

295PB8976 185

295PB8179 180

295PB7375 160

29SPB8562 150

295PB7174 140

295PB6857 120

295PB6549 100

295PB9260 90

295PB7465 85

29SPB7567 80

295PB8242 35

UM A

295PC8103 270

295PC8005 260

295PC9115 260

295PC7596 260

295PC7597 260

295PC7806 255

295PC8818 240

295PB5790 240

ESPECIES

Pt, Af

Pt, Pm

Rb, Pt, Pm, Pc, MI

Pt, Af

Pt, Up”

Up"

Af, Up"

Pm, Ga

Pt, Up"

Pt, Pc

Gt, Pt

Bt, Pt, Ac”

ESPECIES

Rb, Pm, Af

Pt, Pm, Fc, Af

Rb, Pt, Fc, Af, Up

Pm, Af

Rb, Af, Up"

Pt, Af

Pt, Af, Up”

Pt, Pm, Af

Pt, Af

Im, Af, Ps

Pt, Af

Pt, Pm, Af

Pt, Af

Pt, Pm, Af

Pt, Af

Pt, Af, Up"

Pt, Pm, Af

ESPECIES

Pt, Af

Pt, Fc, Af, Up"

Pt, Af

Gt, Af, Pt, Up", Ac”

Pt, Fc, Af

Pt, Up”, Ac”

Pt, Af

PEREZ-QUINTERO ET AL.: Moluscos de aguas continentales de Huelva (SO España)

Arroyo del Cavá

Rivera Caliente

Río Múrtigas

Barranco las Murtiguillas

Barranco Riofrío

Barranco Menjuana

Barranco la Extremedera

Barranco Riofrío

Barranco Valdesotello

Barranco de la Villa

Barranco los Cubos

Rivera del Chanza

Barranco de Monteviejo

Barranco la Buharda

Rivera de la Espada

Rivera del Aserrador

Arroyo del Colmenar

Barranco del Fraile

Barranco el Arroyo

Rivera de Calaboza

Barranco Redondillo

Rivera del Chanza

Rivera Matavacas

Arroyo del Alamillo

Rivera Agua de Miel

Arroyo de la Poricona

Rivera de la Ronchona

Rivera de la Golondrina

295PC8625

295PC9604

29SPC9705

29500006

295PC9911

295009910

295009809

29SPC9809

295PC7808

295PC7902

295PC8203

295PC8202

295PB5482

298PC5503

295PB5455

295PB5694

295PB4944

295PB4779

295PB4776

295PB5898

295PB4261

295PB3979

295PB4248

295PB4632

295PB5774

29SPB4635

295PB3853

295PB4051

400

380

360

350

320

310

285

280

215

165

160

150

140

120

110

Pt, Af

Af, Up”, Ps, Pp

Af, Ps

Pt, Af

Pt, Pm, Ga, Af

Rb, Pt, Pm, Ga, Af

Pt, Af, Up”

Pt, Af

Gt, Rb, Pt, Pm, Fc, Af Arroyo Albahacar de Allá

Pt, Af

Gt, Rb, Pt, Pm, Af, Up”, Ac”

Pt, Pm, Af

Pt, Pi

Pt, Af

Gt, Af, Up"

Rb, Pt, Af

Pt, Af

Pt, Fc, Af

Af, Up”, Ac”

Pt, Af

Pt, Af, PI, Up

Pt, Af

Pt, Pm, Ga, Af

Pt, Af

Pm, Ga, Af

Pt, Pm, Af

Bt, Pt, Pm, Ga, Af, Up, Cf

Arroyo Valquemado

Río Múrtigas

Rivera Charcolino

Rivera del Cañuelo

Arroyo las Cañas

Arroyo Petaquera

Rivera del Chanza

Rivera del Malagoncillo

Arroyo de Agualobos

Rivera de Malagón

Arroyo de los Arroyillos

Rivera de la Ronchona

Arroyo del Contrapeso

Arroyo Grande

Arroyo de la Gitana

Barranco de la Chacera

Arroyo Tariquejo

Rivera Grande

Río Piedras

Arroyo del Pilar

Arroyo Grande

Arroyo de la Vera

Arroyo Puentezuelo

Marismas del Piedras

Canal del Piedras

Arroyo del Prado

Barranco Huerto Torres

295PC8122 220

295PC8222 220

295PB6978 206

295PB6778 205

295PC5804 190

295PB4951 180

295PC6304 180

295PB6472 180

295PB4776 175

295PB5180 170

295PB5173 170

29SPB5565 165

295PB3550 35

295PB3354 35

295PB4430 30

295PB4726 25

295PB3936 20

295PB6129 15

295PB3747 15

295PB6128 10

295PB6428 8

295PB4028 8

295PB5720 7

295PB6227 7

295PB6220 0

29SPB5220 5

29SPB3650 5

Pt, Af, Up"

Pt, Pm, Af

Pt, Af, Up"

Pt, Fc

Pt, Af

Rb, Af, Up"

Pt, Af

Pt, Af, Up"

Pt, Af

Bt, Pt, Af, PI, Up, Ac

Pt, Af

Cf, Pt

Pt, Af, Up”, Ac”

Pm, Af

Me, Af

Me, Gt, Af, Pl, Up, Ac, Cf

Ra, Rb, Pt, Pm

Pt, Fc, Af, Up, Ac, Cf

- Rb, Cf

Pt, Bc

Me, Af

O Sociedad Española de Malacología —_—_—_—_———T— Iberus, 22 (2): 33-44, 2004

Fatty acids of Antarctic gastropods: distribution and com-

parison with Mediterranean species

Acidos grasos en gasterópodos antárticos: distribución y compara-

ción con especies mediterráneas

Conxita ÁVILA*”, AmecloBONTANA Mauro, ESPOSITO?* Maria

Letizia CIAVATTA** and Guido CIMINO**

Recibido el 26-1-2004. Aceptado el 22-V-2004

ABSTRACT

Fatty acids of three different lipid pools: free fatty acids (FFA]), storage lipids (triglycerides

and wax esters, SL) and phospholipids (PL) of mantle and viscera of Antarctic gastropods

were analyzed and compared to species from the Mediterranean Sea. We analyzed spec-

imens of the Antarctic species: Bathydoris clavigera Thiele, 1912, Tritonia challengeriana

Bergh, 1884 and Marseniopsis mollis (Smith, 1902), and the Mediterranean species:

Hypselodoris picta (Schultz, 1836) and Dendrodoris limbata (Cuvier, 1804). Fatty acid

composition was very different between viscera and mantle of the same individuals, and

the amounts of polyunsaturated fatty acids were significantly higher in the mantle. There

were higher levels of polyunsaturated fatty acids in mantle phospholipids of Antarctic mol-

luscs than in Mediterranean molluscs. Arachidonic and eicosapentaenoic acids were the

dominant species of phospholipids in Antarctic molluscs, whereas octadecaenoic acid was

the most abundant species in the phospholipid pools of Mediterranean animals. A compar-

ison of the SFA/PUFA (saturated vs. polyunsaturated fatty acids) and MUFA/PUFA

(monounsaturated vs. polyunsaturated fatty acids) indexes in SL and PL of Antarctic and

Mediterranean specimens showed statistically significant differences among them, thus

suggesting a relationship with environmental temperature.

RESUMEN

Se analizan los ácidos grasos de tres tipos de lípidos, ácidos grasos libres (FFA), lípidos

de reserva [triglicéridos y ésteres de ceras, SL) y fosfolípidos (PL), en el manto y las vísce-

ras de gasterópodos antarcticos, y se comparan con especies mediterráneas. Las especies

estudiadas fueron las antarcticas Bathydoris clavigera Thiele, 1912, Tritonia challenge-

riana Bergh, 1884 y Marseniopsis mollis (Smith, 1902), y las mediterráneas Hypselodo-

ris picta (Schultz, 1836) y Dendrodoris limbata (Cuvier, 1804). La composición de ácidos

grasos difirió mucho entre vísceras y manto de algunos especímenes, y la cantidad de aci-

dos grasos poliinsaturados fue significativamente más elevada en el manto. Se encontra-

ron mayores niveles de éstos últimos en el manto de las especies antarcticas que en el de

las mediterráneas. Los ácidos araquidónico y eicosapentanoico fueron los dominantes en

los PL de las especies antarcticas, en los PL de las especies mediterráneas lo fue el ácido

* Centre d'Estudis Avangats de Blanes (CEAB), C.S.I.C., c/ Accés a la Cala Sant Francesc 14, 17300 Blanes,

Girona, Spain.

** Istituto per la Chimica di Molecole di Interesse Biologico (ICMIB) del CNR. Via Campi Flegrei, 34,

Comprensorio Olivetti, 80078 Pozzuoli (NA), Italy.

** Istituto Zooprofilattico Sperimentale del Mezzogiorno (IZSM). Via Salute 2, 80055 Portici (NA), Italy.

' Corresponding author. E-mail: conxitafceab.csic.es. Fax number: 34-972-337806.

Iberus, 22 (2), 2004

octadecanoico. La comparación entre los índices SFA/PUFA (ácidos grasos saturados vs.

poliinsaturados) y MUFA/PUFA (ácidos grasos monoinsaturados vs. poliinsaturados) de

los SL y PL de los dos grupos de especies mostró diferencias estadísticamente significativas

entre ellos, lo que sugiere una relación con la temperatura ambiental.

KEY WORDS: fatty acids, Antarctic, gastropods, opisthobranch molluscs, chemical ecology.

PALABRAS CLAVE: ácidos grasos. Antártica, gasterópodos, moluscos opistobranquios, ecología química.

INTRODUCTION

Fatty acids in invertebrates are

known to have broad biological roles,

including lipid energy reserves, compo-

nents of cellular structures such as bio-

membranes, and regulation of biosyn-

thesis of eicosanoids, among others.

Environmental conditions, such as diet

or temperature, are closely related to

lipid metabolism and may modulate the

activities of the membrane (VooGr,

1983; CULLIS AND HOPE, 1991; URICH,

1994; NELSON, LEIGHTON, PHLEGER AND

NICHOLs, 2002). In fact, the physical

properties of cell membranes are

affected by even minor variations in the

proportions of phospholipids, glycol-

ipids, sterols and fatty acids (CULLIS

AND HOPE, 1991; UricH, 1994). Also, it is

well known that poikilotherms alter

their membrane lipid composition in

response to varying environmental tem-

perature (UrICH, 1994; MOON, HIGASHI,

ZOLTAN AND MURATA, 1995; CHAKKOD-

ABYLU AND THOMPSON, 1984; DEY, BUDA,

WIIK, HALVER AND FARKAS, 1993; HALL,

THOMPSON AND PARRISH, 2000; FARKAS,

FODOR, KITAJKA AND HALVER, 2001). The

most common changes involve re-tailor-

ing of phospholipid heads, sterol

content of membranes and unsaturation

of phospholipid fatty acids (URICH,

1994; CHAKKODABYLU AND THOMPSON,

1984; DEY ET AL., 1993). One rational

explanation of these variations is that

cells compensate the temperature

decrease with the increase of the mem-

brane fluidity (e.2. the content of unsatu-

rated fatty acids in cell membranes

becomes higher at lower temperatures).

For marine organisms, the temperature-

dependent composition of membrane

lipids has been reported in some bacte-

ria (ROTERT, TOSTE AND STEIER, 1993;

SAKAMOTO, HIGASHI, MURATA AND

BRYANT, 1997; QUOC AND DUBACO, 1997;

RUSSELL, 1998; RUSSELL AND NICHOLs,

1999), unicellular algae (SATO, MURATA,

MIURA AND UETA, 1979; THOMPSON,

GUO, HARRISON AND WHYTE, 1992;

LEHMAL, 1999), cnidarians (e.g. CAR-

BALLEIRA, MIRANDA AND RODRÍGUEZ,

2002), molluscs (KATTNER, HAGEN,

GRAEVE AND ALBERS, 1998; GILLIS AND

BALLANTYNE, 1999; FREITES, LABARTA

AND FERNÁNDEZ-REIRIZ, 2002), and fish

(HAZEL, 1984; DEY ET AL., 1993).

Gastropods are one of the most

diverse animal groups, both in form,

behavior and habitats (PONDER AND

LINDBERG, 1997), and their evolutionary

success could be related, among others,

to their extraordinary ability to become

adapted to different environments. This

makes these animals particularly suit-

able for investigating the temperature

effect on the cellular homeostasis. Gas-

tropods have often been chemically

studied for their ability to produce sec-

ondary metabolites with ecological sig-

nificance or potential use as drugs or

pharmacological tools (IRELAND, COPP,

FOSTER, MCDONALD, RADISKY AND

SWERSEY, 1993; ÁVILA, 1995; SHu, 1998),

but surprisingly, they have received

little attention regarding the effect of the

environmental conditions on their lipid

levels (URICH, 1994; KATTNER ET AL.,

1998; Isay AND BUSAROVA, 1984). Actu-

ally, there are many descriptive studies

on the primary lipids of prosobranchs

(see VOOGT, 1983), but there have been

almost no investigations on opistho-

branchs. In fact, only some very early

studies are reported on Aplysia kurodai

ÁVILA ET AL.: Fatty acids of Antarctic gastropods

Table I. Data on the specimens studied quantitatively.

Tabla 1. Datos de los especímenes estudiados cuantitativamente.

No. of specimens

Species analyzed Depth (m)

Bathydoris clavigera 1 462

Iritonia challengeriana 1 446

Marseniopsis mollis 1 221

Hypselodoris picta 3 2-12

Dendrodoris limbata 2 2-12

(Baba, 1937) (TANAKA AND TOYAMA,

1959), Aplysia fasciata Poiret, 1789 and

Pleurobranchaea meckeli Meckel in Leue,

1813 (TIBALDI, 1966), and more recently

on the pteropod Clione limacina (Phipps,

1744) (KATTNER ET AL., 1998).

During our ongoing research on

natural products of marine inverte-

brates, we repeatedly observed high

contents of fatty acids in extracts of

Antarctic organisms, mainly opistho-

branch molluscs and sponges. There-

fore, we decided to further investigate

this fact by carrying out a comparative

study on the fatty acid content in

Antarctic and Mediterranean gas-

tropods. We selected two opisthobranch

species and a single prosobranch species

from Antarctica, and two Mediterranean

opisthobranch species. We report here

the composition and tissue distribution

of their fatty acids in three different

lipid pools: phospholipids (PL), storage

lipids (SL, i.e. triglycerides and wax

esters), and free fatty acids (FFA).

MATERIALS AND METHODS

Materials

The species studied here were the

opisthobranchs Bathydoris clavigera

Thiele, 1912 and Tritonia challengeriana

Bergh, 1884 and the prosobranch Marse-

niopsis mollis (Smith, 1902), from Antarc-

tica, and the opisthobranchs Hypselodoris

picta (Schultz, 1836) and Dendrodoris lim-

bata (Cuvier, 1804) from the Mediter-

ranean (Table 1). The species T. challenge-

riana had been named Marioniía cucullata

y Wet (w) or Dry

Geographic area Size (cm) (8) weight (a)

Weddell Sea 9.5 45 w

Weddell Sea 5 3.5 W

Weddell Sea 4.5 22.6Ww

Mediterranean Sea 1.510 1.8-2.8 d

Mediterranean Sea 5-8 0.8-1.2 d

(Couthouy in Gould, 1852) in the past,

but its taxonomy has been recently re-

vised by MUNIAÍN AND SCHRÓDL (1999).

Mediterranean species were selected due

to our previous knowledge of their

chemical ecology and their availability,

while Antarctic species were selected be-

cause of their availability and their size

(which should be large enough to allow

for chemical analysis).

Preliminary qualitative studies were

carried out with several specimens of

these and other species not reported

here, in order to test and improve the

chemical methodology. Antarctic speci-

mens used in this quantitative study

were collected during the German expe-

dition ANT XIIM/3 (EASIZ I) to the

Eastern Weddell Sea in January 1996

(Table D) (ARNTZ AND GUTT, 1997). The

Mediterranean specimens were col-

lected by scuba-diving in the Gulf of

Naples (Italy) in June 1996 (Table 1). All

biological samples were immediately

frozen and kept at -30C until the chem-

ical analyses were performed.

Dissection and extraction of mol-

luscs

The specimens were carefully dis-

sected in order to separate mantle and

viscera. Since only a limited number of

the Antarctic specimens was available,

we used sub-samples from the animals

in order to have pseudo-replicates. We

analyzed two sub-samples for T. challen-

geriana and three for the other two

Antarctic species, and two sub-samples

for each type of tissue analyzed. Each

body section was extracted separately

Iberus, 22 (2), 2004

Table II. Content (ug per mg of total lipid extract) of SL, PL and FFA in mantle and viscera of the

studied gastropods.

Tabla II. Contenido (ug per mg del extracto lipídico total) de SL, PL y FEA en manto y vísceras de los

gasterópodos estudiados.

Mantle Viscera

Hypselodoris picta es

Dendrodoris limbata 16.1 194

Marseniopsis mollis 344.8 58.0

Bathydoris clavigera 208.3 146.5

Iritonia challengeriana 2428 242.0

by following the method of Bligh and

Dyer (HAMILTON, HAMILTON AND

SEWELL, 1992). The sample (ca. 8.5 mg)

was homogenized by blending with a

mixture of CHCl3 (8.5 ml) and MeOH

(17 ml) for two minutes. Then 8.5 ml of

CHCIl were added to the solution and

blended for 30 sec. more. Distilled water

(8.5 ml) was added to the solution and

the mixture was blended again for 30

sec. The suspension was filtered

through paper on a Buchner funnel and

the filtrate was recovered. The residue

was transferred into the blender and the

extraction was repeated. The CHCIs3

layer of the combined filtrates was sepa-

rated and dried at reduced pressure to

give a lipid extract containing glyc-

erides, fatty acids, sterols and phospho-

lipids.

Fractionation of the lipid extracts

The CHCl3 soluble fractions were

separated by Si02 column (typically 70

mg silica per 1 mg of extract). Briefly, the

lipid extract was solved in petroleum

ether (typically 50 pl per 1 mg of extract)

and loaded onto a column. Then 0.1 ml

of 37% aqueous ammonia were added to

the column and elution started with

petroleum ether /diethyl ether (95:5 v/v)

to give triglycerides, wax esters and

sterols. Then glacial acetic acid (0.1 ml)

was added to the column and the elution

was completed by diethyl ether and a

mixture of chloroform /methanol/water

(60:40:2 v/v), to give free fatty acids and

polar glycerides, respectively. The

PL FFA

Mantle Viscera Mantle Viscera

14.6 31.0 44.4 1508

90.3 1193 80.6 47.0

293.1 265.4 193.1 61.2

80 338.4 ZO es

1505 220.9 250.0 35.1

results of the separation by the SiO»

column were checked by TLC for accu-

racy. In order to be further analyzed, the

samples were transformed into fatty acid

methy]l esters (FAMESs).

Preparation of FAMEs

Fractions containing free fatty acids

were concentrated at reduced pressure

to a volume of 0.5 ml and reacted in

open vials with a saturated solution of

CH2N2 in Et20 (0.5 ml). The reaction

was performed for 30 min at room tem-

perature. The excess of CH2N2 was

removed by bubbling a stream of nitro-

gen. Solutions were concentrated under

nitrogen flow to ca. 0.3 ml and analysed

by GC-MS under the analytical condi-

tions reported below.

Phospholipids and storage lipids

(triglycerides and wax esters) were con-

verted to FAMEs by a base catalyzed

transesterification with Na2C0O3 in dry

methanol. Briefly, lipid samples were

transferred to graduate screw-top vials

and treated with 1.5 ml of saturated

sodium carbonate in dry methanol. The

reaction solution was heated at 40C for

2 h, cooled at room temperature, trans-

ferred to a separating funnel and

extracted with 5 ml diethyl ether and 8

ml of brine. The upper phase was

removed and the extraction was

repeated three times. The organic layers

were combined, reduced to small

volume (ca. 1 ml) and analyzed by GC-

MS under the conditions reported

below. We did not separate triglycerids

ÁVILA ET AL.: Fatty acids of Antarctic gastropods

Table III. Relative amounts (mean % w/w + SD) of the main fatty acids identified in mantle sec-

tions of Antarctic molluscs. SL: storage lipids (triglycerids and wax esters). PL: phospholipids.

EFA: free fatty acids. nr: below the measurement limit. -: this sample could not be analyzed.

Tabla 11. Cantidad relativa (porcentaje medio del peso húmedo + SD) de los principales ácidos grasos

identificados en las secciones del manto de moluscos Antárticos. SL: lípidos de almacenamiento (triglicé-

ridos y ésteres de ceras). PL: fosfolípidos. FFA: ácidos grasos libres. nr: por debajo del límite de resolución.

-: muestra no analizada.

Marseniopsis mollis

SL PL FFA SL

16:0 28.8+7.2 16.245.5 5.0+1.9

17:0 8.5+3.1 4.8+2.8 1.1£0.7

18:0 22.1£6.7 6.121 3.3+1.4

16:1 w7 4.5+2.1 0.6+0.2 1.2+0.5

18107/09 14.5+44 7.1430 5.3+1.4

20:1 11.0+3.9 14.145.6 12.2+6.5

18:2 m6 nr nr nr

20:4 m6 1.0+0.4 5.6+1.8 26.6+3.1

20:5 w3 2.8+1.3 29.2+8.6 15.9+4.9

22:5 06 nr 4.9+1.5 5.5+2.1

22:6 m3 1.4+0.8 6.5+2.4 19.7+3.4

from wax esters, since this was not the

aim of this study.

GC-MS analysis

Analysis of FAMEs was carried out

on a Fisons MD800 Mass Spectrometer

coupled to a Fisons GC8000 Chromato-

graph equipped with a JW Scientific

DB5-MS column (30 m x 0.25 mm x 0.25

pm). Helium was used as carrier gas at a

flow rate of 1 ml/min. Each sample (1

ul) was injected in split mode (1:20). The

oven temperature was programmed ini-

tially at 100%C for 3 min, and then

increased to 300"C at 3C/min; the

injector and the transfer line tempera-

tures were 260”"C and 240"C, respec-

tively. Mass spectra were recorded in

continuous scan mode from 50 to 450

u.m.a. with an ionization current of 70

eV; the source temperature was set at

200*C. FAMEs were identified by both

retention time, previously determined

on a standard mixture with alkyl chains

from C-12 to C-24, and by library-

assisted interpretation of mass-spectra.

Percentages were measured by analysis

of the peak areas in the chromatogram,

by using HP G1034C Chemstations soft-

Bathydoris clavigera Iritonia challengeríana

PL FFA SL PL FFA

OE Sp ozl nr 1.340,4

OOO 0:22 A0:90:6, C0/6+072

9183.10 06412401 8.8524: 6.0£3.2 7.6428

9.0:2.8 38:10 28.5+/.7 13.9+3.6 14.8:4.4

0020 oz 0 tol oi0.2. 1 9734

9.1428 10.441 10.4x5.1 10,2:2.8 9.5+3.6

aldo 007) nr nr 1.140.4

13.844.1 22.216.3 20.117.2 43.817.9 40.9+5.4

A A o

104 3:94 03:27 050149 16:053.37 01:02

BOE IS LOAUZO AA Ole

ware. The results were expressed as rel-

ative percentages (% w/w) of the total

fatty acid content, and were compared

by using t-tests to determine statistically

significant differences.

RESULTS

The total lipid content in the viscera

was similar in all the studied animals,

with means of 28.0+2.4 mg/g of wet

tissue in Antarctic samples and 25.2+3.3

mg/g in Mediterranean samples. In the

mantle, the total lipid content was con-

sistently higher in the Mediterranean

molluscs (15.8+3.1 mg/g of wet tissue)

respect to the Antarctic ones (3.2+0.4

mg/g of wet tissue). Three different

lipid pools: free fatty acids (FFA),

storage lipids (SL) and phospholipids

(PL) were considered for each body part

(Table II). The fatty acid composition of

FFA, SL and PL was determined from

mantle and viscera, and it is reported

here for the Antarctic molluscs (Tables

III and IV). Although fatty acid compo-

sition of FFA, SL and PL was not very

different, the relative percentages varied

Iberus, 22 (2), 2004

Table IV. Relative amounts (mean % w/w + SD) of the main fatty acids identified in the viscera of

Antarctic molluscs. SL: storage lipids (triglycerids and wax esters). PL: phospholipids. FFA: free

fatty acids. nr: below the measurement limit.

Tabla IV. Cantidad relativa (porcentaje medio del peso húmedo + SD) de los principales ácidos grasos

identificados en las vísceras de moluscos Antárticos. SL: lípidos de almacenamiento (triglicéridos y ésteres

de ceras). PL: fosfolípidos. FEA: ácidos grasos libres. nr: por debajo del límite de resolución.

Marseniopsis mollis Bathydoris clavigera Tritonia challengeriana

SL PL FFA SL PL FFA SL PL FFA

16:0 5.2414 45418 2.2£1.2 29:20 4.3107, 2.5£0.6- —16.7£3.1 "O OS ino

17:0 5,341.4 8.7£0.6 0,420.17 0.7:0.2 * 1.8£06- 05:01 TESEI ES

18:0 12.4:0.3 29.610.6 3.6+1.6— 9.1£0,4- 14,2+1.8- 4.3£0.9 5:9£1.159/ 14220352200

16:1 07 4.7:x0.2 4.541.3 T.120.7 3.3204 26:10" 6.941.4 ""30.3£5:0 12 DES P2O mE

18:1w7/09 135+1.0 17.9:0.9 18.7:1.0 18.3:0.8 12.0:1.1 15.341.4 23.4:2.1-1 05145456

20:1 24.222.2 17.243.2 13.308 15.541.060 18.7:1.4 16.206. — 2.340.900 40-19 SONDA

18:2 06 4.0:0.9 5.411.1 —3.0:0.8 '-5.3:0.5 3.8:0,3 1.1:0.5* 8.9523 1 Moss 000

20:4 006 BUEDI" 1.910.952 LS Ml el2 7 9/920.9 1 9:32100 nr 16.9+1.8 10,3+1.5

20:5 03 9.6114 4.6%0.6 20.221 10.8:0.9 8./+0.9 11.4+0.8 nr 69:21 GEN

22:5 06 4.0+1.5 nr TZ 20.9 TODAS nr 1.6+0.3 1.5+0.6

22:6 m3 9.3E0.2 "OE03 MOROS ZO SOS Sc a nr 3.0:0.7 “1:820%2

from mantle to viscera in both Antarctic

and Mediterranean molluscs. Lipids

from mantle of Antarctic animals con-

tained the highest levels of polyunsatu-

rated fatty acids (PUFA) counting for ca.

50%. Also, the fatty acid composition

varied considerably in SL and PL of the

same individual (Table II.

In general, the SL of the mantle of

Antarctic molluscs consisted largely of

saturated (SFA) and monounsaturated

(MUFA) components, with high

SFA/PUFA and MUFA/PUFA ratios

(Table V). Particularly, the SFA/PUFA

ratio in SL of Mediterranean species was

significantly lower than for Antarctic

species. The PL of the mantle of Antarctic

molluscs were featured by a high content

of PUFA, with similar ratios for the

SFA/PUFA and MUFA/PUFA ratios

(Table V). Accordingly, the analysis of the

SL composition of the mantle of Antarctic

molluscs revealed that palmitic acid

(16:0) and stearic acid (18:0) were the

main acyl residues, whereas arachidonic

acid (20:4 w6) and eicosapentaenoic acid

(20:5 3) were the dominant species in PL

(Fig. 1). Compared to Antarctic gas-

tropods, the mantle PL of Mediterranean

molluscs contained higher levels of

MUFA (Table V), with a particularly large

content of octadecaenoic acid (15.98+12.4;

18:1 07/09) (Fig. 1). The MUFA /PUFA

ratio was significantly higher than the

Antarctic value (Table V).

Mantle sections of T. challengeriana

contained levels of palmitoleic acid (16:1

07) considerably higher than those of

palmitic acid and stearic acid, but the

other two Antarctic species revealed an

opposite composition (Table IMM). A

similar trend was found in the viscera

(Table IV). The overall percentage of

PUFA, mainly arachidonic and eicos-

apentaenoic acids, was very similar in

mantle PL from M. mollis, B. clavigera

and T. challengeriana (49.2%, 44.3% and

58.0%, respectively) although the spe-

cific composition varied according to

the species (Table III).

The extracts of the viscera contained

similar amounts of FFA in Antarctic and

Mediterranean animals (58.1+17.2 ug

and 61.1+14.1 ug per mg of lipid extract,

respectively). However, the content of

FFA in the mantle was consistently

higher in the Antarctic organisms

(226.9+24.4 ug and 62.5+18.1 ug per mg

of lipid extract, respectively in Antarctic

and Mediterranean molluscs). The GC-

ÁVILA ET AL.: Fatty acids of Antarctic gastropods

Table V. Fatty acid ratios in mantle SL (storage lipids: triglycerids and wax esters) and PL (phosp-

holipids) in Antarctic and Mediterranean gastropods. SFA: Saturated fatty acids; PUFA: polyunsa-

turated fatty acids; MUFA: monounsaturated fatty acids. *: significantly different than the Antarc-

tic species value (p<0.01; t-test).

Tabla V. Relación de ácidos grasos en SL del manto (lípidos de almacenamiento: triglicéridos y ésteres de

ceras) y PL (fosfolípidos) en gasterópodos Antárticos y Mediterráneos. SEA: Acidos grasos saturados;

PUFA: ácidos grasos poliinsaturados; MUFA: ácidos grasos monoinsaturados. *: significativamente dife-

rente del valor obtenido en especies Antárticas (p<0.01; t-test).

Antarctic Mediterranean

SFA/PUFA 2.45 0.39*

MUFA/PUFA 2.68 1.24

MS analysis of mantle FFA showed a

trend in fatty acid distribution similar to

that found in mantle PL, with SFA and

MUFA predominant in Mediterranean

animals and PUFA more abundant in

Antarctic organisms (Fig. 2). However,

the fatty acid composition of mantle

FFA in Antarctic molluscs proved to be

rather different from that found in

mantle PL and SL of the same species

(Table III). Also the FFA profile in the

mantle of Mediterranean molluscs

showed very few similarities to the fatty

acid distribution in FFA, PL and SL from

the viscera of the same species (Tables

II and IV). Analysis of FFA composition

revealed that arachidonic acid (20:4 w6),

eicosapentaenoic acid (20:5 m3) and

docosahexaenoic acid (22:6 w3) predom-

inated significantly in Antarctic mol-

luscs (p<0.05; t-test) whereas Mediter-

ranean animals were mainly featured by

a higher percentage of octadecadienoic

acid (13.75+0.98; 18:2 w6) which was

almost absent in Antarctic animals

(0.76+0.66) (Fig. 2).

DISCUSSION

In this study we analyzed the lipid

composition of the opisthobranchs B.

clavigera, T. challengeriana, H. picta and

D. limbata, and the prosobranch M.

mollis. Although the data are of limited

value due to the few number of individ-

Antarctic Mediterranean

0.40 0.52

0.47 9%

uals analyzed from Antarctica, we

believe that the results provide useful

information on their fatty acid composi-

tion, distribution and comparison with

Mediterranean species.

The total lipid content was very

similar in the viscera of Antarctic and

Mediterranean molluscs, but it proved

to be higher in mantle sections of

Mediterranean animals respect to the

Antarctic ones. May be this difference

could be related to the abundance in the

Mediterranean species of non-polar

components, such as terpenoids (ÁVILA,

CIMINO, FONTANA, GAVAGNIN, ORTEA

AND TRIVELLONE, 1991; ÁVILA, CIMINO,

CRISPINO AND SPINELLa, 1991) that were

less abundant in the extracts of the

Antarctic species studied here.

T. challengeriana, contrary to the

other Antarctic species, contained levels

of palmitoleic acid (16:1 w7) in mantle

considerably higher than those of

palmitic acid and stearic acid. As a

similar trend was found in the viscera,

we believe this may reflect dietary pref-

erences. Lipid biomarkers have been

recently used to clarify Antarctic tropho-

dynamics in krill (see PHLEGER, NELSON,

MOONEY AND NICHOLS, 2002). Perhaps

further studies in Antarctic opistho-

branchs will also help to understand

their poorly known trophic relation-

ships with other benthic organisms.

In theory, lipids from the viscera

should be more dependent on factors

Iberus, 22 (2), 2004

Mediterranean species

Antarctic species

*p=0.009

*p=0.0006

7 ; n.s.

A A E.

20:4 20:53 22:50 2246

16:0 16:1 18:0 18:1 20: 1

Figure 1. Relative percentage (mean + SD) of phospholipid fatty acids (PL) in the mantle of the

studied gastropod molluscs from the Antarctic and the Mediterranean. Statistical differences were

determined by t-tests. *: p<0.05. n.s.: not significant. Acids: 16:0 palmitic acid, 16:1 palmitoleic

acid, 18:0 stearic acid, 18:1 octadecaenoic acid, 20:1 eicosanoic acid, 20:4 arachidonic acid, 20:5

eicosapentaenoic acid, 22:5 docosapentaenoic acid, 22:6 docosahexaenoic acid.

Figura 1. Porcentaje relativo (media + SD) de ácidos grasos asociados a fosfolípidos (PL) en el manto de los

moluscos gasterópodos estudiados de la Antártida y del Mediterráneo. Las diferencias estadísticas se determi-

naron mediante t-tests. *: p<0,05. n.s.: no significativo. Nombres de los ácidos: 16:0 ácido palmítico, 16:1

ácido palmitoleico, 18:0 ácido esteárico, 18:1 ácido octadecaenoico, 20:1 ácido eicosanoico, 20:4 ácido ara-

quidónico, 20:5 ácido eicosapentaenoico, 22:5 ácido docosapentaenoico, 22:6 ácido docosahexaenoico.

such as diet and reproductive cycles,

whereas fatty acid composition of

mantle should be far more responding

to environmental conditions, such as

temperature or depth. The fatty acid

composition in SL and PL of mantle

extracts were approximately similar in

all animals studied, although the mantle

PL of Antarctic species showed a higher

content of unsaturated fatty acids (Fig.

1). In fact, the overall levels of PUFA in

PL were similar in the three species of

Antarctic molluscs, but were consis-

tently higher than those of molluscs

from the Mediterranean. MUFA /PUFA

ratios in mantle PL were significantly

divergent in Antarctic and Mediter-

ranean molluscs, and suggested a differ-

ent composition of membrane phospho-

lipids from the two groups. In fact, fatty

acids in PL have positional specificity,

being the sn-1 and sn-2 position of glyc-

erol usually occupied by saturated (or

trans-unsaturated) and polyunsaturated

groups, respectively. At a molecular

basis, the fatty acid composition of the

mantle of Antarctic specimens sup-

ported the presence of a large fraction of

PL with polyunsaturated /polyunsatu-

rated or monounsaturated /polyunsatu-

rated chains, whereas the analysis of the

Mediterranean specimens indicated an

opposite trend in PL with saturated or

monounsaturated species at both the 1-

position and the 2-position. In a study

on fish (DEY ET AL., 1993), it was sug-

gested that some phospholipids, such as

those containing oleic/docosahexaenoic

and oleic/eicosapentaenoic acids, play

an important role in the membrane

ÁVILA ET AL.: Fatty acids of Antarctic gastropods

[_— ] Mediterranean species

Antarctic species

*p=0.02

*p<0.0001

E E

*p=0.004

*p=0.002

FOTO AAESIOAASA TS: 2 LOMO ADOS LSD

Figure 2. Relative percentage (mean + SD) of free fatty acids (FFA) in the mantle of the studied gas-

tropod molluscs from the Antarctic and the Mediterranean. Statistical differences were determined

by t-tests.

*. p<0.05. n.s.: not significant. Acids are as in Figure 1, and 18:2 is octadecadienoic acid.

Figura 2. Porcentaje relativo (media + SD) de ácidos grasos libres (EFA) en el manto de los moluscos

gasterópodos estudiados de la Antártida y del Mediterráneo. Las diferencias estadísticas se determinaron

mediante t-tests.

el ácido octadecadienoico.

homeostasis. In particular, an increase of

the unsaturated fatty acid percentage in

the sn-1 position of phospholipids, such

as that due to the replacement of

palmitic acid by oleic acid, may affect

the membrane structure in order to

maintain its functional integrity at cold

temperatures (DEY ET AL., 1993). Our

data, therefore, are in agreement with

this, having found high levels of

polyunsaturated or monounsaturated

fatty acids in position sn-1 of mantle

phospholipids in Antarctic animals.

Our results also showed a very high

content of FFA in the Antarctic samples.

Free fatty acids may be produced due to

degradation of SL and PL during han-

dling and storage. Consequences of slow

frozen storage autolysis are well know in

fish research (HARDY, MCGILL AND GUN-

STONE, 1979) and these may have

affected our FFA results, since our proce-

*: p<0,05. n.s.: no significativo. Nombres de los ácidos como en la Figura 1, y 18:2 es

dure consisted in a very fast storage of

samples at -30%C until extraction, while

only a storage a -80”C completely blocks

enzymatic activity. However, there are

some evidences suggesting a physiologi-

cal meaning for these FFA levels. First,

degradation did not occur at a similar

rate in mantle and viscera of the animals,

since we detected higher quantities of

FFA in the mantle. Moreover, no appar-

ent correlation was observed between

the FFA composition of the mantle and

that of the other lipid pools from both

mantle and viscera. We believe that the

production and occurrence of high levels

of FFA may be a distinct characteristic of

these Antarctic species, reflecting the

chemical-physical properties of the cold-

adapted metabolism of these organisms

(e.g. lipases). Further research should

investigate this possibility with larger

numbers of specimens.

Iberus, 22 (2), 2004

Besides the reported high amounts,

the specific composition of FFA in the

mantle of Antarctic and Mediterranean

molluscs was also remarkably different

(Fig. 2). While Antarctic species were

characterized by higher levels of PUFA,

Mediterranean animals showed a domi-

nance of saturated and monounsaturated

species. It seems probable that the differ-

ent distribution of FFA may indicate an

environmental adaptation. Whether and

how the accumulation of FFA is related in

any way to the membrane homeostasis

remains to be thoroughly investigated. lt

is interesting to note, however, that the

high concentration of FFA can be one of

the ways to transcend the positional

specificity of fatty acids in phospholipids

(MEAD, ALFIN-SLATER, HOWTON AND

PopjAk, 1986). In fact, the formation of

phospholipids involves the transfer of an

acyl group from CoA to either sn-1 or sn-

2 positions of the corresponding

lysophosphoglycerides. Such transacyla-

tion is catalysed by acyltransferases,

which are enzymes sensitive to the chem-

ical features of the fatty acid chains. As

discussed above, the final result of this

preference is the positioning of saturated

fatty acids at the 1-position and of cis-

unsaturated fatty acids at the 2-position.

It has been demonstrated that the posi-

tion-dependent specificity of acyltrans-

ferases can be overridden by the fatty

acid concentration (MEAD ET AL., 1986). In

the Antarctic molluscs, the presence of

high levels of PUFA, therefore, may be

needed for the synthesis of phospho-

lipids with polyunsaturated chains at

both the 1-position and the 2-position of

BIBLIOGRAPHY

ARNTZ, W. AND GUTT, J., 1997. The expedtition

Antarktis XIII/3 (EASIZ I) of RV Polarstern

to the Eastern Weddell Sea in 1996. Berichte

zur Polarforschung/Reports on Polar Research,

ZAS Pp:

AVILA, C., CIMINO, G., FONTANA, A., GAVAGNIN,

M., ORTEA, J. AND TRIVELLONE, E., 1991. De-

fensive strategy of two Hypselodoris nudi-

branchs from Italian and Spanish coasts. Jour-

nal of Chemical Ecology, 17(3): 625-636.

glycerol. Further studies are, however,

necessary to clarify this point.

Our results show that gastropod

molluscs, and opisthobranchs in partic-

ular, possess biochemical mechanisms

of adapting the fatty acid composition

of phospholipids to temperature. Simi-

larly, Viarengo and coworkers

(VIARENGO, ACCOMANDO, ROMA,

BENATTI, DAMONTE AND ORUNESU, 1994)

demonstrated that fatty acid differences

between Antarctic and Mediterranean

scallops were related to cell membrane

adaptation to low temperatures.

Although the number of species investi-

gated in our study is very limited, we

suggest that these data are representa-

tive of the lipid adaptation in gastropod

molluscs from cold and temperate envi-

ronments.

ACKNOWLEDGEMENTS

Thanks are due to K. Iken (Univer-

sity of Alaska Fairbanks) for her help

along. this study and suggestions to

improve the manuscript. W. Arntz , T.

Brey and K. Beyer and the crew of ANT

XI1/3 (1996) are acknowledged for their

helpful support. C. Ávila had a MEC

contract within the Spanish Antarctic

projects ANT95-1011 (1996-97) and

ANT97-0273 (1998-99); for these, thanks

are also due to A. Ramos, S. Agustí and

R. Sardá. Partial funding for C. Ávila

came from ANT97-1590E. Traveling

between Germany, Italy and Spain was

supported by HA1997-0063 and HI1997-

0001 from the Spanish Government.

ÁVILA, C., CIMINO, G., CRISPINO, A. AND SPI

NELLA, A., 1991. Drimane sesquiterpenoids

in Mediterranean Dendrodoris nudibranchs:

Anatomical distribution and biological role.

Experientia, 47: 306-310.

ÁviLA, C., 1995. Natural products of opistho-

branch molluscs: a biological review. Ocea-

nography and Marine Biology, An Annual Re-

view, 33: 487-559.

ÁVILA ET AL.: Fatty acids of Antarctic gastropods

CARBALLEIRA, N. M., MIRANDA, C. AND Ro-

DRÍGUEZ, A. D., 2002. Phospholipid fatty acid

composition of Gorgonía mariae and Gorgonía

ventalina. Comparative Biochemistry and Phy-

siology, 131B: 83-87.

CHAKKODABYLU, R. S. AND THOMPSON, G. A. JR.,

1984. The mechanism of membrane response

to chilling. Journal of Biological Chemistry, 259:

8706-8712.

CuLLiIs, P. R. AND HOPE, M. J., 1991. Physical pro-

perties and functional role of lipids in mem-

brane. In Vance, D. E. and Vance J. (Eds. ):

Biochemistry of lipids, Lipoproteins and mem-

branes: 1-41. Elsevier Science Publishers B.

V. Amsterdam.

DeEy, L, BUDA, C., Wrux, T., HALVER, J. E. AND

FARKAS, T., 1993. Molecular and structural

composition of phospholipid membranes in

livers of marine and freshwater fish in rela-

tion to temperature. Proceedings of the Natio-

nal Academy of Sciences USA, 90: 7498-7502.

FARKAS, T., FODOR, E., KITAJKA, K. AND HAL-

VER, J. E., 2001. Response of fish membranes

to environmental temperature. Aquaculture

Research, 32 (8): 645-655.

FREITES, L., LABARTA, U., AND FERNÁNDEZ-KEI-

RIZ, M. J., 2002. Evolution of fatty acid pro-

files of subtidal and rocky shore mussel seed

(Mytilus galloprovincialis, Lmk. ). Influence

of environmental parameters. Journal of Ex-

perimental Marine Biology and Ecology, 268(2):

185-204.

GILLIS, T. E., AND BALLANTYNE, J. S., 1999. Mi-

tochondrial membrane composition of two

Arctic marine bivalve mollusks, Seripes gro-

enlandicus and Mya truncata. Lipids, 34: 53-57.

HALL, J. M., THOMPSON, R. J. AND PARRISH, C.

C., 2000. Changes in the fluidity and fatty acid

composition of cell membranes from the sea

scallop (Placopecten magellanicus) during short-

term cold acclimation. Journal of Shellfish Re-

search, 19(1): 602.

HAMILTON, S., HAMILTON, R. J]. AND SEWELL, P.

A., 1992. Extraction of lipids and derivative

formation. In Hamilton, R. J. and Hamilton,

S. (Eds. ): Lipid analysis: 13-64. Oxford Uni-

versity Press, Oxford.

HARDY, R., MCGILL, A. S. AND GUNSTONE, F. D.,

1979. Lipid and autoxidative changes in cold

stored cod (Gadus morhua). Journal of the

Science of Food and Agriculture, 30: 999-1006.

HAZEL, J. R., 1984. Effects of temperature on the

structure and metabolism of cell membranes

in fish. American Journal of Physiology, 246:

460-470.

IRELAND, C., CopPP, B. R., FOSTER, M. P., Me-

DONALD, L. A., RADISKY, D. C. AND SWERSEY

J. C., 1993. Biomedical potential of marine na-

tural products. In Attaway, D. H. and Za-

borsky, O. R. (Eds. ): Marine Biotechnology

vol. 1: 1-30. Plenum Press, New York and

London.

IsaY, S. V. AND BUSAROVA, N. G., 1984. Study

of fatty acid composition of marine orga-

nisms —I. Insaturated fatty acids of Japan sea

invertebrates. Comparative Biochemistry and

Physiology, 77B: 803-810.

KATTNER, G., HAGEN, W., GRAEVE, M. AND AL-

BERS, C., 1998. Exceptional lipids and fatty

acids in the pteropod Clione limacina (gas-

tropoda) from both polar oceans. Marine Che-

mistry, 61: 219-228.

LEHMAL, H., 1999. Adaptation to low tempe-

ratures: lipids in ice diatoms. Berichte zur Po-

larforschung/Reports on Polar Research, 317:

1116)

NELSON, M. M., LEIGHTON, D. L., PHLEGER, C.

F. AND NICHOLs, P. D. 2002. Comparison of

growth and lipid composition in the green

abalone, Haliotis fulgens, provided specific

microalgal diets. Comparative Biochemistry

and Physiology, 131B: 695-712.

MEAD, J. F., ALFIN-SLATER, R. B., HOWTON, D.

R. AND POPJAK, G., 1986. Phosphoglyceride

metabolism. In: Lipids- Chemistry, Biochemistry

and nutrition: 405-428. Plenum Press, New

York.

MOON, B. Y., HIGASHI, S., ZOLTAN, G. AND MU-

RATA, N., 1995. Unsaturation of the mem-

brane lipids of chloroplasts stabilize the pho-

tosynthetic machinery against low tempera-

ture photoinhibition in transgenic tobacco

plants. Proceedings of the National Academy of

Sciences. USA, 92: 6219-6223.

MUNIAÍN, C. AND SCHRODL, M., 1999. Taxono-

mic revision of Magallanic and Antarctic Tri-

toniidae (Suborder: Dendronotacea). Abs-

tracts of the 2nd International Workshop of Ma-

lacology, “Systematics, phylogeny and biology

of opisthobranch molluscs” (Mentfi, Sicily,

Italy): 29.

PHLEGER, C. F., NELSON, M. M., MOONEY, B. D.

AND NICHOLS, P. D., 2002. Interannual and

between species comparison of the lipids,

fatty acids and sterols of Antarctic krill from

the US AMLK Elephant Island survey area.

Comparative Biochemistry and Physiology, 131B:

733-747.

PONDER, W. F. AND LINDBERG, D. R., 1997. To-

wards a phylogeny of gastropod molluscs: an

analysis using morphological characters. Zo-

ological Journal of the Linnean Society, 119: 83-

265.

Quoc, K. P. AND DUBACO, J. P., 1997. Effect of

growth temperature on the biosynthesis of

eukariotic lipid molecular species by the cya-

nobacterium Spirulina platensis. Biochimica

and Biophysica. Acta, 1346: 237-246.

ROTERT, K. R., TOSTE, A. P. AND STEIERT, J. K.,

1993. Membrane fatty acids analysis of An-

tarctic bacteria. Federation of European Micro-

biological Societies, Microbiology Letters, 14:

253-257.

Á3

Iberus, 22 (2), 2004

RUSSELL, N. J., 1998. Molecular adaptations in

psychrophilic bacteria: potential for bio-

technological applications. Advances in Bio-

chemical Engineering and Biotechnology, 61: 1-

Zi:

RUSSELL, N. J]. AND NICHOLs, D. S., 1999. Pol-

yunsaturated fatty acids in marine bacteria

- a dogma rewritten. Microbiology, 145: 767-

769.

SAKAMOTO, T., HIGASHI, S., MURATA, N. AND BR-

YANT, D. A., 1997. Low-temperature induced

desaturation of fatty acids and expression

of desaturase genes in the cyanobacterium Sy-

nechoccusi sp. PCC 7002. Federation of European

Microbiological Societies, Microbiology Letters,

152: 313-320.

SATO, N., MURATA, N., MIURA, Y. AND UETA, NÑN,,

- 1979. Effect of growth temperature on the

biosynthesis of eukariotic lipid fatty acid

composition in the blue-green algae Anaba-

ena variabilis and Anacystis nidulans. Biochimica

and Biophysica Acta, 572: 19-28.

SHU, I. Z., 1998. Recent natural products based

drug development: a pharmaceutical in-

dustry perspective. Journal of Natural Pro-

ducts, 61: 1053-1071.

TANAKA, T. AND TOYAMA, Y., 1959. Fatty oils of

aquatic invertebrates. XXI. Fatty oil of the

sea hare and its sterol components. XXI. Un-

saponifiable components other than sterol

in the fatty oil of the sea hare. Memories of the

Faculty of Engineering Nagoya Univ., 11: 182-

195:

THOMPSON, P. A., GUO, M. X., HARRISON, P. J.

AND WHYTE, J. N. C., 1992. Effects of varia-

tion in temperature. I. On the biochemical

composition of eight species of marine phy-

toplankton. Journal of Phycology, 28: 481-488.

TIBALDI, E., 1966. Ricerche preliminari sugli

acidi grassi di alcune specie di molluschi ma-

rini. Atti Accademia Nazionale dei Lincei, Ren-

diconti, (8) 40: 921-925.

UrIcH, K., 1994. The structure and metabolism

of lipids (Chapter 15). In: Comparative animal

biochemistry, 595-599. Springer-Verlag, Berlin.

VIARENGO, A., ACCOMANDO, R., ROMA, G., BE-

NATTI, U., DAMONTE, G. AND ORUNESU, M.,

1994. Differences in lipid composition of cell

membranes from Antarctic and Mediterra-

nean scallops. Comparative Biochemistry and

Physiology, 109B (4): 579-584.

Voocr, P. A., 1983. Lipids: their distribution and

metabolism. In Hochachka, P. W. (Ed.): The

Mollusca 1 (Metabolic biochemistry and mo-

lecular biomechanics): 329-370. Academic

Press, New York.

O Sociedad Española de Malacología Iberus, 22 (2): 45-49, 2004

Why are beached Acanthochitona fascicularis (Linnaeus,

1767) (Mollusca: Polyplacophora) from Italy missing their

dorsal girdle elements?

¿Por qué los ejemplares de Acanthochitona fascicularis (Linnaeus,

1767) (Mollusca: Polyplacophora) encontrados en una playa de Italia

pierden los elementos del perinoto?

Bruno DELEUANGELO*, Bruno ANSEEUW**, Yves TERRYN*** and

Antonio BONFITTO****

Recibido el 25-111-2004. Aceptado el 21-VII-2004

ABSTRACT

An unusual form with reduced dorsal girdle of the well known and extremely variable

European chiton Acanthochitona fascicularis (Linnaeus, 1767) is hereby illustrated, dis-

cussed, and compared with the regular form. Hundreds of beached specimens and several

live found specimens with similar dorsal girdle were studied and compared. Different theo-

ries and hypothesis are proposed as an explanation for this unusual form and as a conclu-

sion we finish with some open questions.

RESUMEN

En el presente trabajo se estudia una forma inusual con reducción de elementos del peri-

noto del poliplacóforo Acanthochitona fascicularis (Linnaeus, 1767). Se compara, se ilus-

tra y se discuten los datos obtenidos con la forma regular de esta especie. Los autores

aportan su hipótesis sobre la presencia de esta inusual forma y concluyen que hay algu-

nas cuestiones abiertas.

KEY WORDS: Polyplacophora, Acanthochitona fascicularis, Mediterranean Sea, girdle.

PALABRAS CLAVE: Polyplacophora, Acanthochitona fascicularis, Mediterráneo, perinoto.

INTRODUCTION

Many years ago, a great number of

specimens of Acanthochitona fascicularis

(Linnaeus, 1767) were beached after a

storm at “Calambrone”, a locality near

Livorno (Tuscany, Italy). A friend of the

first author picked up many of these

and prepared part of them in an

alcohol /glycerine solution, the rest were

preserved in alcohol. All collected speci-

mens had an unusual feature in

common: a “smooth” dorsal girdle with

reduced sutural tufts. Later on, similar

* Vía Mugellese 66D, 59100 Prato, Italy; bruno.dellangeloCelsag,it

** Mispelstraat 18, 9820 Merelbeke, Belgium; chitonGfpandora.be

*** Kortrijksepoortstraat 109, 9000 Gent, Belgium; loricatafpandora.be

** Museo di Zoologia dell'Universita* di Bologna, Via Selmi 3, 40126 Bologna, Italy; bonfitroalma.unibo.it

Iberus, 22 (2), 2004

living specimens were found at two

localities in Italy, which made a compar-

ative study possible.

Abbreviations:

BDA Private Collection of Bruno Del-

l'Angelo, Italy.

BA Private Collection Bruno Anseeuw,

Belgium.

YT Private Collection Yves Terryn,

Belgium.

MATERIAL AND METHODS

Material examined: Acanthochitona

fascicularis (Linnaeus, 1767)

e Calambrone, Livorno prov., Italy,

56 specimens beached in Feb. 1991

(BDA, Fig.7), including 21 preserved in

alcohol (Fig.6). The dimensions vary

from 28 x 15 mm to 40.5 x 18.5 mm. Only

one spm is of smaller dimensions, 13 x

6.8 mm.

e Calambrone, Livorno prov., Italy, 5

specimens beached on Feb. 1991 (BDA),

about 28.5 x 14 mm, with plates covered

with Bryozoa and other organisms (Fig.5).

e Punta Faro, Messina Strait, Italy,

under rocks at 2-3 m: 2 specimens, about

25 x 14 mm (estimated) strongly curled

and 18 x 12 mm curled (BDA).

e Off Capraia Is]., Italy, dredged at

about -200 m, inside a large semi porous

rock (about 70 x 70 cm): 5 specimens live

taken (S. Savona collection, Italy).

e Hundreds of specimens from

diverse European (Portugal, Spain,

France, Italy, Croatia, Greece) and

African (Morocco, Algeria, Tunisia,

Israel) localities (BDA, BA and YT col-

lections).

SEM-images were made using a SEM

JEOL mod. JSM-5200. Specimens were

disarticulated by boiling in a KOH-solu-

tion until all soft parts were dissolved,

followed by thorough rinsing.

RESULTS

The mollusc-beaching phenomenon

at Calambrone is well known (CAMPANI,

1983), and the number of molluscs

species beached over a period of several

years is rather high (257 species). The

beaching is caused by interaction

between different sea currents and by

the characteristic sea floor. Among these

beached molluscs, three species of

chitons have been reported: Ischnochiton

rissor (Payraudeau, 1826), Chiton oli-

vaceus Spengler, 1797 and Acanthochitona

fascicularis (Linnaeus, 1767) (reported in

Campani's list as Acanthochitona aenea).

The beaching of chitons is generally

uncommon, and usually concerning few

specimens. The beaching in February

1991 is therefore particularly significant

especially as the features of the A. fascic-

ularis beached are so different from

typical shallow-water A. fascicularis.

Some of these specimens have been

illustrated in the book on chitons from

the Mediterranean Sea (DELL'ANGELO

AND SMRIGLIO, 2001).

When we first saw these beached

“smooth” specimens, we thought they

were regular A. fascicularis with

damaged dorsal girdle. In fact, it

seemed as if the dorsal girdle elements

had been removed by some cause as all

spicules were gone and the sutural tufts,

which normally bear long spicules, were

very short , almost truncate. Beside this,

all other features looked macroscopi-

cally identical to normal A. fascicularis,

i¡.e. general shape and tegmentum sculp-

ture. As these specimens were dead,

some external influence could have

caused this phenomenon. For example

the specimens could have died during

the storm resulting in curled up speci-

mens which could have been dorsally

damaged by scraping and rolling upon

rocks during the beaching process. This

could explain why only the dorsal

girdle was removed and not the ventral

part which seemed to be intact.

However, the tegmentum showed no

obvious signs of damage. It was as if

something had meticulously removed

the dorsal girdle, without touching the

rest of the animal. We deliberately

removed the dorsal girdle elements (by

scraping with a knife) from a similar

“regular”, alcohol /glycerine preserved

specimen and the result was a specimen

DELLANGELO ET AL.: Acanthochitona fascicularis missing dorsal girdle elements

Figures 1-4. Acanthochitona fascicularis beached, with “smooth” dorsal girdle. 1: girdle, tuft; 2:

girdle, tuft, lateral view; 3: isolated tuft (right: top, left: inside the girdle); 4: girdle, ventral spicu-

les. Figures 5-7. Acanthochitona fascicularis beached. 5: one of the five specimens with the plates

covered with bryozoans and other organisms (28.2 x 14.3 mm); 6: one of the specimens preserved

in alcohol (28.5 x 17.6 mm); 7: one of the specimens prepared in an alcohol/glycerine solution

(38.7 x 17.5 mm).

Figuras 1-4. Acanthochitona fascicularis, con el perinoto dorsal “liso”. 1: perinoto, protuberancia; 2:

perinoto, protuberancia, vista lateral; 3: protuberancia aislada (derecha: parte superior, izquierda:

dentro del perinoto); 4: perinoto, espículas ventrales. Figuras 5-7. Acanthochitona fascicularis. 5: uno

de los cinco especímenes con las paclas cubiertas por briozoos y otros organismos (28,2 x 14,3 mm); 6:

especimen conservado en alcohol (28,5 x 17,6 mm); 7: especimen preparado en solución de alcohol/glice-

rina (38,7 x 17,5 mm).

Iberus, 22 (2), 2004

that looked almost identical to the

beached ones. The external cause

remains therefore a possible explanation

of the phenomenon.

The disarticulation of one of the spec-

imens was necessary for a complete analy-

sis and comparison with a typical speci-

men of A. fascicularis. When disarticulated

no dorsal girdle appeared. Normally,

when submitting a chiton to this process,

the soft parts of the animal will dissolve

and it will result in the 8 isolated valves,

the radula and a thin layer of dorsal and

ventral girdle bearing the girdle elements.

In the beached specimen, only the ventral

layer was present. The sutural tufts, which

are normally attached to the dorsal layer,

were loose. We concluded that the speci-

men had no dorsal girdle, but what caused

this lack of girdle?

Other material beached at Calam-

brone during February 1991 provided

new insight into this problem. Among

the molluscs beached were five speci-

mens of A. fascicularis with valves

severely covered with corals and/or

eroded, but with the dorsal girdle in

normal condition (Fig. 5). The only dif-

ference from regular A. fascicularis is the

reduced tufts, but this is known to occur

within members of this species. Also,

several of the alcohol preserved speci-

mens still have some of their dorsal

girdle elements present (i.e. areas with

fine and short spicules) and reduced

tufts, which supported the notion that

the lack of dorsal girdle in the previous

material was, in fact, caused by external

influences.

DISCUSSION

We have considered several hypothe-

ses that might explain the phenomenon

of beached Acanthochitona “fascicularis”

with reduced or detached dorsal girdles.

First, it is possible that the specimens were

subjected to an acidic environment, either

before or after their death. The problem is

that this acidic environment would also

“etch” the tegmentum and there is no evi-

dence of this visible on the specimens. We

consider this hypothesis unlikely. Another

possible cause is that the specimens lived

below the carbon compensation depth,

where seawater dissolves calcareous mate-

rial quite aggressively, but this is only

known to appear at great depths (>4000

m) so this can be rejected, as the specimens

were found on the shore and it is highly

improbable that they were transported

from great depths by the storm.

Could this phenomenon have been

caused by polluted water? Is it perhaps

caused by something similar as the “red

tides” which are known to kill chitons in

great numbers in South Africa for

example? (fide A. Seccombe, pers.com.

1997; B. Anseeuw and Y. Terryn, pers.obs.

2002: beaching of Dinoplax gigas Gmelin,

1791). Has an animal removed the dorsal

girdle?

One possible hypothesis is that the

specimens could be a distinctive ecolog-

ical form (ecotype) of fascicularis, i.e.

with reduced dorsal girdle, or even an

unknown species. We are unaware of

any previous mention of this distinction

in literature regarding chitons. In fact, it

is highly improbable that a species

could live without their protecting

dorsal girdle, so this hypothesis was

also considered as highly improbable. It

is however known that the dorsal girdle

elements can be quite variable, even

within specimens from the same local-

ity. For example specimens from a local-

ity which are found on exposed habitats *

can have reduced dorsal girdle ele-

ments, but will also generally show

eroded tegmentum, a feature that was

not observed in the beached specimens.

The most logical remaining hypothe-

sis for us was that some unknown

posthumous process led to the removal

of the dorsal girdle. However, we were

then at a loss to explain our later observa-

tion of living specimens with similar

“naked” appearance. This fact made

everything much more complicated as

these specimens were alive but without a

dorsal girdle. Thus, we still do not have

an explanation about why all these speci-

mens have a reduced dorsal girdle. It is

also possible that these specimens could

represent a species or race that is distinct

from typical A. fascicularis but the only

DELLANGELO ET AL.: Acanthochitona fascicularis missing dorsal girdle elements

difference between both that we have

observed so far is the absence of the

dorsal girdle elements. On the other

hand, it is highly improbable that a

lineage of chitons would lose their dorsal

girdle, as it would seem to make it quite

unprotected and vulnerable. Perhaps a

DNA analysis could test the “different

species” hypothesis but this can only be

done easily on fresh alcohol-preserved

specimens. Perhaps the future will bring

some fresh alcohol-preserved specimens

but until then we can only speculate. We

would be happy to hear of anyone who

has observed similar specimens or of

BIBLIOGRAPHY

CAMPANI, E., 1983. Molluschi spiaggiati in

localita” Calambrone (LD). Interpretazione dati

anni 1970-1982. Quaderni del Museo di Storia

Naturale di Livorno, 4: 59-74.

similar observations for other chiton

species, as more information could even-

tually lead to a plausible explanation.

ACKNOWLEDGEMENTS

We wish to thank Giacomo Di Pacot

(Livorno, Italy), Sergio Savona (Livorno,

Italy) and Salvatore Ventimiglia (Punta

Faro, Messina, Italy) for putting at our

disposal the material they collected, and

Douglas Eernisse (California State Uni-

versity) for comments and suggestions

in preparation of this paper.

DELLANGELO, B. AND SMRIGLIO, C., 2001.

Living Chitons from the Mediterranean Sea.

Evolver, Roma, 255 pp.

CNI )

yr nd Ss rindo

Ea dy TN

0 1)

e ano possable aha

CUNATA

el )

ml aecrltiei

rt e

mir E li

mpriva Heipak. Y

O a '

> E ey A

» QA

15 4,

2147 Í

il carl Ma ;

DonvHs

¡0 *ke ved

Joy se still

A hd tigoabonk bid l

LEVES £

y K

“E

vé a

5 tr Ló dro

tuenT

O Sociedad Española de Malacología Iberus, 22 (2): 51-79, 2004

The genus Calyptraea (Gastropoda, Caenogastropoda,

Calyptraeidae) in the East Atlantic

El género Calyptraea (Gastropoda, Caenogastropoda, Calyptraei-

dae) en el Atlántico oriental

Emilio ROLÁN*

Recibido el 14-11-2004. Aceptado el 2-[X-2004

ABSTRACT

The genus Calyptraea is revised for the East Atlantic. Up to now, only C. chinensis was

recognized but this species is only present in the Mediterranean Sea, and in the European

and North African Atlantic. Two new African species confused up to now with C. chinen-

sis are described, one with a range from Western Sahara to Guinea Conakry and Benin,

and the other from Mauritania to Angola. Another South African species, also considered

C. chinensis, is in fact a different taxon.

RESUMEN

Se revisa el género Calyptraea en el Atlántico oriental. Calyptraea chinensis, la Única

especie reconocida actualmente, se distribuye sólo en el Mediterráneo, y en el Atlántico

europeo y norteafricano. Dos nuevas especies, hasta ahora consideradas como C. chinensis,

se encuentran en Africa; una de ellas se extiende desde el Sáhara Occidental hasta Guinea

Conakry y Benin, y otra desde Mauritania hasta Angola. Otra especie de Sudáfrica, tam-

bién considerada hasta el momento como C. chinensis, es una especie diferente.

KEY WORDS: Calyptraea chinensis, Europe, West Africa, new species.

PALABRAS CLAVE: Calyptraea chinensis, Europa, África occidental, nuevas especies.

INTRODUCTION

The genus Calyptraea Lamarck, 1799

is represented in the East Atlantic by

only one species, in the opinion of most

recent authors (NICKLES, 1950; BERNARD,

1984; FRETTER AND GRAHAM, 1981;

SABELLI, (GIANNUZZI-SAVELL AND

BEDULLI, 1991; POPPE AND GOTO, 1991;

ROLÁN AND RYALL, 1999 and CHIARELLI,

2002). Against this recent situation,

more species of this genus are known

from the fossil records (CASTAÑO, CIVIS

AND GONZÁLEZ DELGADO, 1988; VERA-

PELÁEZ, LOZANO-FRANCISCO, MUNÑIZ-

SOLÍS, GILI, MARTINELL, DOMENECH,

PALMQVIST AND GUERRA-MERCHÁN,

1995). The systematics and the phy-

logeny of the genus Calyptraea were

studied by FISCHER (1950).

Being a common species with a wide

distribution, it was to be expected that

some of the names employed in the past

to designate C. chinensis are actually syn-

onyms (most from the Mediterranean)

(see SABELLI ET AL., 1991 and below).

* Museo de Historia Natural, Universidad de Santiago, 15782 Santiago de Compostela. e-mail:

emiliorolanPinicia.es

Iberus, 22 (2), 2004

- A general description of this species

is given in FRETTER AND GRAHAM (1962,

1981) together with bibliographic infor-

mation on different aspects. Other refer-

ences to this species will be mentioned

below under distribution.

After studying numerous samples of

material from many European and West

African localities, the comparison of shells,

anatomy and other characters indicated

the existence of more than one species.

This is the subject of the present work.

MATERIAL AND METHODS

Some of the material examined as

well as numerous specimens and shells

were obtained by the author from expe-

ditions to several West African localities:

Morocco, Mauritania, Canary Islands,

Senegal, Ghana, Sáo Tomé ls., Principe

Is., Annobón Is. and Angola. Many

samples from the MNHN collection and

also from some private collections men-

tioned below were also studied. In vivo

observations were made by the author.

RESULTS

Abbreviations

AMNH American Museum of Natural

History, New York

BMNH The Natural History Museum,

London

IPM Instituto Portugués de Malacologia,

Lisbon

LS Linnean Society, London

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Muséum Nationale d'Histoire

Naturelle, Paris

ZSM Zoologisch Staatssammlung,

Miúnchen

CJH collection of José María Hernández,

Gran Canaria

CER collection of Emilio Rolán, Vigo

CFS collection of Frank Swinnen,

Lommel, Belgium

sp specimen with soft parts

s shell

f fragment

j juvenile. As the smaller specimens are

adult males (Bacci, 1951), only in

very small shells was the category of

juvenile employed.

Family CALYPTRAEIDAE Lamarck, 1809

Genus Calyptraea Lamarck, 1799

Calyptraea chinensis (Linnaeus, 1758) (Figs. 1-27, 65-67, 83-85, 91, 94, 97, 100,

103-107, 113-116)

Patella chinensis Linnaeus, 1758. Syst. Nat., ed. X, p. 781, no. 656; ed. XIL, p. 1257, no. 749. [Type

locality: M. Mediterraneo].

Patella sinensis Gmelin, 1791. Syst. Nat., ed. XII, p. 3692.

Patella albida Donovan, 1802. Nat. hist. Brit. Shells, YV, pl. 129.

Patella squamulata Renier, 1804. Prodr. Observ. [not available: published in a paper rejected by

ICZN opinion 316/1954].

Patella muricata Brocchi, 1814. Conch. foss. subap., p. 16.

Calyptraea laevigata Lamarck, 1822. Hist. nat. anim. sans vert. VL IL p. 21.

Calyptraea succinea Risso, 1826. Hist. Nat. Eur. Mer., p. 255.

Calyptraea vulgaris Philippi, 1836. Enum. moll. Siciliae, L, p. 119.

Calyptraea polii Scacchi, 1836. Catal. Conch. Regn. Neapolitani, p. 17.

Calyptraea spirata Nardo, 1847. Sinon. modern...Golfo Veneto. In SABELLI ET AL. (1990), p. 160.

Calyptraea canarica Buonamni fide JEFFREYS (1865). Brit. Conch., 3, p. 275.

Calyptraea mamma Krynicki fide Middendorff in JEFFREYS (1865). Brit. Conch. 3, p. 276.

Calyptraea políi var. fusca Issel, 1878. Croc. del Viol.-Testacei, p. 31.

Calyptraea sinensis var. coralligena Pallary, 1900. Coquil. marins d'Oran, p. 329.

Calyptraea sinensis var. elliptica Pallary, 1900. Coquil. marins d'Oran, p. 329.

352

ROLÁN: The genus Calyptraea in the East Atlantic

Type material: 2 syntypes, in LS (Figs. 113-116); the larger one is here designated lectotype (Figs.

113, 114)

Other material examined: Great Britain: 36 s, Shells Bay, Studtland, Dorset (MNHN). Atlantic

France: 15 s, Saint Michel, Normandy, in shells on sand, intertidal (CER); 15 sp, 16 s, 103, Les Cochons

Noires, Roscoff, Finistere, 20 m (MNHN); 4 sp, Roscoff, on shells of sand bottom, intertidal (CER);

3 c, NE canyon de la Cassidaigne 48” 06.7” N 05* 55” E, 150-250 m (MNHN); 35 sp, Saint Quay Por-

trieux, Baie de Saint Brieuc, 10-30 m, on shells (CER); 4 s, Plougastel, Tinduff, port (MNHN); 33 s,

Anse de Bertheaume, 20-30 m (MNHN). Atlantic Spain: 40 sp, Ribadeo, intertidal, on shells (CER);

12 sp, 23 s, Vigo, Samil, 12 m, on shells and rocks (CER); 155 sp, 30 s, O Grove, on shells, sandy

bottom, intertidal (CER); 53 sp, 30 s, Vigo, Cies Islands, 9 m, on shells and rocks (CER); 20 sp, Punta

do Arroás, Vigo, intertidal (CER); 3 s, Vigo, San Simón, 5 m, on shells, sandy bottom (CER); 55 sp,

Vigo, Toralla, 20 m, on shells, sandy bottom (CER); 3 j, Conil de la Frontera (MNHN); 6 s, Cádiz

Bay, 18 m, from sandy sediment (CER); 1 c, El Puerto de Santa María, Cádiz, beach sediment

(MNHN); 6 s, Sotogrande, Torre Guadiaro, Cádiz, circalittoral (MNHN),; 15 c, Torre de la Peña,

Tarifa, in beach sediment (MNHN). Portugal: 30 sp, 9 s, Tavira, Terra Estreita 37” 06' N 07” 38.4”

W, 3 m (MNHN); 2 sp, Ria Formosa, Ilha do Farol, Casi dos Hangares, 3 m (IPM); 6 sp, 7 s, Algarve,

between Sagres and Faro, 40-50 m (MNHN); 106 sp, 70 c, Chenal do Olháo, 37* 00' N 07* 51 W, 3-

7 m (MNHN); 6 sp, 12 s, Baia de Baleeira, 37” 00.7" N 8? 55” W, 12-17 m (MNHN); 20 sp, 8 s, Baia

do Peixe, 37” 00' N 08? 58” W, 23 m (MNHN); 1 sp, Algarve 36* 56' 871”N 08* 17' 45”W, 69-105 m

(IPM). Mediterranean Spain: 114 sp, 165 s, La Atunara, La Línea de la Concepción, 27 m, on shells,

in sandy bottom (CER); 2 s, La Atunara Beach, La Línea, 36” 10.2” N 05* 19.3” W, 30 m (MNHN); 1

s, La Atunara Beach, La Línea, 36” 05.19 N 05* 19 W, 40-45 m (MNHN); 15 s, Málaga, 20-40 m

(MNHN); 5 sp, 23 s, 30 j, Rincón de la Victoria, Málaga (MNHN); 4 s, Marbella port (MNHN); 3 s,

Marbella (CJH); 1 c, Tarajal, Ceuta (MNHN). Mediterranean France: 5 c, Lion Gulf, N/O “Europe”

IFREMER/DEPRO 96, 42” 20” N 03? 27.1” W, 650-725 m (MNHN); 2 sp, Lacaze-Duthiers, Banyuls-

ECOMARGE, 42” 30' N 03? 25.20E, 130-190 m (MNHN); 2 sp, Cap Béar, Banyuls-ECOMARGE, 42”

29.40" N 037 10.40E, 66-67 m (MNHN); 1 sp, Cap Béar, Banyuls-ECOMARGE, 42” 30” N 03? 10.50E,

62 m (MNHN); 7 sp, 1 sp, Cap Béar, Banyuls-ECOMARGE, 42” 31" N 03* 10.10E, 70 m (MNHN);

39 s, Marseille N/O “Georges Petit”, Grand Congloué, 83 m (MNHN); 7 s, Le Brusc/Cap Sicié,

Provenza, 40-100 m (MNHN). Morocco: 13 c, Playa Grande, Tanger (MNHN); 1s, Essaouira, 31?

31” N 09? 47” W, beach sediment 2 sp, Cap Béar, Banyuls-ECOMARGE, 42” 29.40” N 03? 10.40E, 66-

67 m (MNHN); 1 c, 16j, N/O Vamneau, 30% 40” N 04? 55” W, 20 m (MNHN); 3 sp, 4c,7j, N/O

Vanneau, 30” 30” N 09? 43" W, 40 m (MNHN); about 300 s and j, N/O Vanneau, 34? 54' N 09* 58”

W, 110 m (MNHN); 4 s, N' Dig (Rincon), circalittoral (MNHN); 3 s, 10 km N Agadir (CFS). Algeria:

3 s, (MUNHN); 1 sp, 1 s, 1 j, Oran (MNAHN). Italy: 4 c, Tuscan archipelago (MNHN); 12 s, Naples

(MNAHN); 2 s, Palermo and Messina (MNHN). Tunis: 3 s, Canal d' Ajim, Djerba, 10-32 m (MNHN);

56 c, Gulf of Gabes, 10-15 m (MNHN). Rumania: 4 sp (MUNHN). Madeira: 19 s, Pontinha-Cais do

Lazareto, 60 m (CES); 14 s, Lido, Funchal (CES); 26 c, Funchal Bay (CFS); 15 s, Funchal Bay, 25-30

m (CES); 22 s, Funchal Bay, 50 m (CES); 26 s, 1 f, Funchal Bay, 70-80 m (CES); 12 s, Funchal Bay, 100

m (CES); 3 s, Funchal Bay, 130 m (CES); 3 s, Funchal Bay, 150 m (CES); 4 s, Funchal Bay, 180 m (CES).

Canary Islands: 2 s, Gando Bay, Gran Canaria, 15 m (CES); 2 s, Maspalomas, Gran Canaria (CES).

Western Sahara: 6 s, S. Peña Grande, 24” 45 N 15* 25 W (CJH); 1 s, Villacisneros, 30 m (CJH).

Description: The original description

of Linnaeus is: “Testa integra subconica

laevi labio laterali”. For a more complete

description, see FRETTER AND GRAHAM

(1981). The shell (Figs. 1-10) has been il-

lustrated in many papers, but the colour

variability was only well represented in

GIANNUZZI-SAVELLI ET AL. (1997) and

MACEDA ET AL. (1999). The protoconch

(Figs. 11-26) is described in FRETTER AND

GRAHAM (1981) as “having 0.75 to 1

smooth whorls but this is often

eroded.... The protoconch is not clearly

separated from the teleoconch. Its diam-

eter is 600-700 um”. This description is

correct but the nucleus of the proto-

conch, according to the definition of

VERDUIN (1976), has between 194 and

230 um. From this nucleus, the proto-

conch increases in size quickly to reach

3-3.5 times the diameter of the nucleus,

and 0.75 to 1 whorl at its end, although

the limit with the teleoconch is not

clearly defined. The total diameter of

the protoconch in the shells studied was

between 630 and 741 um. Seen from the

Iberus, 22 (2), 2004

side'the protoconch is situated on the

teleoconch without a deep suture be-

tween them. Its surface is apparently

smooth but, at high magnification (Figs.

19, 24, 26), numerous, short axial striae

disposed in spiral bands may be ob-

served. This description is valid for 98%

of the more than three hundred proto-

conchs studied: for some variations, see

below, in Remarks. The teleoconch is

rather smooth, of a patelliform shape,

rounded, rather solid, opaque and not

glossy. Ratio between width and height

is usually between 2.5 and 3.1. Only ex-

ceptional populations may have differ-

ences in shape, the shells being higher

or flatter (see below). Many shells are

smooth, not shiny, with or without

prominences, sometimes with oblique

dorsal threads near the border. Within

the same populations, a variable num-

ber of shells show scarcely separated

prominent spines (Figs. 10, 100).

The colour of the shell is most fre-

quently white but it may be yellowish,

cream, pink, brown or violet. The colour

of each population is variable, with

white or brown sometimes predominat-

ing.

Dimensions: FRETTER AND GRAHAM

(1981) indicated up to 15 mm, which is

similar to the maximum dimension

mentioned in most papers; LOCARD

(1892) mentions up to 20 mm. After

examining many specimens, it is

uncommon for the species to reach more

that 20 mm, but there is a shell (CER)

from Pobra do Caramiñal, north-west

Spain, with a diameter of 24.7 mm; in

the MNHN there is another from

Málaga of 25 mm and, finally, the record

is in CJH where there is a shell from

Malaga of 30 mm in diameter.

The animal was described in

FRETTER AND GRAHAM (1962, 1981). The

Vigo and O Grove specimens which

were studied live were whitish, with

milk-white, large, irregular spots on the

border of the mantle forming groups

(Fig. 94). La Atunara (southern Spain)

specimens studied preserved in alcohol

had orange spots on the mantle border,

with opaque translucent white and

cream dots all around.

Due to the importance of the penis

(Fig. 97) in comparison with other

species, it should be described in more

detail. It is placed behind the right ten-

tacle and is elongate with a bilobed

knob at its apex. These two lobes are dif-

ferent, the larger one being like a spoon,

and the smaller one like a finger, with

an arrangement similar to a hand with

the four fingers fused and forming a

concave prolongation, and the thumb in

opposition.

The radula (shown in BANDEL, 1984

and FINET, WUEST AND MAREDA, 1992)

(Figs. 83-85, 91) is taenioglossate; rachid-

ian tooth narrow and with a prominent

central cusp with 4, sometimes 5, smaller

cusps at each side. Lateral teeth with a

prominent and wide cusp at the conflu-

ence of the internal and the external

edges; this cusp may have serrate bor-

ders. Internal edge with few wide cusps

(usually 4) while the external edge has

between 7 and 12. Marginal teeth elon-

gate, curved, sharp pointed and the in-

ternal tooth with two borders with few

cusps (4-6), while the external one has

few cusps (2-4) on its internal part.

The spawn of C. chinensis (Figs. 103-

105) was studied by LEBOUR (1937),

WYarT (1957) and recently described by

TRONCOSO, URGORRI AND CRISTOBO

(1988) in populations from north-west

Spain. It is formed by numerous spheri-

cal-ovoid capsules, with a circular

section (Figs. 103-104), which are fixed

in groups to the substrate by a narrow

peduncle (Fig. 105). The development of

this species is direct; "TRONCOSO ET AL.

(1988) have shown the intracapsular

veliger stage, and here we present two

more advanced stages (Figs. 106, 107).

The numerical data in TRONCOSO ET

AL. (1988) are the following: capsules

1.2-3.0 mm, number of eggs per capsule

2-18 and each egg 0.6-1.0 mm (the

reason for this last size is that it refers to

two larvae in a capsule). Specimens of

15-20 mm from Vigo and O Grove had

8-15 capsules per spawn, of 10-16 eggs

each, 380-490 um in diameter. From La

Atunara, spawn from specimens of 12-

17 mm had 8-18 capsules each, with 6-17

eggs each, of 340-430 um in diameter.

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 1-9: Calyptraea chinensis: 1-3: Shell, 13.2 mm, Vigo (CER); 4-5: Shell, 13.5 mm, La

Atunara, La Línea de la Concepción, Spain (CER); 6: Shell, 8.8 mm, Vigo, Spain; 7-9: Shell, 15.8

mm, St. Brieuc Bay, Bretagne, France (CER).

Figuras 1-9: Calyptraea chinensis: 1-3: Concha, 13,2 mm, Vigo (CER); 4-5: Concha, 13,5 mm, La

Atunara, La Línea de la Concepción, España (CER); 6: Concha, 8,8 mm, Vigo, España; 7-9: Concha,

15,8 mm, St. Brieuc Bay, Bretaña, Francia (CER).

Iberus, 22 (2), 2004

Figure 10. Juvenile shell of Calyptraea chinensis, 11 mm, Vigo. Figures 11-19. Protoconchs of C.

chinensis: 11, 12: Vigo; 13: Bretagne, France; 14: Samil, Vigo; 15, 16: La Atunara, Línea de la

Concepción; 17: O Grove; 18: Marbella, Spain; 19: La Atunara.

Figura 10. Concha de un juvenil de Calyptraea chinensis, 11 mm, Vigo. Figuras 11-19. Protoconchas

de C. chinensis: 11, 12: Vigo; 13: Bretaña, Francia; 14: Samil, Vigo; 15, 16: La Atunara, Línea de la

Concepción; 17: O Grove; 18: Marbella, España; 19: La Atunara.

SÓ

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 20-27. Protoconchs of C. chinensis. Fig. 20: Small protoconch, Madeira, 150 m (CFS); 21:

Large protoconch, Funchal, Madeira, 80 m (CES); 22: Large protoconch, Western Sahara, (CJH);

23: Large protoconch, La Atunara, Spain (CER). 24: Detail of the sculpture of a large protoconch,

La Atunara (CER); 25: Small protoconch, Marseille, Tombant, France, 83 m (MNHN); 26: Microsculp-

ture of a small protoconch, Tombant; 27: Large protoconch, Marseille, Tombant, 83 m (MNHN).

Figuras 20-27. Protoconchas de C. chinensis. Fig. 20: Protoconcha pequeña, Madeira, 150 m (CFS);

21: Protoconcha grande, Funchal, Madeira, 80 m (CFS); 22: Protoconcha grande, Sáhara Occidental,

(CJHA); 23: Protoconcha grande, La Atunara, España (CER). 24: detalle de la escultura de una proto-

concha grande, La Atunara (CER); 25: Protoconcha pequeña, Marsella, Tombant, Francia, 83 m

(MNAN); 26: Microescultura de una protoconcha pequeña, Tombant, Francia; 27: Protoconcha

grande, Marsella, Tombant, 83 m (MNHN).

Iberus, 22 (2), 2004

Figures 28-38. C. africana spec. nov. 28-30: Holotype, 24.4 mm, Luanda, Angola (MNCN); 31:

Shell, 18.4 mm, Palmeirinhas (CER); 32: Shell, 16.6 mm, Nouadhibou, Mauritania; 33: paratype,

18.8 mm, Luanda (MNHN); 34-38: Guinea Conakry, shells (MNHN): 34: 5.0 mm; 35-36: 5.3

mm, 37-38: 5.4 mm.

Figuras 28-38. C. africana spec. nov. 28-30: Holotipo, 24,4 mm, Luanda, Angola (MNCN); 31:

Concha, 18,4 mm, Palmeirinhas (CER); 32: Concha, 16,6 mm, Nouadhibou, Mauritania; 33:

Paratipo, 18,8 mm, Luanda (MNHN); 34-38: Guinea Conakery, conchas (MNAN): 34: 5,0 mm; 35-

36: 5,3 mm, 37-38: 5,4 mm.

ROLÁN: The genus Calyptraea in the East Atlantic

Distribution: The species is common

and widely distributed (see Figure 117)

in Atlantic European and Mediterranean

areas (JEFFREYS, 1882; LOCARD, 1898);

United Kingdom (JEFFREYS, 1865; EALES,

1961; FRETTER AND GRAHAM, 1962, 1981;

MCMILLAN, 1973; SEAWARD, 1985, 1990),

Ireland (MINCHIN, MCGRATH AND

DUGGAN, 1987), France (BUCcQUOY,

DAUTZENBERG AND DOLLFUS, 1883,

LOCARD, 1892, DAUTZENBERG AND

FISCHER, 1925; PERRIER, 1964; BOUCHET,

DANRIGAL AND HUYGHENS, 1978; FINET,

WUEST AND MAREDA, 1992), Belgium

(BACKELJAU, 1986), Spain (HIDALGO,

1917; ROLÁN, 1993; ROLÁN AND OTERO-

ScHmITr, 1996), Portugal (NOBRE, 1940;

MACEDO, MACEDO AND BORGES, 1999),

Mediterranean (STOLFA-ZUCCHI, 1970;

PARENZAN, 1970; SABELLI AND SPADA,

1977, D'ANGELO AND GARGIULLO, 1978;

SIABEREL ET AL, 1991 RINALDI, 1991;

GIANNUZZI-SAVELLI ET AL., 1997;

ARDUINO, LOCATELLI, ORLANDO AND

REPETTO, 1995), including the Black Sea

(fide FRETTER AND GRAHAM, 1981),

North Africa (PALLARY, 1900, 1912),

Morocco (PASTEUR-HUMBERT, 1962),

Canary Is. and Madeira (NORDSIECK

AND GARCÍA-TALAVERA, 1979), and for

all European coasts (NORDSIECK, 1968;

POPPE AND GOTO, 1991). It is not present

in the eastern Channel and in the North

Sea (FRETTER AND GRAHAM, 1981). The

range is between 24” N and 56" N.

Bathymetrically, C. chinensis is fre-

quently a species collected intertidally

at low tide, and also circalittoral, with

live collected specimens in our material

from 190 m. It was recorded by the Por-

cupine Expedition in 1819 m (LOCARD,

1898), but only as empty shells.

The records of C. chinensis from South

Africa (Figs. 78-80) in BARNARD (1963),

KENSLEY (1973), KILBURN AND RIPPEY

(1982) and STEYN AND Lussi (1998)

belong to a different species. They will be

treated in a further separate paper.

Many other records of Calyptraea chi-

nensis from the West African coast corre-

spond to two other different species,

which are studied below.

Remarks: The European and north

African populations are well repre-

sented by the lectotype and paralecto-

type at LS.

Despite the variability of the species

(colour of the shell, presence or absence

of spines, size, etc.) the protoconch is

relatively constant in size and

microsculpture in most of the shells

studied. However, out of about two

hundred protoconchs examined from La

Atunara, seven had a larger size (870-

930 um, Fig. 23). At first it was thought

that it could be a different species

endemic to the Strait of Gibraltar, but no

other differences were found.

Later, we examined numerous

samples from the Canaries and Madeira

which had some peculiarities: they were

smaller and higher than European and

Mediterranean shells and more fre-

quently brown in colour, therefore

showing some similarities to C. inexpec-

tata (see below). However, the proto-

conch was similar to those studied from

France and Atlantic and Mediterranean

Spain, with similar size and sculpture.

Some shells of this population with

larger protoconchs of almost 1 mm in

diameter (Fig. 21 in comparison with

Fig. 20) were found. Similarly, some

large protoconchs were found in the

Western Sahara population (Fig. 22).

Finally, in material collected in Tombant,

Marseilles, France, Calyptraea with both

small (Fig. 25) and large (Fig. 27) proto-

conchs were found. In this case, the few

adult shells with large protoconch were

also large and very flat (Figs. 75-77),

with a width /height ratio of 6.5. This is

very unusual, but in considering

whether they were conspecific no other

differences were found.

We cannot explain why in several

populations, some unusual and scarce

shells have larger protoconchs which

are distinct from natural variation. It is

possible that, in this species with intra-

capsular development, some of the eggs

might be used as food by viable larvae.

If so, larger protoconchs would be due

to greater food availability, and so,

increased growth. TRONCOSO ET AL.

(1988) mention that in a capsule with

only two eggs, the size of both larvae

was of 1 mm, while in another capsule

Iberus, 22 (2), 2004

Figures 39-49: Calyptraea africana. 39, 40: Juvenile shells of C. africana; 39: Guinea Conakry; 40:

Ivory Coast. Figures 41-49: Protoconchs: 41: Guinea Conakry; 42, 43: Angola; 44, 45: Ghana: 46,

47: Ivory Coast; 48, 49: Banc d'Arguin, Mauritania.

Figuras 39-49: Calyptraea africana. 39, 40: Conchas juveniles de C. africana; 39: Guinea Conakry;

40: Costa de Marfil. Figuras 41-49: Protoconchas: 41: Guinea Conakry; 42, 43: Angola; 44, 45:

Ghana: 46, 47: Costa de Marfil; 48, 49: Banc d'Arguin, Mauritania.

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 50-56. Protoconch variation in C. africana: 50: from Mauritania; 51: from Senegal; 52,

53: from Guinea Conakry; 54-56: from Principe.

Figuras 50-56. Variación en la protoconcha de C. africana: 50: de Mauritania; 51: de Senegal; 52, 53:

de Guinea Conakry; 54-56: de Príncipe.

Iberus, 22 (2), 2004

with 18 larvae, the average size was 0.6 A strange shell of Calyptraea (Figs.

mm. 81, 82) was collected in La Palma,

These larger protoconchs were not Canary Islands. It seems to be very dif-

found in the populations examined from ferent from C. chinensis, but it could be

Galicia, Spain and Bretagne, France. introduced material from ships.

Calyptraea africana spec. nov. (Figs. 28-56, 86-88, 92, 95, 98, 101, 108-112)

Calyptraea chinensis auct. (non Linnaeus, 1758).

? Calyptraea solida von Martens, 1874 (non Reeve).

Type material: Holotype (Figs. 28-30) in MNCN (15.05/46621); Paratypes in the following collec-

tions: MNHAN (1), AMNH (1), BMNH (1), ZSM (1), CER (30), CJH (1), CFS (1), all from Luanda

(Angola).

Other material studied: Western Sahara: 17 s, Port Etienne, Mission Gruvel (MNHN). Mauritania:

12 sp, 3 miles W Kiaone 20? 02” N 16? 22" W, 6 m (MNHN); 6 s, 20? 10' N 16? 30” W, 16 m (MNHN);

15 sp, 2 s, 20” 20" N 16? 22” W, 10 m (MNHN); 15 c, Cansado Bay, Mission Gruvel (MNHN); 2 sp,

5 s, Nouadibhou, dredged 3 m (CER); 3 s, Baie de l' Etoile, dredged in front Club Nautique, 4 m

(CER); 155 s, 25 j, Banc d' Arguin, intertidal sediments (CER). Senegal: 2 c, Salin La Ventite (CES);

12 c, Thiaroye NNE Dakar, 7-8 m (MNHN); 1 c, N Cayar, 15” 12' 5"N 15? 54 8”"W, 25 m (MNHN);

126 s, 178 j, between Dakar and Gorée, dredging 20-40 m (CER); 3 s, Casamance, 12” 20.7” N 16?

53.1” W, 15 m,R/V “Louis Sauger” (MNHN); 1 s, 2 j, Cap Vert, Epave, 15 m (CER); 4 sp, M' Bao,

Dakar, 9-10 m (MNHN); 1 sp, 89 s, Almadies, Dakar, 20 m, on shells in sandy bottom (CER); 18 s,

Dakar (MNAHN); 2 j, Tacoma, 25 m (MNHN); 2 c, N Casamance 12* 53.8" N 17? 03.5" W, 19 m (MNHN);

3 sp, Region de Sine-Saloum, mangroves (MNHN); 3 c, N Casamance, Kafountine 12” 54.4” N 17?

01.5” W, 17 m (MNHN); 17 sp, N Casamance 12” 50.4” N 16? 58.8” W, 13-14 m (MNHN); 5 sp 4 s,

Casamance, Zinguinchor, 3-5 m (MNHN); 17 s, S Casamance, frontier of Guinea Bissau, 4-6 m

(MNHN); 1 s, Carabane Bólon, 3-4 m (MNHN); 3 c, S Casamance, Cap Skirring, in beach sediment

(MNHN). Guinea Bissau: 10 sp, Essoukkoundiak Bólon, near frontier, 5-6 m (MNHN); 2 s, Bissa-

gos Is., Mission L. Gain (MNAHN). Guinea Conakry: 60 sp, 6 s, W Ile Knebomby N/O “André Nizery”

Sedigui I, Sta. 376, 5 m, 9” 48” N 13” 53” W (MNHN); 12 s, W lle Knebomby N/O “André Nizery”

Sedigui l, Sta. 376, 5 m, 9 48' N 13? 53" W (MNHN); 36 s, W Kaporo N/O “André Nizery” Sedigui

[, Sta. 270, 6 m, 9” 36" N 13” 38” W (MNHN); 3 j, W Ile Quito, N/O “André Nizery” Sedigui Il, Sta.

515, 26 m, 10” 00' N 15? 43” W (MNHN); 2 c, NW lle Tamara, N/O “André Nizery” Sedigui l, Sta.

265, 10 m (MNHN); 2 j, SW lle Tamara, N/O “André Nizery” Chalgui 7, Tr. 17, 18 m, 9” 28' N 13?

53" W (MNHN); 3j, W Yomponi river N/O “André Nizery” Sedigui II, Sta. 724, 21 m, 10? 24” N 15?

21” W (MNHN); 12 j, frontier Sierra Leona, N/O “André Nizery” Chalgui 7, Tr. 6, 12 m, 9 01” N

13? 30” W (MNHN); 1 s, W Kaporo N/O “André Nizery” Sedigui l, Sta. 276, 18 m, 9” 36' N 14” 06'

W (MNHN); 2 s, W Kaporo N/O “André Nizery” Sedigui l, Sta. 275, 16 m, 9” 36' N 14” 03” W

(MNHN); 26 s, W Morébaya river N/O “André Nizery” Sedigui I, Sta. 168, 12 m 9” 24” N 13? 38'

W (MNHN); 5 s, 9” 40” N 14? 05” W 18 m (MNHN). Ivory Coast: 1 sp, 120 s, 6 j, Radiale Grand

Bassam N/O Antéa Benchaci I, Sta. 12D, 30 m 5* 09.2 N 3* 47.2? W (MNHN); 1 s, Batrevie Beach,

Sassandra (MNHN); 1 c, Abidjan (MNHN). Ghana: 52 sp, 22 s, 65 j, off Miamia, 30 m, on shell, in

sandy and rocky bottom (CER). Cameroun: 1 s, Wouri Cap Nachtigal 03? 44” N 09* 22” E, 13 m

(MNAHN); 1 s, Wouri Cap Nachtigal 03” 37' N 09* 16' E, 40 m (MNHN); 1 sp, Victoria /Limbé, Bota,

8-10 m (MNHN). Sáo Tomé and Principe: 1 c, Baia de Ana Chaves (MNHN); 63 s, Principe, “Calypso”,

15-18 m (MNHN); 12 s, Santo Antonio, Principe, 10 m, on sandy sediments (CER). Gabon: 2 s, Cap

Esterias-Pointe Idolo (MNHN); 2 s, Cap Esterias, Libreville (MNAHN). Congo: 1 sp, 12 s, 12 j, Pointe

Noire, Mondaine beach, 1-5:m (MNHN); 4 j, Conkouati, chalutier “Kounda”, 17-19 m (MNHN); 28

sp, 10 s, Orstom Beach, Pointe Noire, 3-7 m (MNAN); 1 sp, Pointe Noire, Songolo, 6 m (MNHN).

Angola: 1 sp, Cabo Ledo, Luanda, 10-40 m (MNHN); 39 sp, Corimba, Luanda, 20 m, on shells in

rocky bottom (CER); 3 s, Corimba, Luanda, 10-20 m (MNHN); 10 s, Cacuaco, Bengo, 10-40 m (CES);

4 sp, 3 s, Cacuaco, Bengo, 7 m, on shells in muddy bottom (CER); 1 s, Praia Etambar, beach sedimen

(MNHN); 4 sp, 1 s, Cacuaco, Bengo, infralittoral rocks (MNHN); 6 sp, 1 s, Samba, Luanda, 2 m, on

shells in sandy bottom (CER);4 sp, Samba, intertidal (CER); 5 sp, Palmeirinhas, 15-20 m, on shells

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 57-64. Calyptraea inexpectata. 57-59: Holotype, 5.3 mm, lle de Los, Guinea Conakry, 35

m (MNHN); 60, 61: Shell, Western Sahara, 5.8 mm (CJH); 62-64: Shell, Western Sahara, 4 mm

(CJH). Figures 65-67. Calyptraea chinensis, form similar to C. ¿nexpectata: 65: shell, 6.1 mm, Caja

de Lazareto, Madeira, 60 m (CFS); 66, 67: Shell, 5.8 mm, Caja de Lazareto, Madeira (CES).

Figuras 57-64. Calyptraea inexpectata. 57-59: Holotipo, 5,3 mm, lle de Los, Guinea Conakry, 35 m

(MNHN); 60, 61: Concha, Sáhara Occidental, 5,8 mm (CJH); 62-64: Concha, Sáhara Occidental,

4,0 mm (CJH). Figuras 65-67. Calyptraea chinensis, forma similar a C. inexpectata: 65: concha, 6,1

mm, Caja de Lazareto, Madeira, 60 m (CFS); 66, 67: Concha, 5,8 mm, Caja de Lazareto, Madeira

(CES).

Iberus, 22 (2), 2004

Figures 68-74: C. inexpectata. 68: Shell, Guinea Conakry, 5.5 mm; 69-74: Protoconchs; 69, 70:

From Guinea Conakry; 71: Senegal; 72: Guinea Conakry; 73: Senegal; 74: Guinea Conakry.

Figuras 68-74: C. inexpectata. 68: Concha, Guinea Conakry, 5,5 mm; 69-74: Protoconchas; 69, 70:

Guinea Conakry; 71: Senegal; 72: Guinea Conakery; 73: Senegal; 74: Guinea Conakry.

ROLÁN: The genus Calyptraea in the East Atlantic

in rocky bottom (CER); 15 s, off Luanda, 50 m, in sandy sediments (CER); 40 sp, 20 s, Mussulo,

Luanda, 10-20 m (MNHN); 1 j, Lobito, intertidal (CER); 8 sp, San Antonio, Benguela, 5-10 m (MNHN);

4 s, Praia Santiago, Bengo, beach sediment (MNHN); 1 c, Mossamedes Bay, 5-10 m (MNHN);

Type locality: Luanda, Angola.

Etymology: The specific name derived from the African coasts where the species was collected.

Description: Shell (Figs. 28-40) with a

patelliform shape and usually ovoid at

its base (circular only in juveniles),

rather fragile, transparent, smooth and

shiny. It has an elevated apex, but

usually the profile of the shell appears

to be slightly concave, and the shell is

rather depressed. Width /height ratio is

about 3.3-4.5.

The protoconch may have two dif-

ferent forms. The first one (Figs. 40-49)

has approximately 1 and */4 whorls,

with a very small nucleus which is usu-

ally 60 to 100 um in diameter. The proto-

conch diameter increases quickly, to

reach 10/12 times the size of the nucleus

at the end. The total diameter of the pro-

toconch is difficult to measure because

the protoconch-teleoconch boundary is

not evident, but may have between 700-

900 qm. Seeing this protoconch laterally,

we normally find that the periphery is

more prominent than the teleoconch on

which it is placed, resulting in a deep

suture (Figs. 41, 43, 45). The second type

of protoconch (Figs. 50-56) has only 1

whorl and a larger nucleus (85-157 ym)

and the rate of whorl expansion is

smaller (at the end the diameter of the

whorl is 3-8 times that of the nucleus)

and the total diameter of the protoconch

is between 500-750 um. There are no

clear intergradations between these two

types of protoconchs.

The teleoconch is smooth, glossy,

transparent; most of the shells have

sparse widely separated elevated

spicules, which are not usually present

in a short circle around the apex. These

spicules (Figs. 28, 33-35, 38, 39, 101) are

variable from one shell to another, more

common in some populations and rare

in others. Internally the shell is glossy.

The colour of the shell is transparent

whitish. The apex is sometimes yellow-

ish, or even dark brown, and this colour

may extend along the suture; occasional

shells may be almost entirely light

brown. Aperture ovoid with the peris-

tome usually at one level but sometimes

curved to match the surface to which it

is applied. Internally, a partition arises

from the shell along a curved line

ruming from the apex to near the aper-

ture posteriorly.

Dimensions: Variable between popu-

lations: shells from Mauritania to Ivory

Coast sometimes reach 20 mm in

maximum diameter, but one exceptional

shell from Dakar (MNHN) was 28.5

mm; in comparison, specimens from

Angola are usually larger, about 25 mm

in diameter, but can reach 28-32 mm,

with a record of 35.0 mm.

Animal (described from Ghana spec-

imens) (Fig. 95) with a rounded foot,

which has two pointed extremes on the

anterior border; the sole, seen with mag-

nification, has many very small violet

spots which give this tone to the whole;

the head is elongate posterior to the ten-

tacles, the snout being rather short and

broad; the tentacles, milk white at the

centre, sometimes with dark lines at the

external border; eyes near the base in a

posterior widening; the mantle has very

numerous and small milk-white dots,

which are grouped irregularly forming

variable blotches. The penis (Fig. 98)

(examined in some males of 5 mm) is

bilobed at its extreme, one of the lobes

being formed by two masses together; a

narrow worm-like filament is present

near the tip. In living males, the penis is

bent dorsally towards the posterior part

of the animal; the females have in this

place an atrophied stub.

Seen dorsally, by transparency, a

dark intestinal tract can be observed

surrounding the apex which curving to

the left and returning parallel to the

right side. Some Angolan specimens

observed live also fit this description.

Radula (Figs. 86-88, 92) with a

narrow rachidian tooth with a not very

prominent central cusp and up to 12

Iberus, 22 (2), 2004

Figures 75-77. Calyptraea cf. chinensis, 17.0 x 2.6 mm, Tombant, Est du Grand Congloué, Mar-

seille, 83 m (MNAHN). Figures 78-80. Calyptraea aff. chinensis, South Africa, 19.4 y 20.2 mm

(CER). Figures 81, 82. Calyptraea sp., La Palma, Canary Ids., 11.1 mm (CER).

Figuras 75-77. Calyptraea cf. chinensis, 17,0 x 2,6 mm, Tombant, Est du Grand Congloué, Marseille,

83 m (MNAN). Figuras 78-80. Calyptraea aff chinensis, África del Sur, 19,4 y 20,2 mm (CER).

Figuras 81, 82. Calyptraea sp., La Palma, Canarias, 11,1 mm (CER).

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 83-90. Radulae of Calyptraea. 83: C. chinensis, O Grove, specimen of 12.4 mm; 84, 85: C.

chinensis La Atunara, Línea de la Concepción, shell of 13.4 mm; 86-88: C. africana, Guinea

Conakry, shell of 10.0 mm; 89-90: C. inexpectata, Tacoma, Dakar, shell of 12.0 mm.

Figuras 83-90. Rádulas de Calyptraea. 83: C. chinensis, O Grove, ejemplar de 12,4 mm; 84, 85: C.

chinensis La Atunara, Línea de la Concepción, concha de 13,4 mm; 86-88: C. africana, Guinea

Conakry, concha de 10,0 mm; 89-90: C. inexpectata, Tacoma, Dakar, concha de 12,0 mm.

Iberus, 22 (2), 2004

smaller cusps on each side. Lateral tooth

slightly prominent and with a narrow

cusp in the confluence of the internal

and the external edges; internal edge

with numerous and irregular cusps

(usually up to 11), while the external

edge has more numerous cusps (22-30).

Marginal teeth elongate, curved, sharply

pointed; the internal with two borders

with up to 20 cusps on the internal

border and less on the external (4-6),

while the external teeth have up to 15

cusps on its internal part.

The spawn is formed by numerous

ovoid elongate capsules (Fig. 108-111)

which are fixed to the substrate and sat

upon by the animal. On some occasions,

when the animal was removed from the

substrate, some capsules remained

around the foot (Fig. 110). In the larger

Angolan specimens, the spawn was

composed of up to 48 capsules, with

57/90 eggs in each. In the case of a small

specimen of 12 mm, there were 18 cap-

sules with 13 - 35 eggs of 170-260 um di-

ameter. In Ghana specimens, animals be-

tween 8-13 mm had 17-26 capsules/each

and between 10-30 eggs per capsule,

with an average egg diameter of 166 um.

At hatching, the shells already have the

juvenile morphology (Fig. 112).

Distribution: The shells mentioned as

Calyptraea chinensis from West Africa or

from countries in this area are probably

best referred to the present species:

DAUTZENBERG (1912), KNUDSEN (1950)

and NICKLÉS (1950) recorded it in

several localities; NICKLÉS (1947) also for

Senegal; Lamy (1923) for Bissagos

Islands; BUCHANAN (1954) for Ghana;

TOMLIN (1923) and FERNANDES AND

ROLÁN (1993) for Sáo Tomé and

Principe; BERNARD (1984) for Gabon;

COLLINGTON (1960) for Congo; GOFAS,

AFONSO AND BRANDAO (1985) and

ROLÁN AND RYALL (1999) for Angola.

In our material, the species is repre-

sented by shells and specimens collected

from south of Western Sahara /Maurita-

nia (21.5 N) down to Angola (15* S),

including the Gulf of Guinea islands of

Sáo Tomé and Principe (Fig. 117). It was

not collected in the Cape Verde Islands

or in Annobon.

This species usually occurs in

shallow water. The deepest live-col-

lected specimens from the Ghana area

are from 20-30 m, but are usually found

in shallower waters in Mauritania,

Senegal, Guinea Conakry and Angola.

Discussion: The new species C.

africana differs from C. chinensis in nu-

merous characters (Table I). The most

important are: C. chinensis is usually

smaller, more rounded in form and

rarely appearing ovoid; the shell is more

solid at a similar size, not transparent,

more variable in colour and the external

surface is not glossy; the projections on

the external surface, when they are pre-

sent, are denser and not very prominent;

the protoconch has a larger nucleus,

larger spire (1 */4 versus */4 - 1), spiral

rows of micro-undulations, and seen

from the side the protoconch is more dis-

tinct from the teleoconch by a deeper su-

ture. The penis is also different, C. chi-

nensis has an elongate penis expanded

into two lobes at its apex like a spoon

with a thumb in opposition, while C.

africana is bilobed and has on one side

two masses together and a narrow fila-

ment. The spawns are also different,

with capsules more elongate in C.

africana and more spherical sometimes

slightly ovoid in C. chinensis; the eggs are

smaller in C. africana than in C. chinensis.

KNUDSEN (1950) probably examined

the spawn of C. africana because one of

the populations mentioned was col-

lected at 17 m, in the middle of the dis-

tribution of this species. The other two

populations were probably from a dif-

ferent species because C. africana has not

been collected alive at 34-50 m.

The presence of two kinds of proto-

conchs with different characters without

intergradations made us suspect that we

could be working with two different

species. This could correspond to two

sympatric sibling species. Furthermore,

the most common type of protoconch,

the larger one, is the only form observed

in Angolan populations; the smaller one

appears more frequently in the Principe

population and also in shells from

Ghana, Ivory Coast, Guinea Conakry

and Senegal, but is always present in

ROLÁN: The genus Calyptraea in the East Atlantic

Table I. Differences in characters of the shell, protoconch, radula and spawn of Calyptraea chinen-

sis, C. africana spec. nov. and C. ¿nexpectata spec. nov.

Tabla I. Diferencias entre los caracteres de la concha, protoconcha, rádula y puesta de Calyptraea chi-

nensis, C. africana spec. nov. y C. inexpectata spec. nov.

Profile of the shell

Form of the aperture

Border of the shell aperture

Transparency of the shell

Shell width / height ratio

Most common coloration

Other possible colorations

External sculpture

Usual shell diameter

Maximum diameter

Protoconch: diameter

Protoconch: diameter of the nucleus

Protoconch: number of whorls

Diameter nucleus/ diameter at the

end of the protoconch ratio

Elevation of the protoconch

from the teleoconch

Protoconch sculpture

Sculpture at the beginning of teleoconch

Spawn: capsules

Eggs size

Animal mantle

Penis

Depth limits of the live

collected specimens

C. chinensis

hooked

rounded

uniform

2.5-3.1

white

pink, brown, violet

usually smooth, sometimes

with numerous prominences

up to 15 mm

25.0 mm

600-741 jm

194-230 jm

3/4-1

3-3.5

slightly elevated and

suture not deep

spiral bands of axial striae

spherical scarcely ovoid

340-490 jm

transparent with some white

spots grouped in the border

bilobed with distally a

opposing thumb on a spoon

0-19 m

C. africana Spec. nov.

tendency to be concave

ovoid in adult

uniform

yes

3.3-4.0

white or white with the

apex yellowish or brown

subsutural brovn band

usually smooth, sometimes

with very elevated and

separate prominences

up to 25 mm

35.0 mm

form 1: 700-900 jm

form 2: 500-750 m

form 1: 60-100 um

form 2: 89-157 pm

form 1: 11/4

form 2: 1

form 1: 10-12

form 2: 3-8

elevated with deep suture

ovoid

160-270 um

transparent with irregular

white spots

bilobed like two fused

eggs, with other

prominence and a fine

worm like filamente

3-30m

C. inexpectata spec. nov.

tendency to be convex

rounded irregularly

frequently undulate

or irregular

1.8-2,4

totally white or brown

apical brown

sometimes wrinkled,

frequently numerous scarcely

elevated prominences

like scales

up to 7.5 mm

12.0 mm

471-614 um

142-215 pm

2.1-4,5

slightly elevated and

suture not deep

Spiral grooves

unknown

transparent with very evident

radiating milk white lines

at the border

elongate, straight, with

two angulations

25-93 m

Iberus, 22 (2), 2004

small numbers (about 10%). For this

reason, we undertook a detailed study

to find other differences which can

usually be found in two sibling species.

In the present case, the comparison of

animals, shell characters, radula, etc. did

not show other differences and it was

not possible to prove that these different

protoconchs were the result of sampling

two species. For the time being we

decided to consider the shells with these

two kinds of protoconchs as conspecific.

The holotype is a shell with the large

type of protoconch, from Luanda,

Angola, an area where the second type

of protoconch has never been found.

Currently there is no explanation for

this difference, and it is expected that

the future study of DNA will provide

additional information.

Calyptraea inexpectata spec. nov. (Figs. 57-74, 68-74, 89, 90, 93, 96, 99, 102)

Type material: Holotype (Figs. 57-59) deposited in MNHN with 1 paratype from type locality; 10

paratypes more, from W Kaporo, Sta. 275, 16 m, 9” 36" N 14” 03” W; 2 more from W Pte. Goro, Sta.

544, 41 m, and 1 more from W Cap Verga, Sta. B>CH, 20 m, 10 12" N 13? 06' W, all them from N/O

“André Nizery” Sedigui l and II. Other paratypes, from 40 m between Dakar and Gorée, in the fol-

lowing collections: MNCN (15.05 /46622) (1), AMNH (1), BMNH (1), ZSM (1), CER (68), CJH (1),

CFS (1).

Other material examined: Western Sahara: 2 s, 23” 05' N 16” 00' W, 27-36 m (C]H); 1 sp, (CJH); 1

s, 22? 35 N 16 58” W, 86 m (CJH); 2 s, 23 05” N 167 00 W, 25-50 m, (CJH); 1 sp, (CJH); 3 s, 22? 35'

N 16? 58” W, 86 m (C]H). Mauritania: 1 s, 17” 42” N 16* 12 W, 46 fms, (CJH); 8 s, 17” 22 N 16" 17'

W, dredged at 70-100 m (CJH); 2 s, (MNHN); 1 s, N/O N' Diago 18” 12” N 16? 20” N, 50 m

(MNHN); 1 s, N/O N' Diago 19* 12 N 16? 40' N, 47 m (MNHN); 3 s, N/O N' Diago 19* 06' N 16?

34' N, 37 m (MNHN) 1 s, N/O N' Diago 17” 42 N 16? 25' N, 50 m (MNHN); 4 s, N/O N' Diago

17* 18' N 16 26" N, 76 m (MNHN); 1 s, N/O N' Diago 17? 36' N 16" 23” N, 65 m (MNHN); 1 s,

N/O N' Diago 19* 00' N 16? 30” N, 31 m (MNHN); 1 c, 17? 17” N 16? 30” W, 85 m (MNHN) 1 s,

17* 45' N 167 23" W, 300-600 m (CFS). Senegal: 1 sp, 12 s, Tacoma, 25 m (CER); 2 s, 3 j, Cap Vert,

Epave, 15 m (CER); 5 sp, 225 s, SW lle Madeleine, Dakar, 48 m (MNHN); 3 j, SW Madeleines,

Dakar, 455-465 m (MNHN >); 4 c, Dakar 14* 23' 5"N 17? 24' 5"W, 65-70 m (MNHN); 53 s, Gorée, 20

m (CES); 3 sp, dredged near Goree, 15-25 m (CER); 20 s, S Gorée, Dakar, 33-42 m (MNHN); 16 s,

6 j, S Gorée, 98 m (MNHN); 6 s, Dakar 14” 27” N 17? 33” W, 145-200 m (MNHN); 13 j, Dakar, 6 m

(CER); 8 s, 8 j, Dakar, 20 m (CER); 15 s, E Gorée, 20 m (MNHN); 49 s, S Gorée, “G. Treca”, 65 m

(MNAHN); 1 sp, 14 s, 8 j, Tacoma, 20-25 m (CER); 5 s, Delta of Saloum, (CJH); 1 sp, N Casamance

13” 01.8" N 17? 25.5" N “Louis Sauger”, 533 m (MNHN); 1 s, Casamance, 12” 20,7” N 16? 53.1” W,

R/V “Louis Sauger”, 15 m (MNHN); 2 s, N Casamance 12” 32” N 17” 28.8" N “Louis Sauger”, 45

m (MNHN); 18 sp, N Casamance 12” 46.9 N 17” 29.9 N “Louis Sauger”, 45 m (MNHN); 39 s, 40

j, between Dakar and Gorée, dredged at 20-40 m (CER). Guinea Conakry: 1 s, W Kaporo N/O

“André Nizery” Sedigui I, Sta. 276, 18 m, 9” 36' N 14” 06” W (MNHN); 5 c, 3 j, Baie de Sangarea

N/O “André Nizery” Sedigui Il, Sta. B12-13CH, 35 m, 9” 42” N 15” 33" W (MNHN); 1 s, W lle

Tannah, Sedigui I, Sta. 84. 9” 12” N 13? 49.5” W, 33 m (MNHN); 1 s, (CJH); 1 s, 099 56' N 15? 58”

W, 36 m (CJH). Benin: 2 s, Ouidah 06* 10' N 02 05' E, 200 m (UNHN).

Type locality: lle de Los, Guinea Conakry, Expedition Sedigui l, Sta. 262, 9? 30” N 13? 59” W, 35 m.

Etymology: The specific name alludes to the fact that this species appeared while comparison between

C. chinensis and C. africana was being made, and was not expected when the study was begun.

Description: Shell (Figs. 57-64, 68)

with a patelliform shape and usually

rounded at its base, relatively solid, not

transparent, smooth but not shiny,

sometimes with a rough surface. It has

an elevated apex, and the profile usually

shows a right or slightly convex line.

The shell is not depressed. Ratio

between width and height is between

1.8 and 2.4. The protoconch (Figs. 69-74)

has approximately 1 whorl, with a

nucleus of 142-215 um in diameter.

From this nucleus, the protoconch diam-

eter increases quickly, to about 2.7-4.5

times that of the nucleus. The total

diameter of the protoconch in the mater-

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 91-93: Drawing of the radulae of Calyptraea. C: central tooth; L: lateral tooth; M1, M2:

marginal teeth. 91: C. chinensis, specimen of 13 mm, Vigo, Spain; 92: C. africana, specimen of 11

mm, Luanda, Angola; 93: C. inexpectata, specimen of 12 mm, Tacoma, Dakar.

Figuras 91-93: Dibujo de los dientes radulares de Calyptraea. C: diente central; L: diente lateral; MI,

M2: diente marginal. 91: C. chinensis, ejemplar de 13 mm, Vigo, España; 92: C. africana, ejemplar

de 11 mm, Luanda, Angola; 93: C. inexpectata, ejemplar de 12 mm, Tacoma, Dakar.

Iberus, 22 (2), 2004

Figures 94-96: Diagram of the animal of Calyptraea. 94: C. chinensis, Vigo, Spain; 95: C. africana,

Miamia, Ghana; 96: C. inexpectata, Dakar, Senegal. Figures 97-99. Penis of Calyptraea: 97: C. chi-

nensis upper row, from Vigo, Spain, specimens; lower row: first 2, from O Grove, Galicia, Spain;

the last 2 from La Atunara, Spain; 98: C. africana: upper row and first of the second from Miamia,

Ghana; the rest form Luanda, Angola; 99: C. inexpectata, Dakar, Senegal.

Figuras 94-96: Esquema del animal de Calyptraea. 94: C. chinensis, Vigo; 95: C. africana, Miamia,

Ghana; 96: C. inexpectata, Dakar, Senegal. Figuras 97-99. Pene de Calyptraea: 97: C. chinensis fzla

superior, ejemplares de Vigo, España; fila inferior: los 2 primeros, de O Grove, Galicia, España; los 2

últimos, de La Atunara, España; 98: C. africana: fila superior y primero de la segunda fila de Miamia,

Ghana; el resto de Luanda, Angola; 99: C. inexpectata, Dakar, Senegal.

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 100-102. Sculpture of the shells. 100: Calyptraea chinensis, 101: C. africana; 102: C. inex-

pectata.

Figuras 100-102. Escultura de las conchas. 100: Calyptraea chinensis; 101: C. africana; 102: C.

inexpectata.

ial studied was 471-614 um. In lateral

view the periphery of the protoconch is

not very prominent from the teleoconch,

and the suture is shallow (Figs. 72, 73).

The protoconch surface lacks any sculp-

ture except for growth lines.

The teleoconch has at its beginning

numerous growth lines crossed by spiral

grooves (Figs. 69, 70, 74) which radiate

from the protoconch;, the teleoconch is

opaque and not glossy, and in many

shells spiral striae exist. Most of the

shells have numerous wavy scales (Fig.

102), closer between them, which are

usually absent in a short circle around

the apex (Fig. 68). These scales are vari-

able among populations, and are some-

times rare. Internally the shell is glossy.

The colour of the shell is whitish or

brown, usually uniform, each shell with

a single colour. Aperture rounded with

the peristome frequently irregular,

perhaps due to irregular substrate.

Internally, a partition arises from the

shell along a curved line running from

the apex to near the aperture posteriorly.

Dimensions: in most of the popula-

tions the shells are only 5-7 mm, but in

some, the largest shells reach 12 mm in

maximum diameter.

The animal (Fig. 96), examined in

five specimens (three females and 2

males) of the Dakar population, is

whitish translucent with numerous

milk-white Oopaque spots on the tenta-

cles, the lips and the body. The mantle

has very marked milk-white radiated

fusiform blotches which reach the

border. All of them are fused into a con-

tinuous line a little below the border,

but continue towards the interior in

other finer lines. The buccal lips have a

darker tone. The males are very small (3

mm diameter). The penis observed in

the two males studied (Fig. 99) is behind

the right tentacle and difficult to see

because it is in a dorsal position; it is

cylindrical and simple, having two

angulations.

Radula (Figs. 89, 90, 93) with a

rachidian tooth narrow and with the

central cusp not very prominent and up

Iberus, 22 (2), 2004

112

105

110 111

Figures 103-112. Spawn and larval specimens of Calyptraea. Figures 103-107. C. chinensis from Vigo.

103, 104: egg capsules; 105: group of egg capsules. 106, 107: larval specimens after the veliger period.

Figures 108-112. C. africana from Miamia, Ghana. 108, 109: egg capsules; 110: position of the eggs

with the animal on the substrate; 111: spawn removed the animal; 112: larval specimen.

Figuras 103-112. Puesta y ejemplares larvarios de Calyptraea. Figuras 103-107. C. chinensis de Vigo.

103, 104: cápsulas; 105: grupo de cápsulas; 106, 107: ejemplares larvarios posteriores a la etapa velígera.

Figuras 108-112. C. africana de Miamia, Ghana. 108, 109: cápsulas; 110: posición de las cápsulas con

el animal sobre el sustrato; 111: puesta una vez retirado el animal; 112: ejemplar larvario.

to 5 smaller cusps at each side. Lateral

teeth with a cusp in the confluence of

the internal and the external edges

which has the same size as those in the

internal edge (usually up to 5) while the

external edge has more numerous cusps

(16-20). Marginal teeth elongate, curved,

sharply pointed and the internal ones

with up to 22 cusps on the internal

border and 4-6 on the external. The

external teeth have up to 6 cusps on its

internal part.

The spawn of C. inexpectata was

probably examined by KNUDSEN (1950),

because two of the populations men-

tioned in this work are from 34 and 50

m, Which is the usual depth for this

species. He mentioned 23 and 28 cap-

sules per brood, with 9-29 eggs per cap-

sule and a total between 239-274 eggs.

Only one spawn of this species could be

examined, but in poor conditions to

count and measure capsules and eggs.

Distribution: C. inexpectata is known

from Western Sahara (23? N) and Mauri-

tania to Benin (6 N) (Fig. 117). It was

not found in Ghana in spite of the many

samples obtained from this country.

ROLÁN: The genus Calyptraea in the East Atlantic

Figures 113-116. Types of Calyptraea chinensis in LS: 113, 114: Lectotype; 115, 116: Paralecto-

ope:

Figuras 113-116. Tipos de Calyptraea chinensis en LS: 113, 114: Lectotipo; 115, 116: Paralectotipo.

This species has not been found in

shallow water. The living material has

been collected from 25 to 53 m, and

empty shells from up to 600 m.

Discussion: C. inexpectata is different

from the other species mentioned in the

present work. The differentiating char-

acters are listed in Table l, but they will

be commented as follow:

C. chinensis is usually larger; the

sculpture (when it is present) is more

prominent and less dense; the shell

width / height ratio is larger (2.5-3.1 vs.

1.8-2.4); the protoconch is rather similar,

but a little larger and also the nucleus

diameter is «slightly larger, showing

microsculpture which does not exist in

C. inexpectata. The beginning of the pro-

toconch in C. chinensis lacks the grooves

present in C. inexpectata. The radula has

fewer cusps on the lateral tooth, and the

cusp in the confluence of the internal

and external borders is larger; also,

there are fewer cusps on the marginal

teeth; the central tooth is wider.

C. africana has a larger shell, usually

about 25 mm in diameter, and up to 28-32

mm, while C. inexpectata seldom reaches

Iberus, 22 (2), 2004

Co chinensis

C. inexpectata

UE C. eficana

Figure 117. Distribution range of C. chinensis, C. africana and C. inexpectata.

Figura 117. Área de distribución de C. chinensis, C. africana y C. inexpectata.

7Ó

ROLÁN: The genus Calyptraea in the East Atlantic

12 mm. Also, C. africana is ovoid in its

aperture, more depressed, more transpar-

ent, shiny, with external sculpture (when

present) formed by isolated and raised

spicules, in contrast to the scaly concen-

tric lines of C. inexpectata. The protoconch

of C. africana is larger (in the commonest

form), with a smaller nucleus, more

prominent and deeper suture between

the protoconch and the teleoconch. The

radula has more cusps and some differ-

ent details, such as the larger cusp in the

ACKNOWLEDGEMENTS

The author would like to thank the

persons and institutions who provided

the material used in this work: Gonzalo

Rodríguez Casero for North Spain;

Jacques Pelorce for Senegal; Rosina

Acuña for Spain; Juan Carlos Sotelo for

France; José María Hernández for

Canary ls., Western Sahara and Maurita-

nia; Frank Swinnen for Canary ls.,

Morocco and Madeira; Teresa Borges

and Pedro Nunez for Portugal, and

MNHN. Kathie Way and Amelia

MacLellan sent information and pho-

tographs of the type material of Calyp-

BIBLIOGRAPHY

ARDUINO, G., LOCATELLI, B., ORLANDO, F. AND

REPETTO, G., 1995. Catalogo illustrato delle

conchiglie marine del Mediterraneo. Amici del

Museo “F. Eusebio”, Alba, 198 pp.

BaAccI, G., 1951. L'ermafroditismo di Calyptraea

chinensis L. e di altri Calyptraeidae. Pubbli-

catione della Stazione Zoologica di Napoli, 23: 66-

90.

BACKELJAU, T., 1986. Lijst van de recente mariene

mollusken van Belgie. Documents de Travail,

29: 1-106 pp.

BANDEL, K., 1984. The radulae of Caribbean

and other Mesogastropoda and neogas-

tropoda. Zoologische Verhandelingen, 214: 1-

188, 22 pls.

BARNAROD, K. H., 1963 “1962”. Contribution to

the knowledge of the South African marine

Mollusca. Part III. Gastropoda: Proso-

branchiata: Taenioglossa. Annals of the South

African Museum, 47 (1): 1-199.

BERNARD, P. A., 1984. Coquillages du Gabon.

Pierre A. Bernand, Libreville, Gabon, 140 pp.

confluence of the external and internal

edges of the lateral tooth.

These two species co-occur, in

Sahara-Mauritania, Senegal and Guinea

Conakry. C. africana was collected in

beach sediments at a few meters depth,

while C. inexpectata was collected in

deeper water, at least 30-50 m, but

sometimes deeper (shells collected fre-

quently in 70-100 m, up to 600 m),

where no shells of C. africana were

found.

traea chinensis at LS. Rachel Collin and

Angel Luque made suggestions and

Javier Conde corrected English. Jesús

Méndez of the Centro de Apoyo Cientí-

fico y Tecnológico a la Investigación

(CACTI) of the University of Vigo made

the SEM photographs. The colour pho-

tographs were made with the help of

Jesús S. Troncoso in the Departamento

de Ecología of the University of Vigo.

This work was supported by a grant

from the Ministry of Science and Tech-

nology of Spain (reference VEM 2003-

20047).

BOUCHET, P., DANRIGAL, F. AND HUYGHENS, C.,

1979. Living seashells. Molluscs of the English

Channel and Atlantic Coasts. Blandford Press,

Poole, Dorset, 144 pp.

BUCHANAN, J. B., 1954. Marine Molluscs of the

Gold Coast West Africa. Journal of the West

African Science Association, 1 (1): 30-45, 1 pl.

BucQuoY, E., DAUTZENBERG, PH. AND DOLL-

FUS, G., 1883. Les mollusques marins de Rous-

sillon. Tome I: Gastropodes avec atlas de 66

planches. J.-B. Bailliere and Fils, Paris, 570

pp., 66 pls.

CASTAÑO, M. J., CIVIS, J. AND GONZÁLEZ DEL-

GADO, J. A., 1988. Los moluscos del Plioceno

de la Palma del Condado y Moguer (Huelva).

Aproximación paleoecológica. Iberus, 8 (2):

173-186.

CHIARELLL S., 2002. Nuovo catalogo delle conchiglie

marine del Mediterraneo. Societá Italiana di

Malacologia, Roma, no pages number.

Iberus, 22 (2), 2004

COLLINGTON, J., 1960. Observation faunistique

et écologiques sur les Mollusques testacés

de la baie de Pointe-Noire. Bulletin de

l'LF.A.N., 22, sér. A (2): 411-461.

D'ANGELO, G. AND GARGULLO, S., 1978. Guida

alle conchiglie Mediterranee. Fabrici Ed., Milano,

223 pp.

DAUTZENBERG, P., 1912. Mollusques Marins.

Annales de l'Institut Oceanographique, 5, fasc.

3: 1-111, 3 pls.

DAUTZENBERG, P. AND FISCHER, P. H., 1925. Les

mollusques marins du Finistere et en particulir

de la Région de Roscoff. Les Presses Univers.

de France, Paris, 180 pp.

EALES, N. B., 1961. The littoral fauna of the British

Islands. Cambridge University Press, Cam-

- bridge, 306 pp.

FERNANDES, F. AND ROLÁN, E., 1993. Moluscos

marinos de Sáo Tomé y Príncipe: actual-

ización bibliográfica y nuevas aportaciones.

Iberus, 11 (1): 31-47.

FINET, Y., WUEST, J. AND MAREDA, K., 1992. Gas-

tropods of the Channel and Atlantic Ocean: Shells

and Radulas. L'Informatore Piceno, Ancona,

75 pp-

FISCHER, P. H., 1950. Remarques sur la systé-

matique et la phylogénie du genre Calyp-

traea. Journal de Conchyliologie, 90: 231-234.

FRETTER, V. AND GRAHAM, A., 1962. British

Prosobranch Molluscs. Ray Society, London,

759 pp:

FRETTER, V. AND GRAHAM, A., 1981. The proso-

branch molluscs of Britain and Denmark.

Part. 6, Cerithiacea, Strombacea, Hipponi-

cacea, Calyptraeacea, Lamellariacea,

Cypraeacea, Naticacea, Tonnacea, Het-

eropoda. The Journal of Mollusca Studies, supl.

9: 285-363.

GIANNUZZI-SAVELLI, R., PUSATERI, F., PALMERI,

A. AND EBREO, C., 1997. Atlante delle conchiglie

marine del Mediterraneo. Vol. 2. La Conchiglia,

Roma, 258 pp.

GOFAS, S., AÁFONSO, J. P. AND BRANDAO, M.,,

1985. Conchas e moluscos de Angola. Universi-

dad de Agostinho Neto/Elf Aquitaine, An-

gola, 139 pp.

HIDALGO, J. G., 1917. Fauna malacológica de Es-

paña, Portugal y las Islas Baleares. Moluscos

testáceos marinos. Junta para Ampliación de

Estudios e Investigaciones Científicas, Serie

Zoológica, 30, Madrid, 752 pp.

JEFFREYS, J. G., 1865. British Conchology, vol 3.

John van Voorst, London, 393 pp, 8 pls.

JEFFREYS, J. G., 1882. On the Mollusca procured

during the “Lightning” and “Porcupine” Ex-

peditions. Part V. Proceeding of the Zoological

Society, London, 1882: 656-687.

KENSLEY, B., 1973. Sea-shells of southern Africa.

Gastropods. Maskew Miller Ltd., Cape Town,

236 pp.

KILBURN, R. AND RIPPEY, E., 1982. Sea Shells of

Southern Africa. Macmillan South Africa, Jo-

hamnesburg, 249 pp.

KNUDSEN, J., 1950. Egg capsules and develop-

ment of some marine prosobranchs from

tropical West Africa. Atlantide Report, 1: 85-

130.

Lamy, M. E., 1923. Mollusques Testacés. Comptes

rendus du Congres des Societés savantes en 1922:

1-16.

LEBOUR, M. V., 1937. The eggs and larvae of the

British Prosobranchs with special reference

to those living in the plankton. Journal of Ma-

rine Biology Association U. K., 22: 106-166.

LOCARD, A., 1892. Les coquilles marines des

cótes de France. Annales de la Société Lin-

néenne de Lyon, 37: 1-385.

LOCARD, A., 1898. Expeditions scientifiques du

Travailleur et du Talisman. Mollusques Tes-

tacés. vol 2. Mason et Cie., Paris, 515 pp., 8 pls.

MACEDO, M. C. C., MACEDO, M. I. C. AND

BORGES, J. P., 1999. Conchas Marinhas de Por-

tugal. Verbo, Lisboa, 516 pp.

MCMILLAN, N. F., 1973. British Shells. Frederick

Warne and Co., London, 196 pp.

MINCHIN, D., MCGRATH, D. AND DUGGAN, C.

B., 1987. Calyptraea chinensis (Mollusca, Gas-

tropoda) on the West coast of Ireland: a case

of accidental introduction?. Journal of Con-

chology, 32: 297-301.

NICKLESs, M., 1947. La collection de mollusques

testacés de l'L.F.A.N. Catalogues 1. Institut

Francais d'Afrique Notre. Catalogues I: 1-23.

NICKLES, M., 1950. Mollusques testacés marins de

la cóte occidentale d“Afrique. Manuels ouest-

africains, 2, Lechevalier, 269 pp.

NOBRE, A., 1940. Fauna malacológica de Portugal,

[. Moluscos marinhos e das águas salobras. Com-

panhia Editora do Minho, Barcelos, 806 pp,

87 pls.

NORDSIECK, F. AND GARCÍA-TALAVERA, F., 1979.

Moluscos marinos de Canarias y Madera (Gas-

tropoda). Aula de Cultura de Tenerife, 208

pp., 46 pls.

NORDSIECK, F., 1968. Die europáischen-Meeres

Gehíiuseschneken. G. Fischer, Sttutgart, 273

pb:

PALLARY, P., 1912. Exploration scientifique du

Maroc. Mission Zoologique. Malacologie.

Empire chérifien. Archieve scientifique du Pro-

tectorat francaise, 2: 1-108, 1 pl.

PALLARY, P., 1900. Coquilles marines du lit-

toral du Départment d'Oran. Journal de

Conchyliologte, 48 (3): 211-422, pl. 6-8.

PARENZAN, P., 1970. Carta d'identita delle

conchiglie del Mediterraneo, vol. 1. Bios Taras,

Taranto, 283 pp.

PASTEUR-HUMBERT, C., 1962. Les mollusques

marins testacés du Maroc. Travaux de l'Insti-

tut Scientifique Chérifien, ser. Zoologie, 23: 1-245.

ROLÁN: The genus Calyptraea in the East Atlantic

PERRIER, R., 1964. La faune de la France en tableaux

synoptiques illustrés, 9. Bryozoaires,

Braquiopodes, Mollusques, Protocordés. Dela-

grave, Paris, 170 pp.

POPPE, G. T. AND GOTO, Y., 1991. European

Seashells. C. Hemmen, Wiesbaden, 352 pp.

RINALDI, E., 1991. Le conchiglie della costa ro-

magnola. Essegi, Ravenna, 189 pp.

ROLÁN, E. AND RYALL, P., 1999. Checklist of

the Angolan marine molluscs. Reseñas Mala-

cológicas, 10: 5-132.

ROLÁN, E. AND OTERO-SCHMITT, J., 1996. Guía

dos moluscos de Galicia. Galaxia, Vigo, 318 pp.

ROLÁN, E., 1983. Moluscos de la Ría de Vigo, l.

Thalassas, 1, suplemento 1: 1-383.

SABELLI, B. AND SPADA, G., 1977. Guida illus-

trata all'identificazione delle conchiglie del

Mediterraneo. Supplemento a “Conchiglie”, 12

(28) 1, pl

SABELLI, B, GAANNUZZI-SAVELLI, R. AND BEDULLI,

D., 1991. Catalogo annotato dei molluschi marini

del Mediterraneo. Libreria Naturalistica Bolog-

nese, Bologna, 348 pp.

SEAWARD, D. R., 1985. Sea area atlas of the Ma-

rine Molluscs of Britain and Ireland. The Con-

chological Society, London, 53 pp + 756 figs.

SEAWARD, D. R., 1990. Distribution of the marine

molluscs of north west Europe. The Concho-

logical Society of Great Britain and Ireland,

London, 114 pp.

STEYN, G. S. AND LUSSI, M., 1998. Marine Shells

of South Africa. Ekogilde, Hartebeespoort,

264 pp.

STOLFA ZUCCHI, M. L., 1970. Gasteropodi recenti

dell'Adriatico Settentrionale tra Venezia e

Trieste. Memorie Museo tridentino di Scienze

naturali, 19 (1): 123-243.

TOMLIN, J. R., 1923. The marine mollusca of Sáo

Tomé. IL. Journal of Conchology, 17 (3): 81-94.

TRONCOSO, J. S., URGORRI, V. AND CRISTOBO, F.,

1988. Observaciones sobre el comportamiento

larvario de Calyptraea chinensis en las costas

de Galicia. Iberus, 8 (2): 115-120.

VERA-PELÁEZ, J. L., LOZANO, FRANCISCO, M. C.,

MUNIZ-SOLÍS, R., GILI, C., MARTINELL, J.,

DOMENECH, R, PALMOVIST, P. AND GUERRA-

MERCHÁN, A., 1995. Estudio preliminar de la

malacofauna del Plioceno de Estepona

(Málaga, España). Iberus, 13 (2): 93-117.

VERDUIN, A., 1976. On the systematics of recent

Rissoa of the subgenus Turboella Gray, 1847,

from the Mediterranean and European At-

lantic coasts. Basteria, 40: 21-73.

WYATT, H. V., 1957. The reproduction, growth

and distribution of Calyptraea chinensis. Chal-

lenger Society Report, 3: 33-41.

de Es

> ñ

vismshA AH sd ns aa E

] =—

ira

7 mood 0 y

> ar

15 lar y pan

Dep

¿(de

LARES e

sd] AACOFIS y

tz 7 ">

nn 2 AC AIUTINA

TAN EN Aa A 1

MARTA cid * ]

NORMAS DE PUBLICACIÓN

* La revista Zberus publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos relacio-

nados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de texto, meca-

nografiadas, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son tra-

bajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas como un suple-

mento de /berus. Los autores interesados en publicar monografías deberán ponerse previamente en contacto con el

Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha sido publicado, ni se publicará en

otra parte ni en su totalidad ni resumido.

* Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: Serge Gofas,

Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n,

29071, Málaga, España.

* El texto del trabajo podrá estar redactado en cualquier lengua culta moderna.

* En los trabajos que superen las 20 páginas, cada página de exceso será abonada por el(los) autor(es) a precio de coste.

* Los artículos, notas y monografías deberán presentarse mecanografiados sobre DIN A-4, por una sola cara a doble

espacio (incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra-

bajos se entregarán por triplicado (original y dos copias), y se incluirá una versión en disco flexible (diskette), utili-

zando procesadores de texto en sus versiones de DOS o Macintosh. En caso de autoría compartida, uno de los auto-

res deberá hacerse responsable de toda la correspondencia.

* Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los que el

propio Editor considere oportunos.

* Los manuscritos se presentarán de acuerdo al siguiente esquema:

Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traducción

al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a familia o

táxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del autor o autores,

las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés. Dicho resumen deberá

sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del artículo; se sugiere una exten-

sión de 100 a 200 palabras. El resumen deberá estar seguido de una lista de palabras clave, también con su versión

inglesa para su inclusión en los bancos de datos internacionales.

Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabeza-

mientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y métodos,

Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no habituales en

el texto, deberán indicarse tras el apartado de Material y Métodos.

* Las notas breves deberán presentarse de la misma forma, pero sin resumen.

+ Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos

Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un táxon aparezca por primera

vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se den las sinonimias

de los táxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la publicación donde fueron des-

critas, y la localidad tipo si es conocida entre corchetes, según el siguiente esquema (préstese especial cuidado a la pun-

tuación):

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.

Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto.

Si se incluyen una lista completa de referencias de un taxon inmediatamente tras éste, deberá seguirse el mismo esque-

ma (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto).

* Sólo los nombres en latín y los de táxones genéricos y específicos deberán llevar subrayado sencillo o preferentemente

ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni siquiera el Título. Las

unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correctas abreviaturas. En artículos

escritos en castellano, «en los números decimales sepárese la parte entera de la decimal por una coma inferior (.),

NUNCA por un punto (.) o coma superior (*).

* Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER Y

GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto [Smith,

Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publicado más de

un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. cit. La lista de refe-

rencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se citarán los nombres de todos

los autores de cada referencia, sea cual sea su número. Los nombres de los autores deberán escribirse, en letras minús-

culas o VERSALITAS. No deberán incluirse referencias a autores cuando éstos aparezcan en el texto exclusivamente

como autoridades de un taxon. Los nombres de las publicaciones periódicas deberán aparecer COMPLETOS, no

abreviados. Cuando se citen libros, dése el título, editor, lugar de publicación, n* de edición si no es la primera y

número total de páginas. Deberán evitarse referencias a Tesis Doctorales u otros documentos inéditos de difícil con-

sulta. Síganse los siguientes ejemplos (préstese atención a la puntuación):

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. 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 Zoolgica, 3 (5): 21-51.

* Las gráficas e ilustraciones deberán ser originales y presentarse sobre papel vegetal o similar, con tinta china negra y

ajustadas al formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm

(dos) de anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. En caso de pre-

parar figuras para que ocupen el total de una página, se ruega ajustar su tamaño para que puedan caber los pies de

figura bajo ella. Si han de incluirse gráficas de ordenador, deberán imprimirse con impresora láser sobre papel de

buena calidad. Las fotografías, bien contrastadas y sin retocar, deberán ajustarse siempre a los tamaños mencionados.

Al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que estén debidamente

alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma página conlleva una pobre

reproducción final. Las escalas de dibujos y fotografías deberán ser gráficas, y las unidades que se utilicen del sistema

métrico decimal. Considérese la reducción que será necesaria a la hora de decidir el tamaño de las escalas o letras en

las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte deberá etiquetarse con letras mayús-

culas, el resto de las letras deberán ser minúsculas. No deberán hacerse referencias a los aumentos de una determi-

nada ilustración, ya que éstos cambian con la reducción, por lo que debe emplearse una escala gráfica. En su caso, se

recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse en

hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de figura, en una

hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua del trabajo). Utilícese

el esquema siguiente:

Figura 1. Neodoris carví. A: animal desplazándose; B: detalle de un rinóforo; C: branquia.

Las abreviaturas empleadas en las ilustraciones deberán incluirse en la hoja de pies de figura.

Los autores interesados en incluir láminas en color deberán abonarlas a precio de coste (30.000 ptas por página). Por

lo demás, deberán ajustarse a los mismos requisitos que los indicados para las figuras.

Si se pretende enviar gráficas o ilustraciones en formato digital es IMPRESCINDIBLE contactar previamente con el

Editor

* Las Tablas se presentarán en hojas separadas, siempre con numeración romana (l, II, IIL...). Las leyendas se inclui-

rán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particularmente com-

plejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo necesario.

* Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de los cam-

bios necesarios.

+ El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha de envío

a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El Comité Editorial, a la

vista de los informes de los revisores decidirá sobre la aceptación o no de cada manuscrito. El autor recibirá en cada

caso copia de los comentarios de los revisores sobre su artículo. En caso de aceptación, el mismo Comité Editorial, si

lo considera conveniente, podrá solicitar a los autores otras modificaciones que considere oportunas. Si el trabajo es

aceptado, el autor deberá enviar una copia impresa del mismo corregida, acompañada por una versión en disco flexi-

ble (diskette), utilizando procesadores de texto en sus versiones de DOS o Macintosh. La fecha de aceptación figura-

rá en el artículo publicado.

e Las pruebas de imprenta serán enviadas al autor responsable, EXCLUSIVAMENTE para la corrección de erratas, y

deberán ser devueltas en un plazo máximo de 15 días. Se recomienda prestar especial atención en la corrección de las

pruebas.

* De cada trabajo se entregarán gratuitamente 50 separatas. Aquellos autores que deseen un número mayor, deberán

hacerlo constar al devolver las pruebas de imprenta, y NUNCA POSTERIORMENTE. El coste de las separatas adi-

cionales será cargado al autor.

INSTRUCTIONS TO AUTHORS

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,

Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, Spain.

+ Manuscripts may be written in any modern language.

* When a paper exceeds 20 pages, extra pages will be charged to the author(s) at full cost.

+ Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side on A-4

(297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together with a floppy

disk containing the article written with a DOS or Macintosh word processor. When a paper has joint authorship, one

author must accept responsability for all correspondence.

* The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate.

* Papers should conform the following layout:

First page. 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. These 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.

» 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

manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper comma (*).

* References in the text should be written in small letters or SMALL CAPITALS: Fretter and Graham (1962) or FRETTER

AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must include all of them

[Smith, Jones 82 Brown (1970)], thereafter use et al. [Smith et al. (1970)]. Ifan author has published more than one

paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit.

The references in the reference list should be in alphabetical order and include all the publications cited in the text

but only these. ALL the authors of a paper must be included. These should be written in small letters or SMALL CAP-

ITALS. The references need not be cited when the author and date are given only as authority for a taxonomic name.

Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of publisher, place of

publication, indication of edition if not the first and total number of pages. Keep references to doctoral theses or any

other 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. 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 Zoológica, 3 (5): 21-51.

* Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size

when designing figures. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high,

or be proportional to these sizes. Two columns format is recomended. It is desirable to print figures with their legend

below, so authors are asked to take this into account when preparing full page figures. If computer generated graph-

ics are to be included, they must be printed on high quality white paper with a laser printer. Photographs must be of

good contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are

of uniform width. Remember that grouping photographs of varied contrast results in poor reproduction. Take

account of necessary reduction in lettering drawings; final lettering must be at least 2 mm high. In composite draw-

ings, each figure should be given a capital letter; additional lettering should be in lower-case letters. A scale line is

recomended to indicate size, magnification ratio must be avoided as it may be changed during printing. UTM maps

are to be used if necessary. Figures must be submitted on separate sheets, and numbered with consecutive Arabic num-

bers (1, 2, 3,...), without separating “Plates' and “Figures”. Legends for Figures must be typed in numerical order on

a separate sheet, and an Spanish translation must be included. Follow this example (please note the punctuation):

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.

Authors wishing to publish illustrations in colour will be charged with additional costs (30,000 ptas, 180 euros per

page). They should be submitted in the same way that black and white prints.

If the authors want to send Figures in digital format, CONTACT the Editor first.

e Tables must be numbered with Roman numbers (I, II, TIL...) and each typed on a separate sheet. Headings should

be typed on a separate sheet, together with their English translation. Complex tables should be avoided. As a general

rule, keep the number and extension of illustrations and tables as reduced as possible.

e Manuscripts that do not conform to these instructions will be returned for correction before reviewing.

e Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the date

the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two referees. Based of

these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors will receive a copy of

the referees” comments. If a manuscript is accepted, the Editorial Board may indicate additional changes if desirable.

Acceptable manuscripts will be returned to the author for consideration of comments and criticism; a finalized man-

uscript must then be returned to the Editor, together with a floppy disk containing the article written with a DOS or

Macintosh word processor. Dates of reception and acceptance of the manuscript will appear in all published articles.

* Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special

attention to references and their dates in the text and the Bibliography section, and also to numbers of Figures and

Tables appearing in the text.

e Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs

are returned, and will be charged at cost. NO LATER orders will be accepted.

LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Junta Directiva desde el 14 de noviembre de 2000

Presidente Emilio Rolán Mosquera

Vicepresidente Diego Moreno Lampreave

Secretario Luis Murillo Guillén

Tesorero Jorge J. Otero Schmitt

Avda. de las Ciencias s/n, Campus Universitario, 15706 Santiago

de Compostela, España

Editor de Publicaciones Gonzalo Rodríguez Casero

Apartado 156, 33600, Mieres del Camino, Asturias, España

Bibliotecario Rafael Araujo Armero

Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez

Abascal 2, 28006 Madrid, España

Vocales Ramon M. Álvarez Halcon

Benjamín Gómez Moliner

Eugenia María Martínez Cueto-Felgueroso

Jesús Souza Troncoso

José Templado González

La Sociedad Española de Malacología se fundó el 21 de agosto de 1980. La sociedad se registró como una aso-

ciación sin ánimo de lucro en Madrid (Registro N* 4053) con unos estatutos que fueron aprobados el 12 de

diciembre de 1980. Esta sociedad se constituye con el fin de fomentar y difundir los estudios malacológicos

mediante reuniones y publicaciones. A esta sociedad puede pertenecer cualquier persona o institución interesada

en el estudio de los moluscos.

SEDE SOCIAL: Museo Nacional de Ciencias Naturales, c/ José Gutierrez Abascal 2, 28006 Madrid, España.

CUOTAS PARA 2004:

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

Socio Corporativo (en Europa): 48 euros

(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.

ÍNDICE

Iberus 22 (2) 2004

NAGEL, K-O. Observations on the reproductive period of the freshwater mussel Potomida littoralis

(Unionidae)

Observaciones sobre el periodo reproductor del mejillón de agua dulce Potomida littoralis

(Ontonidae) cc 1-8

MARTÍNEZ-ORTÍ, A., APARICIO, M2 T. Y ROBLES, E La malacofauna de la Sierra de Alcaraz (Alba-

cete, España)

The molluscan fauna of' the Alcaraz mountains (Albacete, Spaia) cono no 9-17

PÉREZ-QUINTERO, J. C., BECH TABERNER, M. Y HUERTAS DIONISIO, J. L. Los moluscos de las

aguas continentales de la provincia de Huelva (SO España)

Freshwater Molluscs of Huelva Province (SW Spaia) ooo noni 19-31

ÁvILa, C., FONTANA, A., ESPOSITO, M., CIAVATTA, M? L. AND CIMINO, G. Fatty acids of

Antarctic gastropods: distribution and comparison with Mediterranean species

Ácidos grasos en gasterópodos antárticos: distribución y comparación con especies mediterrá-

A A 33-44

DELLANGELO, B., ANSEEUW, B., TERRYN, Y. AND BONFITTO, A. Why are beached Acanthochi-

tona fascicularis (Linnaeus, 1767) (Mollusca: Polyplacophora) from Italy missing their

dorsal girdle elements?

¿Por qué los ejemplares de Acanthochitona fascicularis (Linnaeus, 1767) (Mollusca: Polyplaco- *

phora) encontrados en una playa de Italia pierden los elementos del perinoto? ........... 45-49

ROLÁN, E. The genus Calyptraea (Gastropoda, Caenogastropoda, Calyptraeidae) in the East Atlan-

tic

El género Calyptraea (Gastropoda, Caenogastropoda, Calyptraeidaejen el Atlántico orien-

A A A 51-79

MIA

ISSN 0212-3010