1123 mol/. 1DCYVYUS Vol. 22 (2) REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Oviedo, diciembre 2004 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 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 9 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 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 Orontes Orontes Syria Orontes Orontes Syria 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 23.6.2000 30.6.1991 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 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 Nagel and Badino, 2001 Nagel and Badino, 2001 MNCN FW-1157 MNCN FW-645 Haas, 1917, 1920; SMF 003054; this study Nesemann and Nagel, 1989 Nesemann and Nagel, 1989 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 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. 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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. 9 Iberus, 22 (2), 2004 Río Alcaraz 8 QRO Reolid Río Salobre de ich ESPAÑA 12 En Ñ / R Ario 6 26 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 10 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. 11 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 12 012 01 50/0 5/0 3/0 (Mv 10 0 0/4 2160 08 Mm 34 1/0 41 0/6 1/0 0/2 2/4 0/2 114 01 01 0/5 01 0/1. 0/2 0/5 0/2 01 0/3 8/0 2/0 0/5 41 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 0 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 0/2 0/3 02010307 01 01 01 1/3 3/8 111 011 0/8 01 1/0 0/2 MM 0/15 014 01 0 1/0 112 0102 4/0 1/0 1/0 5/0 2/6 01 2/0 0/5 1/0 4/0 01 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 13 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. 14 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, IS 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. 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Jor- nadas sobre el Medio Natural Albacetense, Ins- tituto Estudios Albacetenses, 339-357. 17 oe sosdlA) Ba: É SE > GON 4%, sl 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 19 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 20 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 21 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. 22 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. He (1111000 7 j o QB al EAN 0 ll e el al s Ol / PB Ú LS — QAN |] Radix balthica Ce 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. 23 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 24 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) UA a o sE Y | ++ AL HA y + 5 ES AA Ha A. Aaa: AH E El | — dale Los + S QA 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 | | EE Uno pictorum Anodonta cygnea PC / Ele QC PC Mar “as LL . TH (e e 7 ADOS / LB ATA] HL 0B aL ++ arg e 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- 25 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. 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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. 28 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 ZO 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 2,0Pt 2,0Gt, Pt, Bc, Pc 1,9Pt, Fc 1,9Pt 1,6Pt 1,5Pt 1,5Pt Pt 0,7Pt 29 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 30 UTM 295081983 295081879 295080664 2950B1372 2950B2175 295080652 295081368 295080061 2950B1866 295081057 2950B0127 295081851 295082450 295081851 2950B1663 295PB9444 295082045 295PB9244 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 700 640 600 580 580 580 580 560 540 500 460 440 440 440 380 360 360 330 330 320 320 CUENCA DEL GUADIANA-PIEDRAS (23/19/9) H= 3,75 A 600 600 580 580 540 540 480 480 Iberus, 22 (2), 2004 CUENCA DEL TINTO (12/12/6) H= 2,19 ESPECIES Af Pm, Ga, Af Pt 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 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 Af 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 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 Af 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 Pt, Af Pt Pt, Pm Rb, Pt, Pm, Pc, MI Pt Pt Pt, Af Pt, Af Pt, Up” Af Pt Up" Af, Up" Pm, Ga Pt 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, Pm, Af Pt, Af Pt, Af Pt, Pm, Af Pt, Af Pt, Af Pt, Af, Up" Pt, Pm, Af Pt, Pm, Af ESPECIES Pt, Af Pt, Fc, Af, Up" Pt, Af 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 400 400 380 360 360 350 320 310 285 280 280 215 165 160 160 160 150 150 150 140 120 95 95 90 120 110 90 80 Pt, Af Af, Up”, Ps, Pp Af, Ps Pt, Af 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 Af 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 Af Pt, Af, Up”, Ac” Pm, Af Me, Af Pu Me, Gt, Af, Pl, Up, Ac, Cf Mm Ra, Rb, Pt, Pm Pt, Fc, Af, Up, Ac, Cf Pt Ra Mm - Rb, Cf Pt, Bc Me, Af 31 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. 33 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- 34 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 35 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. SL 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 36 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 SY 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 38 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). SL 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- PL 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 39 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 40 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 a Ll 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. A] 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. 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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: 44 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 A5 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 46 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). 47 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 48 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. 49 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 P » QA 15 4, SE 2147 Í il carl Ma ; DonvHs e ¡0 *ke ved Joy se still A hd tigoabonk bid l LEVES £ y K el “E Ye vé a LN 5 tr Ló dro y UA |] e a 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 Si 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 53 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. 54 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). ES 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). SY 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. 58 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 39 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. 60 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. 61 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 62 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). 63 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. 64 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 65 Iberus, 22 (2), 2004 De be dd da Me 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). 66 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. 67 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. 68 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 no 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 no 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 no no 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 no 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 no Spiral grooves unknown unknown transparent with very evident radiating milk white lines at the border elongate, straight, with two angulations 25-93 m 69 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 7O 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 SN 92 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. EN 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. 72 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 73 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), 74 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 75 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. 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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. IT 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. 78 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. TA de Es > ñ vismshA AH sd ns aa E ] =— o ira O 7 mood 0 y > ar 15 lar y pan Dep A e ¿(de He LARES e EN 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. 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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. 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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